ThePrinciplesofBiology 10045931

T HE P R IN C I P L E S OF
B IO L O G Y
H E RBE RT S P E NC E R
IN T W O VOL UM E S
VO L . ll
RE VIS E D A ND E NL A RGE D E DITIO N
1 899
NE W YO RK
D . A PPL E TON AND COMPA NY
1 900
8 8 8 03
P R E FA C E
T O T HE R E VI S E D A ND E N L A R GE D E D I T I ON
OF VOL . II .
T o t he s tate m ent s m a d e i n the p r e f a c e t o th e fi r s t v ol u m e

of thi s re vis e d e di ti on th er e mu st h er e be a dd e d
,
a fe w h a v
i ng p e ci a l f ce t o thi s s e c on d v olu m e
‘
s r e e r en .
O ne of t h e m i s th at the re v isi on has n ot b e e n c a rr i e d

ou t in q u i te th e s am e w a y , bu t i n a w ay s om e wh a t l e s s c om
l
p e te . W h en re vi e wi n g th e fi r s t v olu m e a f ri en d l y c r i tic ,
P r of L l oy d. M or g an , s ai d
B ut th o ug h th e i nt el l e ctu a l w e i g h t h as a l so b ee n a u g m e nt e d , it
“
is a n o p en q u e s ti on w h eth er i t w ou l d n ot h a v e b ee n w i s e r t o l e a v e
i nt a ct a t re at i s e , & c rel e at i n c orre c ti ons a nd a ddi t i o ns t o
.
g g
n ot es a nd a pp e n d i c e s .
I thi n k t h at P r o f . M or g a n is righ t . T h ou gh at th e cl o s e
of th e p r e fa c e to v ol um e I , I w r ote in al l s ec ti on s n ot
m ark e d as n ew, th e e ss enti a l id ea s se t f or th a re th e sam e as
th e y w e re i n th e igina l e di ti on of
or y et th e d er
r ea
who h a s n ot r e a d t hi s st a t e m e nt or d oe s
,
n ot b e ar i t i n mi n d ,
will s u p pos e th a t al l m ost of the e nun ci a t e d c on c e p ti on s
or
a r e of r e e nt c d a te ,
wh er ea s onl y a sm a ll p ar t of th e m a r e .
I f ol l ow , i n thi s s e c on d v ol um e, a
h a v e th e re f or e d e cid e d to
c our s e s om e wh a t li k e t h at s u g g est e d by P r of Mor g an .
s om e wh at lik e , I say , b e c a u s e i n s u n d r y c as es th e am e n d
m e n ts c ou ld n ot be s a tisf a c tor ily m ad e by a pp e nd e d note s .
Vi PR E FAC E TO TH E RE V I SE D E D I T I ON .
B u t th e r e h a s b ee n a furth e r r e a s on for th i s ch a n g e of
m e th o d . An in v alid w h o is ne ar ly e ig h ty c ann ot wi t h p ru
d enc e e n te r u p on w or k which wi ll ta k e l on g to c omp l ete .
H e n c e I h a v e t h ou gh t i t b e tter to m a k e th e n ee df ul a lte ra
ti on s an d a ddi ti ons in w a ys re q u i r i ng r e at l i v el y m od e rate
tim e and l a b our .
Th e a ddi ti on s m a d e t o t his v ol u m e a re l e ss nu m er ou s and
l e ss imp orta nt th an t h os e m a d e t o th e fi rs t v ol u m e . A n ew
ch a p ter en di n g P a rt V, on
“
Th e I nteg ra ti on of th e Or
g ani e W or ld , s er v e s t o r oun d off th e g ene ra l t h eory of

”
E v ol uti on i n i ts a p p li cat i on t o li vi ng t hi ngs B e y on d a n e w .
s e c ti on 28 9 a ) a nd th e v ari ou s f o ot -
n ote s , s e r vi ng c hi efly
th e p u rp os e of e l u cid at i on, th er e a re no e t s of s om e si
g ni fi
c an c e a pp en d e d to C h a p t er s I , I I I , I V, an d V, i n P a rt I V ,
C h a p te r s V an d V I I I , i n P art V, a nd C h a p te r s I X , X , an d
X I I i n P a rt V I .Mor eo v e r t h ere a r e t h re e f u rth e r a pp en

dic e s , D F, and G which h a v e, I t h in k , c on sid e r a bl e si g
’
, ,
n i fi c a nc e as s e r vi n t o m a k e cl ear e r s om e of th e vi e ws
g
e xp res s e d in th e b ody of th e w or k .
T urn i ng f r om th e ad di ti on s to the re vi si on s , I h a v e to
say t h a t the ai d n e e d e d for b ri ngi ng up to d a te th e c onte nts

of this v ol u m e , h as b een gi v en m e b y the g entl e m en w h o
ga v e m e l ik e ai d in r e vis i ng th e fi r st v ol um e : omi tting
P rof P er ki n, wi thi n wh os e p ro vi nc e n on e o f th e c onte nts
.
o f th is v ol u m e f a ll P l an t Morph ol o gy and P l ant P hysi

.
- -
l gy h a v e b een o v e r s een by Mr A G T an sl e y Oriti

o o . . . .
cism e u p on p art s d e ali n g wi th A nim al Mor ph ol og y I owe

t o Mr J T C u nni n gh a m an d P rof E W Mac Bri de
. . . . . .
A n d the s ta te m ents incl u d e d u nd e r A n im al P hysi ol ogy

h a v e b een c h e ck e d by Mr W B H a rdy . . . .
PR E F AC E TO T H E R E VI SE D E D I T I ON .
vi i
F or s
r e a on s l ik e t h os e na m ed i n t he p re fa c e t o th e fi rs t
v ol u m e, I h a v e n ot su bmi tte d th e p roof s of t his re v i se d
s e c ond v ol u m e t o t h e s e g entl e m en : a fa c t which i t i s nee d
ful to m e , si n c e on e or oth er of t h e m m igh t el s e be h el d
na
r e sp on sibl e for s om e e rr or which i s n ot hi s b u t mi n e I t is .
th e m ore re q uisi t e to sa
y t his b e c au s e whil e, i n re s p e ct of
m atte r s of f a c t, I h a v e, sa v e i n on e or tw o cas e s , a c c e p t e d
t h e ir c orre c ti on s a s n ot to be qu e s ti on e d, I h a v e n ot a l w ay s
d one t his i n re sp e c t of m a tte r s of i n f e ren c e , but in s u ndr y

pl a c e s h a v e adh er e d t o my o wn i nter p re ta ti ons .
P e rh a p s I m ay be e xc u s e d for e xp re ssi ng s om e sa t i sfa c
t i on t h a t I h a ve n ot b een obli g e d t o re li n q u i s h th e v i e ws set

f ort h i n 1 8 6 4— 7 . T h e hyp oth es i s of physi ol og ic a l u ni ts — or,
as I w ou ld n ow c a ll t h e m , c on sti t ut i ona l u ni ts — ha s b een
a d o p te d by s e v e ra l l g is ts un d er m odi fi e d f or ms S o fa r
z oo o .
as I am a w a re , the a ll e g e d g en e ra l l a w of or a nic symm e try

g
h as not ca ll e d f o rt h an
y m anife sta ti on s of di ss en t . The
s ugg e s te d th e or y of v erte b ra te s truc tu r e a pp e ar s to h a ve
b e c o m e c u rrent ; an d fr om the i n ve s tigati on s of the l ate
P rof C OP e , ha s
. re c e i v e d v e rific ati on T he c on cl u s i on s
.
d ra w n i n P ar t VI on
“
Mul tiplic ati on h a v e
The L a ws of ,
”
not I b e li e v e b een c ontr o v ert e d

, A nd t h ou gh only s o m e
, .
w ork s on b ota n y h a v e gi v en c u rre n c y to th e d o c tri n e s et

f ort h i n A p p endi x C “
O n C i r c u l a t i on a n d th e For m a ti on
of W oo d i n P l a nt s
ye t I h a v e m e t w i th n o a tt e m pt t o dis
,
p ro v e i t The only v i e ws c ont e ste d by c ert ai n of th e g e n

.
tl e m en a b o v e na m e d a re th o se c on c e rn i n g th e or i i n of
, g
th e t w o g re a t ph aen o g a mic t p e s of pl a nts a n d th e ori gi n
y ,
of th e a nnu l os e t yp e of a n im a l s I h a v e n ot h ow e v e r
.
, ,
pe rh a ps b e c a u s e of na tu r a l bi a s— f ou n d my se lf c om p e ll e d
PR E F AC E TO T H E R E V I SE D E D I T I ON .
to s urr en d er t h e s e v i e w s . My s
re a on s for a dh eri ng t o th e m
will be f ou n d i n n ote s t o th e en ds of C h a p te r s II I a nd IV
i n P a rt I V,
2
and i n A pp en di x D
On n ow fin a lly l ea vi ng bi ol ogic a l s tu di e s i t re m a i ns only
to s ay th at I am gla d I h a v e s ur vi v e d l ong e nou
g h to giv e
this w or k i ts fin ish e d f or m .
B R I G H T ON
Oc tobe r , 1 8 9 9 .
P RE FA C E T O VOL . II .
T HE p r oof sh eets of this v ol u m e , l ik e th os e of th e l a s t

v ol um e, h a v e b ee n l ook e d th rou gh b y D r H ook er and .
P rof H u xl e y ; a n d I h a v e , a s b e for e , to t h a n k th e m for

.
t h e i r v a l u a bl e c ri ticisms , a n d for th e t rou b l e t h e y h a v e

ta k e n i n ch e cki n g t he n u m er ou s s ta te m en t s of f a ct on
which the a rg um ent s p ro c ee d T h e c on sci ou s ne s s th a t
.
th ei r m a n y du ti e s r en d e r ti m e e x t re m e ly p r e ci ou s t o th e m ,
m a k e s m e f ee l how h ea v y is m y o bliga ti on .
P a rt I V , wit h which this v olu m e c om m e n c e s , c onta i n s

.
nu m erou s fi ur e s
g . N e a r ly on e h a lf of th e m are re p e ti
ti on s , m ostly a l t e re d i n sc al e a nd s implifi e d i n e x e cu ti on, of

fig u res or p ar ts of fig ure s , c on ta i ne d i n the w or k s of v ari
,
o us B otan is t s a n d Z ool ogis t s A m ong th e a uth ors wh om I

.
h a v e l aid u n d er c ontri bu ti on, I m ay n a m e B er k e l e y , C ar

p e n t e r
,
C u vi er ,
G r ee n ,
H a r v e y ,
H oo k e r ,
H u xl e y ,
M il n e
E d w a rds , R a lfs , S mi th T he r e m a i ni ng fi gu re s , n u mb eri ng

.
1 5 0, a re f r om or igi na l sk et ch e s a n d di a g ra m s .
T h e s u cc e ssi v e i n st a lm e nts which c o mp os e this v ol u m e,

w e re is su e d to the S u b s c r ib e rs a t th e f oll ow in g d ate s — N o .
1 3 ( pp 1 — 8 0) i n J anu ar y , 1 8 6 5 ; N o 1 4 ( pp 8 1 — 1 6 0) i n
. . .
J u n e , 1 8 65 N o 1 5 ( p p 1 6 1 — 2 4 0) i n D e c e m b er , 1 8 6 5 N o
. . .
1 6 ( pp 2 4 1 — 3 2 0) i n J u n e , 1 8 6 6 ; N o 1 7 ( pp 3 2 1 — 4 00) i n
. . .
N o v e mb er, 1 8 6 6 an d N o 1 8 ( p p 4 01 — 5 6 6 ) i n Mar ch , 1 8 6 7
. . .
L O ND O N Ma r c h
, 23 r d , 1 8 67 .
C ONTE NTS OF VO L . II .
P A R T I V — MOR P H OL OGI C A L
. D E VE L OP ME N T .
CR AP .
I — T HE
. O F MORP H OL OGY
P ROB L E MS
II — T H E M O R P H O L O G I CA L C O M PO S ITIO N O F P L A N T S
.
II I — T H E M O R P H O L O G I CA L C O M PO S ITIO N O F P L A N T S— C onti n u ed
.
I V — T H E M O R P H O L O G I CA L C O M PO S ITIO N O F A N I MA LS
.
V — T HE M O R P H O L O G I CA L C O M PO S ITIO N O F A N I MA LS— Conti nu ed

.
VI — M O R P H O L OG I CA L DI FF E R EN TI A TIO N I N P L A N T S
.
VII — T H E G ENE RA L S HA P ES O F P L A N T S
.
V III —T H E S HA P ES O F B RA N CH ES
.
I X — T H E S HA P ES O F LE AV ES
. .
X — T H E S HA P ES O F F L O WE R S
.
XI — T H E S HA P ES O F V E G E T A L C ELLS
.
X II — C HA N G ES O F S HA P E OT H E R W I SE CA USE D
.
X III — M O R P H O L OG I CA L DI FF E R EN TIA TIO N I N A N I MA LS

.
XI V — T H E G E N E RA L S HA P ES O F A N I MA LS
.
XV — T H E S HA P ES O F V E R T EB RA T E S K ELE TO NS
.
X VI . T H E S HA P ES O F A N I MA L C ELLS
- -
E V I L— S U M MAR Y O F M O R P H O L O G I CA L D E V EL OP M EN T .
P A RT V — P H Y S I OL O GI C A L
. D E VE L OP ME N T .
I —T H E
. P R OBLE M S O F P H YS IO L O G Y
I I — D I FF E R EN T I A TIO NS
. BE T WEEN T H E O U T E R A ND I NNE R TI S
SUES O F P L AN T S
I I I — D I FF E R EN TI A TIO NS
. AM O N G T H E O U T E R TI SSUES O F P L A N T S
I V — D I FF E R EN TI A TIO NS
. AM O N G T H E I NNE R TI SS UES O F P L A N T S
xi
Xfi C ONTENT S
C RA P .
P H YS IO L O G I CA L I N T E GRA TIO N I N P L A N T S
V— .
VI —D I FF E R EN T I A TIO NS BE T WEEN T H E O U T E R A N D I NNE R TI SSUES

.
O F A N I MA LS
VI I — D I FF E R ENTI A TIO NS AM O N G T H E O U T E R TI SSUES
. A N I MA LS
VI II — D I FF E R EN TI A TIO NS AM O N G T H E I NNE R TI SSUES
. A N I MA LS
I X — P H YS IO L O G I CA L I N T E GRA TIO N I N A N I MA LS
.
X — S U M MAR Y O F P H YS IO L O G I CA L D E V EL OP M EN T
.
X — T H E I N T E GRA TIO N O F T H E O RGA N I C W O R L D

‘
.
P A R T VI — L A WS OF M U L T I P L I CA T I ON J
FAC TO R S
I — THE
.
I I — A P RI OR ! P R I N C IP LE
.
I II —O B V E R SE A P RI OR ] P R I N C IP LE
.
I V — D I FF I C UL TI ES O F I N D U C TI V E V E R I F I CA TIO N
.
V — AN T AG O N I S M BE T WEEN GR O W T H A N D A SE X U A L G ENES I S
.
VI — AN T AG O N I S M BE T WEEN GR O W T H A N D S E x U A L G ENES I S
. .
VI L — T H E A N T AG O N I S M BE T WEEN D E V EL OP M EN T A N D G ENES I S ,
A s E xU A L A N D S E x U A L
V II I — AN T AG O N I S M BE T WEEN E X P EN DIT U R E A N D G ENES I S

.
I X — C OI N C ID EN C E BE TWEEN H I GH NU T R ITIO N A N D G ENES I S

.
X — S P E C I A L ITI ES O F T H ESE R EL A TIO NS

.
X I — I N T E R P R E T A TIO N AN D ! U A L I F I CA TIO N
.
V
X I I — MUL TIP L I CA TIO N O F T H E H U MA N RAC E
.
t/ XII I — HU MA N POP UL A TIO N I N T H E F U T U R E

.
A P P E ND I CE S .
A —
.
S UBS TIT U TIO N O F A X I A L F O R F O L I AR O RGA NS I N P L A N T S
B — A CR ITI C I S M O N P R O F O WEN S T H E O R Y O F T H E V E R T EB RA T E
’
.
.
S KELE TO N
C — ON C I RC ULA TIO N A N D T H E F O RMA TIO N O F W OOD I N P L A N T S
.
.
D — ON T H E O R I G I N O F T H E V E R T EB RA T E T Y P E
.
.
D 9 — T H E A NNUL O SE T Y P E
.
E — T H E S HA P ES A N D ARRA N G E M EN T S O F F L O WE R S
.
F — P H YS IO L O G I CA L ( O R C O NS TIT U TIO N A L) UN IT S
.
G —T H E I N H E R IT A N C E O F F UN C TIO N A LLY CA USE D M ODI FI CA TIO NS

-
.
CH A P TE R I .
T H E P RO B L E M S OF M O RP H OLO GY .
§ 1 75 . T H E di visi on of Mor phol ogy f rom P hysi ol ogy i s

,
one whi c h m ay be t ol er a bly w e ll p r e s er v e d s o l on g as w e do

-
not c a rr y ou r i n qu i r i e s b ey on d th e e mpi r ic a l g enera li z a ti on s
of th e i r res p ectiv e ph enom e na ; bu t it is one w hich b ec om e s
i n grea t m ea su r e nom i na l wh en th e ph enom en a are to be

,
ra ti on a lly i n t er p ret e d I t wou ld be p ossibl e a ft er a na lyzi n g

.
,
ou r S ol ar S y s t e m to set d own c ert a i n g enera l t ru th s re sp ect

,
in g th e sizes a n d di st a nc es of its p ri m ary a nd s ec ond ar y

m emb er s om itti ng a l l m enti on of th ei r m oti ons ; and it wou ld
,
be p oss ibl e to set d own c ert a i n oth er g ener a l t ru th s resp e ct

i ng th ei r mo ti ons W ith ou t sp ec ifyi n g th ei r di m en s i ons or
,
p os iti ons fu rth er th an as gr eat er or l ess ne arer or m ore r e

, ,
m ot e Bu t on seeki ng to a ccou nt for th es e genera l t ru th s

.
,
a rr iv e d a t by i n d u cti on w e fi nd ou r se lv e s oblige d t o c on sid er

,
s im u lt a neou sly th e r e l a tiv e s i z e s a n d pl aces of th e m a ss e s ,
a nd th e re l a t iv e a m ou nts a nd di recti on s of th e i r m oti on s .
S imil arly with orga n i s m s T h ou gh w e m ay f ra m e s u n d ry

.
c om p reh ens ive p rop os iti ons r esp ecti n g the arrangem ent s of
th ei r organs c on s id ere d a s so m an y i nert p a rt s ; a nd th ou gh
,
We m a y est a bli s h s ev era l w id e c on cl u s i on s r esp ecti ng th e s ep a

r a t e a n d c o mbi ne d a c ti on s of th ei r or g an s W ith ou t kn ow i n g
,
a nyt hi n g d efi n it e res p ecti n g th e f orm s a nd p os iti on s of th es e
or g ans ; e t w e ca nn o t r e ac h s u c h a r a ti on a l e of th e fa c ts a s
y
3
4. M O RPH OLO G I CA L DE VEL O PME NT .
th e hyp oth esi s of E

v ol u ti on a ims at with ou t c ont empl ati ng
,
s tru ctu r es a nd f u n c ti on s i n th ei r m u tu a l r el a ti ons E v ery .
w h er e st ru ctu res i n g r ea t m ea su re d et erm i n e fu n cti ons ; a nd

e v e r y wh e r e fu n cti on s a re i n c es s a ntly m o di fyin g s t ru ct u r es .
I n N atu r e the two ar e i ns ep a r a bl e co—Op er a tor s an d S ci enc e

c a n giv e n o t ru e i n t er p ret a t i on Of N a tu r e wi th ou t k e e pi n g
the i r co—op er a ti on c ons t a ntly i n Vi ew A n a ccou nt of org anic

.
e v ol u ti on i n its m or e s p e ci a l a s p ect s m u s t be e ssenti a lly a n

, ,
a c cou n t of th e i nter a c t i on s of s t ru ct u r es a n d f u nc ti on s a s
-
,
p erp etu ally a lt ere d by c h anges of c on diti on s .
H enc e wh en t re a ti n g a p a rt Mor ph ol ogic a l D ev e l opm ent

,
a nd P hysi ol ogic a l D e v e l op m ent a l l w e c a n do i s to di rect ou r

,
a tt enti on m a in ly to th e one or t o th e oth er a s the c a s e m ay ,
be I n d ea li ng with th e f ac ts of st ru c tu re w e m u s t c onsid er
.
,
th e f acts of fun cti on only i n su ch g enera l wa y as i s n eed fu l

to expl a in th e f a c ts of st ru c t u r e ; an d conv ers ely wh en d e a l
i ng with the f a cts of fu ncti on .
1 76 . p r obl em s of M orp h ol ogy f a ll i nt o two disti nct

T he
cl a s ses an sweri ng res p ectiv ely to th e two l ea di ng a sp ects of
,
E v ol u ti on I n thi n gs whi ch ev olv e th e re g o on two p roc ess es

.
— i n c re a se of m a ss a n d i nc rea s e of s t ru ctu re I nc rease of .
m a ss i s p r im ar y a nd i n s i m pl e ev olu ti on tak e s p l a c e a lm os t
,
a l one . I nc rea s e of s t ru ctu re is s econd ary a ccomp anyi n g or ,
foll owi ng i nc rease of m ass w ith m ore or l ess regu l ar ity Wher ,
e v er e v ol u ti on r i s e s a b ov e th a t f or m whi ch s m a ll i norg a n ic
b odi es su c h as c ryst al s p resent to u s A s the f und a m ent a l

, ,
.
a n t a g on i s m b etween D iss ol u ti on an d E v ol u ti on c ons i s t s i n thi s ,
th a t whil e th e one is a n i nt eg ra ti on of m oti on a nd di s

i ntegr ati on of m a tt er the oth er i s a n i nt egr ati on of m att er
,
a n d di s i n t eg r a ti on of m oti on ; a n d a s thi s i nt eg r a ti on of m a t
t er a cc omp anyi n g di s i nt egr ati on of m oti on i s a necess a ry ,
a nt eced ent t o th e different i a ti on of th e m a tt er s o i nt e g ra t e d ;
it foll ows th a t qu esti ons c oncerni ng the m od e i n w hich the

p arts a re u nit ed in t o a W hol e mu st be d ea lt with be fore
,
T HE PR O BLE M S OF M O RPH OLO G Y .
5
questi on s c onc erni ng th e m od e in which th es e p a rts b ecom e

*
m odi fi ed .
T hi s is bvi ou sly a m orph ol ogi ca l qu esti on Bu t an

not o .
illu stra ti on or two wi ll m a k e it m a ni fes t th a t f u nd am ent a l

di fferenc es m ay be p rod u ce d b et ween a ggr ega tes by di ffe rences
i n th e d e gr ees of c om p os i t i on of th e i n c r em en t s : th e u l t i m a t e
u n its of th e i nc r e m en ts b e i n g th e s a m e T h u s a n a ccu m u l a .
ti on of thi ngs of a giv en ki nd m ay be m a d e by a ddi ng one at

a tim e . Or the thi ngs m ay be ti e d u p i nto b u n dl es of ten ,
and the t en s pl a c e d t og eth er Or th e t ens m ay be u nit e d .
i nto hu n d re d s a n d a pile of hu nd r eds f or m e d S u ch u nli ke

,
.
n es ses i n the st ru ctu r es of m asses a re h a bitu a lly s een i n ou r

m ercantil e t ran s a cti ons A rt i cl es which th e consum er r e.
c ogn ize s a s si ngl e th e ret a il er k ee p s wra pp e d u p i n d ozens

, ,
th e Wh ol es a l er s en d s i n gr oss a nd th e m anu f a ct u rer su ppli es ,
i n p a ck a ges of a hu n d re d g ros s T h a t is th ey se v er a lly i n .

,
c r e a s e th e i r s t oc k s by u n i t s of S i m pl e of c om p ou n d a nd of , ,
d ou bly c om p ou nd ki n d s S i m il a rly resu lt those di ffe ren c es of

-
.
m orph ol ogic a l com p os iti on whi ch we h a v e fi r st to con s id er .
A norg a nism consi st s of u ni ts T h ese u n it s m ay be a ggre .
g at ed i nt o a m a s s by the a ddi ti on of un it to u nit Or they .
m a y be un it ed i nt o grou p s a n d th e gr ou ps j oi ne d t og eth er , .
Or th ese grou ps of grou p s m a y be s o combi ne d a s to for m a

d ou bly—c om p oun d a ggr eg a t e H en c e th ere a r i ses re sp ecti ng
.
ea ch or g a ni c f or m th e qu es ti on— is i t s c omp os iti on of th e
fi rs t s ec on d thi r d or fou rth or d er — d oes it e xhibit u nit s of

, , ,
a si n gly c omp ou n d e d ki n d on ly
-
or a re th es e c ons olid a t e d ,
i nt o u ni t s of a d ou bly comp ou n d ed ki nd or a t riply com -

,
-
p ou n d ed ki nd ? A nd if it di s pl a ys d ou bl e or t ripl e c omp os i
It s ee m s nee dfu l h ere to s a y, th a t a ll u s i on is i n th i s p a r a gr a ph to a
m a de
p ropo s i t i on res pe cti ng th e u l ti m a t e n a tu re s v

of E ol uti on a nd D i s s ol ut i on,
whi ch i s cont a i ned i n a n e ssa y on The C l ass ific a ti on f

o the Sci ences , p u b
l i sh e d i n Ma rch , 1 8 64 . When t he opp ortuni t y co m e s , I h Op e t o m a k e th e
de finiti on th e re a rri ved a t the b as i s of a re o rga ni za t i on of th e

,
- s econd
p a rt of
F i r st P r i nc ip l es : gi v i ng t o th a t work a h i gh er de v el opm ent ,

a nd a
gre at e r
coh es i on, th an i t at p re sent p osse sse s .
[ The i ntenti on h ere i ndi ca te d was
d ul y c a rri ed o ut in
6 M O RPH OLO G I CA L D E V ELO PM ENT .
ti on the h om ol ogi es of its di fferent p arts b ec om e p robl ems

,
.
Und er the di s gu i ses i n d u ced by th e c on solid a ti on of p r i m a ry ,
s e c ond ar y a n d t er ti a ry un it s it h a s to be a scert a i ne d which

, ,
a n s wer t o whi ch i n th ei r d eg r ee s of com p os iti on

,
.
S u ch qu e st i ons a re m ore i nt ri c at e th a n t h ey a t fi rst a p

p ea r ; si nce b es id es th e ob scu r i ti es c au sed by p rog ress iv e i nte
,
a t i on a n d th os e du e to acc om p a n yi n g m odifi ca ti ons of f or m

g r , ,
fu rth er ob scu r iti es r esu lt f rom th e v a ri a bl e g rowths of u n its

of th e di ffer ent or d er s Ju st a s an ar my m ay be au g m ent e d
.
by recru iti n g ea ch c omp any w ith ou t i nc reas i ng the nu mb er

,
of c omp a n i es ; or m ay be a u gm ent e d by m a ki n g u p th e f u ll
c ompl em ent of c omp ani es i n ea ch regi m ent whil e the nu m ,
ber of regi m ent s r em a i n s th e s a m e ; or m a y be a u gm en t e d

by p u tti ng m ore regim ents i nt o ea ch divi s i on oth er thi ngs ,
b ei ng u nch anged ; or m ay be au g m ent e d by a ddi ng to th e

nu mb er of its divisi on s with ou t a lt e r i n g th e c om p onents of
ea c h divi s i on ; or m ay be au g m en t e d by two or th ree of th es e
p r oc esses at once ; so i n org a ni s m s i n c r ea s e of m a ss m ay

, ,
res u lt f rom a dditi ons of u n its of th e fi rs t or d er or th os e of th e ,
s ec ond ord er or th ose of still hi gh er or d er s ; or it m ay be du e

,
to s im u lt a neou s a dditi on s to u n its of s ev era l or d ers A nd .
thi s l as t m od e of i nt egr ati on b ei ng the genera l m od e p u ts ,
di ffi cu l ti es i n th e way of ana ly si s Ju st as the s t ru ct u re of

.
a n a rmy w ou ld be m a d e l ess ea s y to u n d er s t a n d i f com p a n i es
o ft en ou t grew re gi m e nt s or r egim ent s b e c a m e l a rge r th a n

,
b rig a d es ; s o th ese q u es ti on s of m orphol ogi ca l com p os iti on

a r e c omplic a t e d by th e i n d et er mi n a t e si z e s of th e u nits of
eac h ki n d : rel a tiv ely s impl e u n it s f requently b ec omi ng m ore

-
b u lky th an rel a tively c omp oun d u nits

-
.
§ 1 77 T h e m orph ol ogic a l p robl ems of the s ec ond cl ass

.
a r e th os e h a vi n g for th e i r s u bj ect m a tt er th e ch a n ge s of s h a p e
-
whi ch a c com p a ny ch ange s of a gg reg a ti on T he m ost genera l .
qu es ti ons res p ecti ng th e s t ru ctu re of a n org a n i s m h a vi ng b een ,
a ns were d wh en it i s a s c ert a i ne d of Wh a t u n it s it is c om p os e d
as a wh ol e a n d i n i ts s ev e r a l p a rts ; th ere c om e th e m or e
,
T HE PR O BL E M S OF M O RPH OLO G Y .
7
sp eci a l qu esti on s c onc ern i n g i ts for m— form i n the or di nary

s en s e . A ft er th e c ontra st s c au s ed by v a r i a ti ons i n th e p roc ess
o f i nt eg ra t i on we h a v e to con s id er th e c ontr a s ts cau se d by
,
v ar i ati ons i n the p rocess of di ff erenti ati on T o sp ea k s p eci .
fi ca l l y— the Sh a p e of th e or g an i s m as a wh ol e i rre s p ec tiv e ,
of i ts c om p os iti on h a s t o be a cc ou nt ed for
, Rea s ons h a v e .
to be fou n d for th e u n lik enes s b et ween i ts gen era l ou tli nes

a n d th e g enera l ou tlin e s of a lli e d or g a ni s m s A n d th ere .
h a ve to be a nswere d ki nd red i nqu i r i es r es p ecti n g th e p rop or

ti ons of i ts c omp on ent p a r t s — Why a m on g s u ch of th ese as
,
a r e h om ol og ou s with one a n oth er h av e th er e a r i sen the

,
di fi er ences th a t exi st ? A n d h ow h a v e th ere b ee n p rod u c e d

th e con t ras ts b et w een th e m a nd th e h om ol og ou s p a r ts of
org a n i s m s of th e s a m e typ e ?
Ve ry nu m erou s a re th e h et erogene iti es of f or m p re s enti n g

th em se lv es for i nter p ret a ti on u n d er th e se h e a d s T he u lti .
m a t e m orph ol ogic a l un it s combi ned i n a ny grou p m ay be dif ,
ferenti a ted i n di vid u a lly or c oll e ctiv ely or b oth : e ach of th em

, ,
m a y u nd er g o c h a n g es of sh a p e ; or s om e of t h em m ay be
ch ange d and oth er s not ; or the g rou p m ay be rend ered m u l

tiform by the gr ea t er g rowth of s om e of i ts u nits th a n of
oth er s S imil ar ly wi th th e c om p ou n d u nit s a r i s i ng by u n i on
.
O f th es e simpl e u n it s A gg reg a t es of th e s econ d or d er m a y

.
be m a d e r el ativ ely c ompl e x i n form by i ne qu a li ti es i n th e

,
ra t e s of m u ltiplic a ti on of th e i r c omp on ent u ni ts i n div ers e
di re cti ons ; and a mon g a nu m b er of s u ch a ggr eg a tes nu m er ,
ou s u n lik en esse s m a y be cons t it u t e d by difi e r e n c e s i n th e i r
d eg rees of grow th and by di fferenc es i n th ei r m od es of grow th

,
.
Ma ni festly a t ea ch high er st a ge of com p os iti on th e p os sibl e

,
s ou rces of div er g en ce are m u ltipli e d s till f u r th er .
T h a t facts of thi s or d er ca n be a cc ou nt ed for i n d et a il is

not to be e xp ect e d— th e d a ta a r e w an t i ng A l l t h at w e m a y
.
h op e to do i s to a s c ert a i n th ei r genera l l aw s H ow thi s is to .
be a tt empt e d we will now c ons id er .
§ 1 78 . T he ta s k b e fore us i s to t ra ce th rou gh ou t th es e
8 M O RPH OLO G I CA L DE V ELO P MENT .
ph enom en a the p roc es s of e v ol u ti on ; and to Sh ow how as ,
di s pla y e d in th em i t con form s to th ose fi rs t p r i ncipl es whi c h

,
e v ol u t i on i n g en e r a l c on form s t o T wo set s of f actors h a v e

.
-
to be ta k en i nto a ccou n t L et u s l ook at th em

. .
Th e fa ct or s of th e fi rst c l a ss a r e th ose whi c h t en d di r ectly

t o c h a n ge a n or ga n i c a ggr ega t e i n c omm on w ith e v er y oth er
,
a gg r e g a t e f r om t h a t m or e S i m pl e for m w hi c h i s n ot i n eq u i
,
lib ri u m with i ncid ent for ces t o th a t m ore com pl ex f or m which

,
i s i n equ ilib r i u m w ith th e m We h a v e to m a r k h ow i n c orre

.
,
s p on den ce w ith th e u n iv e r s a l l a w t h a t th e u n i for m l a p s es i n t o
th e m u lti form an d th e l eS S mu ltif or m i nt o th e m ore mu lti

,
for m th e p art s of e a ch or g an i s m a r e e v er b ecomi ng f u rth er

,
di ffe renti a te d ; an d we h a v e to trace the v aryi ng rel a ti ons to

i nc id ent for ces by whi ch fu rt h er di fferen ti a ti on s ar e ent a il ed .
We h a v e t o ob serv e too how each p r im a ry m odifi ca ti on of

, ,
s tru c tu r e i n d u ce d by an a l te r e d di str ib u ti on of for ces b ecom e s

, ,
a p a r en t o f secon d ary m odi fi c a ti on s— h ow t h r ou gh th e n ece s ,
s a r y m u l tipli ca t i on of e f fec t s ch a nge of for m i n one p art

,
b ri ngs a b ou t c h an ges of form i n oth er p a rt s A nd th en we .
h a v e a l s o to n ot e the m et am orph os es c ons tantly b e i n g i ndu ced

by the p roc ess of s egr eg ati on— by the gr a d u a l u n i on of lik e
p a rt s exp ose d to lik e for ces an d th e g ra du a l sep a ra ti on of lik e
,
p a rt s e xp osed to u n lik e forces The f a ct or s of th e s econ d

.
c l a ss whi c h w e h a v e t o k ee p i n Vi ew th r ou gh ou t ou r i nt er r et
p
a ti on s a r e th e f or m a tiv e t en d en c i es of org a n i s m s t h e m s e lv e s
,
— the p roclivi ti es i nh er it e d by th em from a ntece d ent or g a n
i sms and which p as t p rocess es of ev ol u ti on h a v e b equ eathe d

,
.
We h a v e s een it to be i n fera bl e from v a r i ou s ord e rs of fa cts

65 84 97 , , th a t org a n i s m s a r e b u ilt u p of c ert a i n
high ly com pl ex m ol ecu l es whi ch w e di sti n gu i s h e d a s physi o
-
,
l ogi ca l un i ts [ Or consti tu ti ona l u ni ts a s th ey might oth erwi s e

be ca ll e d ] e ac h ki n d of or g ani s m b ei n g b u ilt u p of u nit s p e
-
c u l i a r to i t sel f We r ecogn i zed i n th ese u n i ts p ower s of a r

.
,
ra n gi ng th e m s e lv es i n t o th e for m s o f th e org an i s m s to w hi c h
th ey b el ong ; ana l ogou s to the p owers whi ch the m ol ecu l es of

i norg ani c su b s t ances h av e of a ggrega ti ng int o s p ecific c ryst a l
PR O B LE M S M O RPH OLO G Y
“
TH E OF .
9
li ne form s We h a v e con sequ ently to r egar d thi s p roclivity of

.
the phy s i ol ogi ca l u ni ts a s p rod u ci ng d u ri ng th e d e v e l op m ent

, ,
of a n or g a n i m a co mbi n a ti on of i nter n a l f orce s th a t exp en d

y s ,
th em selves i n wor ki ng ou t a stru ctu re i n equ ilib ri u m wi t h

th e f orc es to whi ch a n ces t ra l org a ni sm s w ere e xp ose d ; bu t
n ot i n e qu ilib r i u m w ith th e for ce s to w hi c h th e e xi s ti n g or g a n
i s m i s e xp os ed if the envi ronm ent h a s b een c h a n ge d

, H enc e .
the p r obl e m i n a l l c a se s i s to a sce r t a i n th e r esu lt a nt of i nt er

,
n a l org a n i z i n g f or c es t en di n g to rep r od u c e th e a nc es t ra l f or m
, ,
a n d e xt ern a l m odifyi n g f orc es t en di ng to c au s e d evi ati ons

,
f rom th a t for m Mor eov er we h a v e to t ak e i nt o a c cou nt

.
, ,
n ot on ly the ch a ra ct er s of im m edi a t e ly p re c e di n g a n c es t or s -
,
bu t a l so th os e of th e i r a nce st ors a nd a ncestoi s of a l l d egree s ,

'
o f re m ot eness S et ti n g ou t w i th ru dim ent a ry typ es w e h a v e

.
,
to c ons id er h ow i n eac h s u ccess iv e s t a ge of e v ol u ti on th e

, ,
st ru ct u res a cqu i r e d d u ri ng p revi ou s s t a g es h a v e be en oh

scu re d by f u rt h er i nt e g r a ti on s a n d fu r th er di f feren ti a ti ons ;
or ,
con v er s e ly h ow th e li nea m ent s of p r i m itiv e org a n isms
,
h a v e al l a l ong c onti nu e d to m anif est th ems elv es u nd er the

s u perp os e d m odific ati on s .
§ 1 7 9 T wo w ays of c arryi ng on the i nqu i ry s u ggest th em

.
s elv es .We m ay g o th r ou gh th e s ev er a l great g rou ps of

or g a n i s ms with th e Vi e w of r ea chi n g by c omp a ris on of p arts
, , ,
c ert a i n g ene ra l t ru th s r es p ecti n g the h om ol ogi es th e for m s , ,
a n d th e r el ati on s of th ei r p a rt s ; a nd th en h a vi n g d ea lt w i t h ,
the ph enom en a i n du c tiv ely m a y r et ra c e ou r s t e p s w i th th e

,
Vi ew of d e d u c tiv ely i nt er p reti n g th e g en era l t ru th s reac h ed .
Or i n st ea d of thu s s ep a r a ti ng th e two i n v es tig a ti ons we

, ,
m a y c a r r y th e m on h a n d i n h a n d— fi rst es t a bli s hi n g e ac h
gener a l t ru th em pi r i c a lly an d th en p roceedi n g to th e ra

,
t i on al e of it T hi s l a st m eth od w ill I thi n k c ond u c e to

.
, ,
b oth b revity a nd cl earn es s L et u s now thu s d ea l with the .
fi rst cl as s of m orph ol ogic al p robl em s .
[ N OT E — In p rep arati on for t re a ti ng of morphol ogic a l de

10 M O RPH OLO G I CA L DE VELO PM E N T .
vel opm ent, su n d ry oth er

genera l con si d er ati ons s h ou ld h av e
b een in c lu d e d i n the foregoi ng ch a p ter w h en origi n a lly pu b
l i sh e d. T hi s s eem s the m os t a pp rop ri a te pl ace for n ow n a m
i ng th em S om e w ere impli ci tly c onta i n e d i n the fi rs t vol
.
u m e bu t i t w ill be w e ll d e fi ni te ly to s t a te th e s e a s w ell as
, ,
the oth er s n ot yet i m pli ed .
I nt er p reta ti on of the for m s of orga ni s m s and th e f or ms

of t h e i r p a rt s m u s t d ep en d m a i nly on th e c on cl u s i ons re
, p
vi ou s l y d r a wn res p e c ti n g t h e i r phyl og en y ; a n d t he d r a wi ng
of s u ch c onc l u s i ons m u s t be gu id e d by recog n i ti on of th e
v a r i ou s f actor s of E v ol u tion a s well a s by r ecogn i ti on of

,
c erta i n ext rem ely g ener a l resu l ts of E v ol u ti on an d c ert a in

c oncomi ta nts of E v olu ti on .
A p r i m a ry one a m on g t h es e is th a t no exi sti n g s p eci es c a n

e xh ibi t m or e th a n a pp roxi m a tely the a nc est r a l s tru ctu r e of
a ny oth e r exi st i n g s p ec i es A s a l l a n ces t or s h a v e di sa pp eare d

.
,
so i n a g rea t er or l e s s d eg r ee th e t r a i ts s p e c ific g ener i c or

, , , , ,
or din a l whi c h di s ti n gu i s h e d th e ea r li er o f t he m h a v e di s a p
,
p ear ed . S etti n g ou t w i th th e fa m ili a r sy m b ol a t ree l et u s , ,
re g a r d i ts p e r iph e r a l t w ig s a s r ep r e s e nti n g e xt a nt s p e ci es ;
l et u s a ssu m e t h a t the i nt eri or of th e tre e i s fill ed u p w ith

s om e su pp orti n g su b st anc e l ea vi n g only the end s of the
,
livi ng t wigs p roj ecti ng ; a nd l et u S su pp ose the t ru nk m a in ,
b ra nch es s econd a ry b ra n ch es t erti a ry b ra nch es &c h av e

, , ,
.
,
d ec ay ed a wa y T h en i f we t a ke th ese d ec a ye d p a rt s to s t a n d
.
for the div ergent a n d re div er g ent li nes of ev ol ut i on which

-
i t w ill be
’
a r e re p r ese nt e d by foss il s i n th e E a r th s c r u s t ,
m a ni fes t fi rs t th a t no one of th e livi ng su p erfi ci a l t wig s ( or

, ,
sp ec i es ) e xhibit s th e a n cestra l org a n i za ti on wh en ce a ny oth er

of th e livi ng s u p er fici a l t w ig s ( or s p eci e s ) h a s b ee n d e v el op ed ;
i t will be m a ni fes t s ec on d th a t the gener i c s t ru ctu re i n

, ,
h er i ted by a ny e xi sti n g s p ec i es m u st be a s tru c tu r e ou t o f

whi ch c a m e su nd ry a lli e d s p eci es— th e for k a s i t we re a t , ,
whi ch a dj a cent t w ig s dive rged ; a nd thi r d th at the a ncest or ,
o f a n or d e r m u s t i n lik e m a nner he s ou ght a t s om e p oi nt

, ,
d eep er d own i n th e symbolic t ree— a pl ace of div ergence of

TH E PR O B L E M S OF M O RPH OLO G Y .
11
the s u b-
b r anch e s rep resenti n g a lli ed gener a S imil ar ly with
.
t he anc es t ra l typ e s of c l a sses s till d ee p er d own i n th e tree or

,
fu rth er b ac k i n ti m e S o th a t phyl og eny b ecom es m or e an d

.
m ore sp ecu l a tiv e as its qu es ti on s b ecom e m or e a n d m ore
r a dic a l . A n d the di ffi cu l ty is m a d e g reat er by th e d efici en cy

of p a l aeon t ol ogic a l e vid en c e .
One obvi ou s c oroll a r y i s th a t a n a n c est r a l typ e f rom w hi c h

su nd ry a lli ed typ es now e xi sti ng div erged w a s S p ea ki n , g ,
g enera lly si m pl er th an th es e ; Si nc e th e div erg ent typ es he

,
c a m e di fi erent by th e s u pe rp os i ng of m odifi c a ti ons a ddi ng ,
to th ei r c om pl exiti es T h ere i s a fu r th er r ea son for inf err i ng

.
th a t the l ea s t s p eci a li ze d m em b er of a ny grou p i s m or e lik e

th e rem ot e a nc est or th a n a ny of th e oth ers ; for e v ery a d a p
fa ti ou st a nd s i n the way of s u b sequ ent re a d a pt a ti on s : it -
p res ent s a g rea t er a m ou nt of s t ru ctu re to be u nd one T o g et .
s om e id ea of th e a ncest r a l typ e w h ere n o ext a nt m e mber of
the grou p i s m a ni fes tly S i m pl er th an th e res t th e m eth od ,
m u s t be to t a k e al l i ts ext ant m em b er s a n d a fter l etti n g ,
th ei r differenc es mu tu a lly c a nc el ob ser ve wh a t rem a i ns com

,
mon to th em a l l .
Bu t th er e a r e di ffic u lti e s st a n di n g i n the w ay of phyl ogeny ,
a n d c on s equ ent ly of m or ph ol ogy mu ch g rea t er th a n th ese

, .
Ret u rn i n g to ou r sy m b oli c t ree it i s cl ea r th a t it wou ld be

,
far fr om eas y to s a y of a ny one t wig w hi c h exti nc t su b

b r an ch b r anch and m a i n b ra nch it b el on ge d to ev en su p
, , ,
p os i n g th a t the g rowth s of a l l p a r ts h ad b een u n iform ly ou t

w a rd s. I m m ense ly m ore p erpl e xi n g th en mu st be th e
, ,
a ffili a ti on if v ar i ou s of th e b r a n c h es s u b b r an ch es &c
-
have ,
, ,
.
s en t ou t b a c kw a r d g rowi n g s h oots which h a v e c om e to th e

-
s u rfa c e on ly a ft er p rol on g e d r et rog r a d e cou rse s a n d if oth e r

,
b ra nch es h av e s ent Sh oots i nt o regi ons oc cu pi ed by a li en

b r anch e s— S h oots b ear i ng t w igs whi c h com e to the su rface
a l on g with th os e t o w hi c h th e y a r e bu t re m ot e ly a lli e d T he .
p robl ems of origi n a nd of s t ru ctu re whi ch or g ani s m s p resent ,
a re m et by b oth of th e di ffic u lti e s thu s s ymb oli z e d .
On e of th e m a r i ses f r om the p rev a l enc e of r et rogra d e

m et am orph oses T h rou gh ou t the a n i m a l w or ld t h e s e a re

.
v a ri ou s ly di s pl ay e d by p a r as i te s m u l titu di nou s i n the i r

,
ki nd s ; for m os t of th em b el ong to typ es mu ch high er i n

org a n iz a ti on . Ch an ge d h a bi ts a nd con sequ ent c h a nged st ru e
t u res h a v e s o t r a n s ferred th em th at only by s tu dy of th ei r
e m b r y on ic s ta g es c a n th e i r ki n ship s be m a d e ou t A n d th ese .
r et rog r a d e m et a m or ph os es c on s pic u ou s a m on g p a r a s it es h a v e
, , ,
i n t he cou rse of e v ol u t i on a ffect e d s om e m emb er s of a l l

,
g rou p s ; for i n a l l g rou p s the stru ggl e for exi stence h a s com
el l e d s om e t o a d op t c a r eer s l e s s t ryi n g bu t l ess p r ofit a bl e
p .
N ot only by forc i n g on m a ny ki n d s of orga nisms simpl er

w a y s of livi ng a n d con sequ ent d eg ener acy h as the u niv ers a l
, ,
c om p e titi on c a u se d ob s c u r i n g t ran s for m a ti on s I t h a s d one

.
thi s a l so by t empti n g m any oth er ki nd s of or g ani s m s

to a d opt w a y s of li fe not simpl er th a n b efore bu t m erely
di fferent P ressu re conti nu a l ly p r om pt s ev er y typ e t o i n
.
t ru d e on oth er typ es s ph er es of a ctivi ty ; a nd so c au ses it to

’
a ssu m e c ert a i n st ru ctu ra l ch ara ct er s of th e t yp es wh ose

S ph e r e s it i nv a d es m as ki n g its p revi ou s ch ara ct er s
, Modi fi .
ca t i on s h enc e a r i s i n g h a v e i n th e g r ea t m a ss of c a s es
,
b een ,
s u p er p osed one on another tim e a fter tim e T h e a qu a tic

.
a ni m a l b ecom e s th r ou gh se v er a l t r an s i t i on s a l a n d a n im a l -
,
a n d th en th e l a n d a n i m a l th r ou gh oth er t ra n s iti on s b e c om e s
-
now a n a e r i a l a n i m a l lik e th e ba t an d n ow a n a qu a tic a n im a l

lik e the wh a l e Cert a i n ki nds of bi r d s f u rni sh ext r em e
.
illu stra ti ons . T h ere wa s th e ch a ng e f rom the fish to the

w at er b rea thi n g a mphibi an an d th en to the a i r b reathi ng
- -
’
a m phibi a n ; t h en c e t o the re p t il e livi n g on th e E a r th s s u r
f ace ; th ence to the flyi ng r eptil e an d the bi r d ; th en c am e

t he divi ng bi rd s j oi ni ng w i th th e i r s é ri a l li fe a li fe p assed
,
p a rtly i n the wat er ; and fi na lly c a m e a typ e like the

p engu i n i n whi c h th e p ower of flight h a s b een l ost and the
,
w a te r h a s a g a i n b ecom e the a lm ost ex c l u s iv e m e di u m ex ce p t ,
for b r ea thi ng . Of c ou r s e th e m ou ldi ngs a nd re m ou ldi ngs of -
s t ru c t u r e r esu lti n g from th es e s u cc ess iv e u nlike m od es of
li fe i n m a ny c ases pu t gr ea t di ffic u lti es i n the way of a scer

,
TH E PR O BL E M S OF M O RPH OLO G Y .
13
t a i ni ng whi c h are the or igi na l c orrespondi n g p a rt s S om e .
p a rt s h a ve b ecom e a b nor m a lly l ar ge ; oth er s h a v e dwi n dl e d

or di s a pp e a re d ; a n d th e r el a tiv e p os iti ons of p art s h a v e often
’
been g rea tly ch an g e d A b a t s W i ng a n d a bi r d s Wi ng a re
’
.
a na l og ou s or g ans bu t th ei r fr a m ewor ks are bu t p ar ti a lly

,
h om ol ogou s Whil e i n the bi r d the t er mi na l p art s of th e

.
fore li m b do littl e t owa r d s s u pp orti ng the wi n g i n th e ha t

-
,
th e w i ng is m a i nl y s u pport e d by enor m ous ly d ev el op e d t er mi

-
na l p arts .
T h e effects of th e s t ru ggl e to s u r viv e which h ere p rompt s

,
a s i mpl er li fe wi th resu lti n g d e g en er a cy a n d th ere a di fferen t
life w ith resu lti ng n ew d ev el opm ent s a re fa r f rom b ei ng the

,
onl y c au s e s of m orph ol ogic a l ob s cu r a ti ons F ul film ent of

.
c ert a in hi gh ly genera l req u i rem ent s giv es c ert a i n c ommon

t ra its to pl a nts of wid ely div erg ent cl a s ses ; and fu lfilment of
cert a in oth er highly gen era l requ i rem ents gives c ert a i n
c omm on t ra its to an im a l s of wid ely div ergent cl a s s es I t wa s .
r em a r k ed i n th e fi r s t v ol u m e th a t the c ar dina l di sti nc

ti on be t ween the ch ara ct er s of pl ants a nd a nim a ls a ris es f rom
the f a ct th a t whil e th e c hi e f food of pl ants is u niv ersa lly
p re s en t the f oo d of a ni m a ls i s s catt ere d H ere it h a s to be
.
a dd ed th a t to u tili z e th e u ni v er s a lly di s t r ib u t e d f ood th e
or di na r y pl a n t n ee d s th e a i d of light an d h a s to a c qu i re
,
st ru ctu res ena bli ng it to g et th at ai d ; whil e the or di nary

a ni m a l to u tili z e th e s c att ere d f ood m u st a cqu i r e th e st ru e
, ,
t u r es n eedfu l for l ocom oti on L et u s cont em pl at e s ep a rat ely

.
the t ra its h en c e r esu lti n g i n the v e get a l w orld a nd the t ra its

h en ce resu lti ng i n th e anim a l world .
T h e f a m ili a r pl a n t a in m eets th e r eq u i rem ent by g row i n g

s tiff l ea v e s e na bli n g it t o p r ess d own th e c om p eti n g g ra ss e s
a r ou nd whi c h wou ld el se sh a d e it ; bu t th e gr ea t m a j or ity of
o r di na ry pl an t s m ee t th e r eq u i re m en t by ra i s i n g th e ms elv e s
i nt o the ai r H ence the need for a s tem a nd h ence the f a ct

.
,
th a t pl ants of wid ely u nlik e n atu res s i m il ar ly form st e ms

whi ch i n a chi evi n g st rength enou gh to su pp ort the f oli a ge
,
a nd res i s t th e w i n d a cqu i re c er t a i n a d a ptiv e s t ru c tu res h av

,
14 M O RPH OLO G I CAL D E V EL O PM E NT .
i ng a g enera l simil ar ity H er e from the edge of a p ool i s

.
a re e d a n d h e re fro m the a dj a cen t cops e i s a h eml ock : t he

,
one h a vi n g gr own t a ll in esca pi n g th e s h a d e of i t s com
p a ni ons and the oth er i n es c a pin g th e sh a d e of th e su rround

ing b rus hwood On be i ng cu t a cros s eac h dis c l os es a t u b e
.
,
a n d ea c h exhibit s s e pt a dividi n g thi s t u b e i nt o c h am b ers .
I n eith er case by the tubu l a r s t ru ct u r e is g a i ned th e gr ea t e st

s tren gt h w i t h th e l ea st m a ter i a l ; bu t th er e is n o m orph o
l ogic a l ki nship b etw een the tu be s nor b etween the s ept a .
S till m or e m ark ed i s the s i m u l a ti on of h om ol ogy by an a l ogy

i n a noth er pl a nt whi c h the a dj a ce n t dit c h m a y f u rn i s h— th e
c omm on H orset a il I n thi s a ga i n w e see a n el ong at ed ver
.
, ,
ti ca l g row i ng p art ra isi ng the foli a ge i nt o the a i r ; and as

-
, ,
b ef ore this is tu bu l ar and divid e d by s ept a A typ e u tt er ly

,
.
a li en f rom th e oth er tw o h a s by su r viv a l o f th e fitt e st b een

, ,
si m il arly m ou ld ed to m eet m ech anic a l needs .
P a ss in g now to th e ob scu r at i on s i n th e a nim al w orld

c au s ed by a lt era ti ons fav ou r i ng l ocom oti on we not e fi rs t th a t ,
the l ocom ot iv e p ower is a t th e ou t set v ery slight A m ong .
m any or d ers of P r otozoa a s a l so a m on g m any l ow type s of

,
Meta zoa vib ratil e cili a a re the m ost gener a l agents of l oc o

,
m oti on — nec ess a rily f eebl e l ocom oti on Reg ar d ed i n th e .
m a s s the Cael en tcra ta wh en not st a ti on ary lik e th e H ydra or

, ,
high er typ es i n th e hyd roid st a ge u su a lly p osses s only s u ch

,
s m a ll s el f m obility a s the sl ow rhythmic a l c ontra cti ons of

-
th ei r u mb rell a di sk s effect or el se su ch a s i s effect ed by b and s

-
,
of cili a or of vib r a til e pl a t e s as i n th e B er oc ,

E v en a m ong .
th ese l ow tpes of M eta zoa h owev er i n which ordi na rily the

, ,
ra di a l s t ru c tu r e i s con s pi cu ou s or bu t slightly ob s cu re d by
,
a n ov oid f or m a s i n th e C ten o hora w e fi nd i n th e C es i us

p , ,
ven er i s e xt rem e ob s cu r a ti on c au s e d by a n e l on g a t i on which

,
f a cilit ates m ov em ent th rou gh the w a t er ; a lik e by the a cti ons

of i ts vib ra til e pl a t e s a n d by i ts u nd u l a ti ons whi c h simu l at e
,
those of su nd ry high er a nim als .
A nd h er e we c om e u p on the ess enti a l f a ct to be rec ogni zed .
E l onga ti on f a v ou rs l oc om oti on i n v ar i ou s w ays th at are

T HE PRO BLE M S OF M O RPH OLO G Y .
15
v
s e era lly t ak en a dv ant a ge of by different typ es of c reatu res .
giv m ss of m ovi ng m a tt e r th e r es i s t a n c e of th e
( )
1 T o a e n a
m e di u m d ecrea s es a l on g wi t h d ecre a s e i n the a r ea Of i t s
t r a nsvers e secti on a nd this impli es i ncrease of l ength : a giv en

,
for ce wi ll m ov e the l en gth ene d m a s s a l on g with g rea t er

f ac ility ( 2 ) Rea chi ng a c ert a i n p oi nt the el onga t e d f or m
.
en a bl es a n a n i m a l to p rog res s by u n d u l a ti ons a s i n th e ,
w a t er fi sh do an d ev en s om e c oel ent era t es and t u r be ll a ri a ns

,
do an d a s on l a n d sn a k es do : l a t era l re si st a nces s ervi ng i n

,
en i ng of th e body s erv e s
e ith e r c a s e a s fu lc r a ( 3 ) L e n gth
.
oth er wi se to a i d l oco m oti on i n the c ree pi n g or b u rrowi ng
worm which u tili z i ng the st a ti c a l resist an c e of its hi nd er

, ,
p art thru st s onw ards its fore p a rt and th en h oldi ng f a st its , ,
fore p a rt by th e a i d of mi nu t e s etce d raws the hi n d er p art ,
a ft er it Bu t e l on ga ti on d ou bly a dv a nt a g eou s at fi rst w hil e

.
, ,
th e body i s its elf th e chi e f i n stru m ent of l oc om oti on gr a du a lly ,
l os es its a dv ant a geou s ness a s s p eci a l i n st ru m ents of l ocom o

ti on a r e d evel op ed ( 4 ) T his we see i n th a t l oc omotiv e
.
a cti on effect ed by limbs which m a n y a n d sm a ll i n th e l ower

, ,
A rthrop oda a nd b ec omi n g few an d l arger i n the high er a t ,
l ength giv e gr ea t a ctivity to a sh ort ened and c onsolid ate d

b ody : a s tage rea che d only th r ou gh st a ges of d ec rea si ng
e l ong a ti on a c c omp an yi n g i n c r ea s e of limb p ower ( )
5 I n -
.
the Ver tebra ta l oc om oti on by u n d u l a ti on s c om es a l on g cer ,
t a i n li nes of ev ol u ti on to be r epl a c ed by th a t limb l ocom o

,
-
ti on which a ccom p ani es th e r i se from wat er life to l an d life - -
the ev olu ti on Of A m phibi a n s e xhibi ti n g th e t r an s iti on (6) .
F u rth er we s ee a m ong m a m m als th a t as limbs b ecome effi

,
ci ent the el ong ate d b ody ce a s es to be it s elf i nst r um ent al i n

l ocom oti on bu t th a t still s om e el ongati on rem ai n s a ch a r ac
,
ter i sti c .
( 7 ) Fi n ally wh ere limb l oc omoti on rea ch es i ts high
,
-
est d e gr e e a s i n bi r d s e l on g a ti on di s a pp ear s
, , .
T h ese c l a sse s of f a mili ar f a cts I h a v e r ec a ll e d to Sh ow

th at in the c ou rs e of ev olu ti on a chi ev em ent by pl ants of the
, ,
a l l ess enti a l el ev a ti on i nt o th e a i r a nd by an im a l s of th e
-
a l l e s senti a l p ower of m ov em ent h a v e d e v el op e d this t ra it

-
16 M O RPH OLO G I CAL D E V E LO PM ENT .
of e onl ga ti on i n v a r i ou s typ es ; and th a t i n eac h ki ngd om

a c qu i s i t i on of th e c om m on t ra it h a s h a d a t en d en cy n ow to
ob sc u re m or ph ol ogi ca l equ iv a l en c e an d n ow to giv e th e a p

,
p ea r a n c e of ki n s hip W h ere th ere i s n on e . A f u rth er

p ur
p os e h a s b een to p rep are th e way for a qu esti on h er ea ft er

t o be di sc u s se d— w h et h er i n the v ar i ou s typ es of e ith er ki ng
,
dom th e el ong a ti on i s effe ct e d i n th e s a m e w ay s or i n di f

,
fer ent w a ys We s h a ll h a v e to ask wh ether th e v ertic a lly

.
g rowi ng p art is a l w a ys lik e th a t of L ess oni a a s impl e i n

, ,
dividu a l or wh eth er a s p oss ibly i n P h aenoga ms it i s a un it ed

, , ,
s eri es of i ndividu a l s ; and si m il a r ly wh ether the el ongat ed

b ody is a lwa ys si ngl e lik e th a t of a m ollu sc or wheth er as
, , ,
pos sibly i n annu l os e anim als, it is a s er i es of u nit ed i n

di vi du al s .
]
CH A P T E R II .
TH E M O RP H OLO GI C A L C O M P O S I T I O N OF P LA N T S .
§
l 8 o E VOL U T I O N
. impli e s i n s en sibl e m odific a ti ons a n d
gra du a l t ra n si ti on s whi c h ren d er d efi niti on di ffi cu l t— whi c h
,
m a k e it i m p ossibl e to s ep a r a t e a b s ol u t ely the ph as es of

or g an iz a ti on fr om one a noth er An d thi s i n defi ni tenes s of
.
di stin cti on to be exp ected a p ri ori we a re compe ll e d to

, ,
r ecog ni z e a p os ter i or i th e m om en t w e be gi n to g r ou p m orph o

,
l ogic a l ph enom ena i nt o genera l p rop os iti ons T h u s on .

,
i nqui ri n g W h a t i s the m orphol ogi ca l u n it W h eth er of pl ants ,
o r o f a n i m a l s we fi nd th a t th e fa c ts re fu s e to be i n cl u d e d i n
,
a ny r igid f or m u l a . T he d oct r i ne th a t a l l org an i s m s a re b u ilt

u p of c e ll s or th a t c ell s a r e th e el e m en ts ou t of whi c h e v e ry
,
ti ssu e i s d ev el op e d i s bu t a pp roxi m a t ely t ru e T h ere a re

, .
li vi ng for m s of whi ch c ellu l a r s t ru ctu r e c ann ot be a ssert ed :
an d i n livi ng f orm s th a t a r e for th e m ost p art c ell u l a r th er e ,
a r e n ev e rth e l ess ce r t a i n p orti on s which a r e not p r od u c e d by
th e m et a m or ph os i s of c ell s S u pp os i n g th a t cl a y wer e th e
.
only m a ter i a l a v a il a bl e for b u ildi n g th e p r op os iti on th a t a l l

,
h ou ses a re bu ilt of b r i ck s wou ld hear a bou t the s a m e r el ati on

,
to th e t ru th a s d oe s th e p r op os iti on th a t a l l org an i s m s a re
,
c omp ose d of c e ll s . T his genera li z a ti on res p ecti ng h ou s es

w oul d be Op en to two critici s m s — fi rst th a t c ert a in h ou ses ,
of a p r i m i t iv e ki n d a re for m e d n ot o f b ri c ks bu t ou t o f ,
,
u n m ou ld e d c l a y ; a nd s ec on d th a t th ou gh oth er h ou s e s con
,
s i s t m ai nly of b r i ck s
yet th ei r chim n ey p ot s d ra i n pip es and
- -
, , ,
48
18 MORP H OL OGI OA L DE V E LO PM E NT .
r idge til es do n ot resu lt f rom c om bi na ti on or m et a m orph osis

-
,
of b r i c k s bu t a r e m a d e di r ectly ou t of th e origi n a l c l a y
, .
A n d of lik e n a t u r es a re th e c ritici s m s which m u st be p a s se d

on th e g ener a li za t i on th a t ce ll s a r e th e m orph ol ogi ca l u ni ts
,
of or g a ni s m s T o conti nu e th e s imil e th e tru th t u r n s ou t t o

.
,
be th a t the p r imi t iv e c l a y or p r ot opl a sm ou t of w hi ch

,
or g an i s ms a re b u il t m ay be m ou ld e d e i th er di re ctly
,
or ,
wi th v a ri ou s d egr ees of i ndi rect nes s i n t o or ga nic s t ru ctu res

.
, .
T h e phy s i ol ogi c a l u n it s whi ch we a r e oblig e d to a s su m e a s

th e c om p onent s of thi s p rot opl a s m m u st a s w e h a v e seen
, , ,
be the p ossessor s of th os e p r oc liviti es w hi c h r esu lt i n

'
th e st ru ctu ra l a rr a n gem ent s of th e org a n i s m T he a ssu m p .
ti on of su ch s t ru ct u r a l a rran gem ent s m ay g o on a nd i n ,
m an y cases d oes g o on by th e S h ort es t rou t e ; w i th ou t th e

,
p ass a ge th rou gh wh a t we c a ll m et a m or ph oses Bu t wh ere .
s u ch st ru ctu ra l a rr a n gem ent s a re reach ed by a c i rcui tou s

rou te t h e fi r s t s ta ge i s th e f or m a t i on of th es e s m a ll a ggr e
,
g a t es whi ch u nd er th e na m e of cells a re cu rrently r eg ard ed

, ,
a s m orph ol ogi c a l u nits .
The ra ti on a l e of t h ese t ru ths a pp ea rs to be f u rni sh e d by

th e hyp oth es i s of ev ol u ti on We set ou t with m ol ecu l es
.
s om e d eg rees high er i n c ompl exi ty th an th ose m ol ecu l es of

n i trog enou s c oll oid a l s u b s t an c e i nt o whi ch or g a n ic m a tt er i s
res olv a bl e ; a nd w e re g ar d th e s e v er y mu ch m or e c ompl e x
m ol ecu l es a s h aving the impli ed g reat er i ns ta bility gr eat er ,
s en sitiv e ne s s to s u rr ou ndi n g i nfl u enc es an d c ons equ ent

,
gr ea t er m obility of for m S u ch b ei ng the p ri m itiv e phy s i o

.
l ogic a l u n i ts organ i c evol u ti on mu s t b egi n with th e form a ti on

,
of a mi nu t e a ggre g a t e of th em — a n a gg re g a t e s h owi n g vit a li ty
by a high er d e gr ee of th a t rea di ne ss to ch ange i ts for m of

a gg r eg a ti on w hi c h c oll oid a l m a tt e r i n g ene r a l di s pl a y s ; a n d
by i ts a bility to u nit e the nit rogenou s m ol ecu l es it m eet s

with i nto c ompl ex m ol ecu l es lik e th ose of whi ch i t i s c om
,
p os ed Obvi ou sly the earli es t form S mu st h a v e b een mi nu te ;

.
,
si nce i n the ab senc e of a ny bu t diff used org ani c m a tt er no

, ,
f orm bu t a mi nu te one c ou ld find nut rim ent Obvi ou sly too .

, ,
TH E M O RPH O LO G I CA L C O MP OS I T I ON OF P L A NTS .
19
it mu s t h a v e b een st ru c tu rel ess ; si n c e a s differen ti ati ons a r e

,
p rod u c ibl e only by th e u nli ke a cti ons of i ncid ent forces th ere ,
c ou ld h av e b een no differen ti a ti ons b efor e s u ch forc es h a d h a d

ti m e to w or k . H enc e di sti n cti on s of p ar t s lik e th ose requ i red
,
t o c ons tit u te a c ell w ere n ec e s s a r ily a b s ent a t fi rst A n d we .
n eed not theref or e be s u rp ri se d to fi nd a s we do fi nd s pe cks, ,
of p rot opl a s m m a n i fe s ti n g lif e a nd yet S h owi ng n o s ign s of

,
or g a n i z a ti on . A fu rth er s t a ge of ev ol u ti on i s
r ea c h e d w h e n th e imp erfectly i n t eg ra te d m ol ecu l e s f or m i ng
on e of th e s e m i nu te a gg r eg a t e s be com e m ore coh erent ; at

,
the s a m e ti m e a s th ey p a s s i nt o a s ta t e of h et erogeneity ,
g ra du a lly i ncrea s i ng i n its defi ni tenes s T h at i s to s a y w e.

,
m a y l ook for the a ssu m pti on by th e m of s om e di s ti n c ti on s of

,
p a rt s s u ch a s we find i n cell s a nd i n wh a t a re ca ll e d u ni
,
c e ll u l a r or g a n i s ms . T h ey ca nnot ret a i n th ei r p r i m or di a l
un i for m i ty ; a n d W hil e i n a few c a se s th ey m ay d ep a rt f rom it
bu t s lightly th ey w ill i n th e gr ea t m a j or ity of cases a c qu i re

, , ,
a d ec id e d m u l ti for m i ty : t h e r e w ill r esu lt th e c om p a r a tiv ely
in t egr a t ed an d comp ara tiv ely di ffe renti a t ed P r otop hyta a nd

P ro tozoa . T he p r od u c ti on of mi nu t e a ggreg a t es
o f p h y s i ol ogi c a l u n it s b e i n g th e fi r s t s t e p a n d th e p a s s a g e
,
o f s u c h mi n u te a gg r e g a te s i n t o m or e c on s olid a t e d a nd m or e
compl e x f or m s b ei n g th e s ec on d s t e p it m u s t n at u r a lly
,
h a pp en th at a l l high er org a n i c typ es s u b s equ ently a ri s i n g by

,
fu rt her i nt egr a t i on s a n d di fferenti a ti ons w ill e v erywh ere b ear

,
th e i m p r ess of thi s ea rli es t ph a se of e v ol u ti on F rom the .
l a w of h ere di ty cons id ere d a s e xt en di n g to the en ti re su c

,
’
cess i on o f livi n g t hi n g s d u r i ng th e E a rth s p as t hi s t or y it ,
foll ow s th a t s i n ce the form a t i on of th es e s m a ll s impl e orga n ,
i s m s mu st h a v e p r ec ed e d the form a ti on of l a rger a nd m or e

com pl ex or ga n i sms th e l a r g er a n d m ore c ompl e x or g a n i s m s
,
mu st i nh er it th ei r ess en ti a l ch a racter s We m ay a nticip at e.
th at the mu l tipli c ati on a nd com bi na ti on of th es e mi nu t e

a ggr e g a tes or cells wi ll be c on s pi cu ou s i n th e ear ly d e v e l op
,
m ent a l s ta g es of pl ant s a n d an i m a l s ; a nd th a t th rou gh ou t

a l l s u b s eq u ent s t a g es c ell p r od u c ti on a nd c ell di f
,
-
ferenti a ti on -
20 M O RPH OLO G I C A L D E VE LO PM ENT .
will be d om i nant ch ar ac teri stic s T he phy s i ol ogic a l u nit s .
p ecu li ar to each higher sp e ci es w ill s p ea ki n g g ener a lly , ,
p a ss th rou gh this for m of a ggr eg a ti on on thei r w ay t owa r d s

t h e fi n a l a rr a ng e m ent th ey a r e to assu m e ; b ec au s e t h os e
p r i m ordi a l p hy s i ol ogi ca l u ni ts from whi ch th ey a re rem otely
d es c en d ed a ggr eg at e d i nto thi s form An d yet j u st a s i n
,
.
,
oth er c a s e s we fou n d r ea s ons for i n f erri n g 1 3 1 ) t h a t th e

t ra it s of an c estra l org ani z a ti on m ay u n d er c ert a i n c onditi ons , ,
be p a rt i a lly or wh olly obli ter a t e d and the u lti m a t e s t ru ctu re ,
a ssu m e d w i th ou t p a ss i n g th r ou gh th e m ; s o h e r e it i s to be , ,
i nfer red th a t th e p rocess of cell for m a ti on m ay i n s om e c a ses -

, ,
be p a s se d ov er T hu s t he hyp oth es i s of e v ol u ti on
.
p r ep a res u s for th os e two r a di c a l m odifi cati ons of the cell

d octri n e whi ch the f a cts oblige u s to m a k e I t l ea ds u s to .
e xp ec t t h a t a s s t ru ctu r e l es s p orti on s of p r otopl as m m u st h a v e
p reced e d cell s i n th e p roc ess of genera l ev ol u ti on ; s o i n th e ,
S p eci a l e v ol u t i on of ea c h high e r or g a n i s m th ere W ill be a n ,
h abitu a l p rodu c ti on of c ell s ou t of st ru ctu rel ess bl a st em a .
An d it l ea d s u s to exp ec t th a t th ou gh g enera lly the phy s i o , ,
l ogi ca l u nits c om p os i n g a s t ru ct u rel e ss bl a stem a w ill di s pl a y ,
th ei r i nh er it ed p rocliviti es by c ell d ev el op m en t and m et a -
m orph os is ; t h ere w ill n ev ert h el es s oc cu r ca s e s i n whi ch the

ti ssu e to be form ed i s f or m e d by di rect t ran s f orm a ti on of the
,
*
bl a ste m a .
L et m e h e re re fe r th os e wh o a re i nte re s te d i n thi s q uesti on ,

t o P rof .
Hu r l e y s ’
cri t i c i s m o n t he ce l l -doc t r i ne , p ubli s h ed i n th e M edic O Chi ru rg i ca l
R evi ew i n 1 8 5 3 .
A c ri ti c v
w h o t hi nk s th e a bo e s t a t e m e nt s a re ra th e r m i s l e a di ng ad
m i ts t h a t th e l o w e s t ty pe s of orga ni s m s yi el d t hem s u p p ort , s a yi ng th a t

of
“
th e re a re ce rt a i nl y m a s s e s p roto pl as m conta i m n rr m a ny nuc l e i , b ut no
t ra c e o f ce l l u l a r s t ruc t u re , in b ot h a ni m a l s a nd p l a nt s . S uch non-c el l ul a r
m a s ses m a y e x is t d uri ng de v e l o pm ent a nd l a te r b ecom e se

pa ra t ed u p i nto
c el l s , b ut th ere a re c e rt a i n low o rg a ni s m s i n whi ch s u ch m x
as s es e i s t i n th e
a d ul t s ta t e . Th e y a re c a l l ed by s o m e b o t a ni s t s non -
c e l l u l a r, by oth e rs
m u l t i -nucl e a tc cel l s . Cl e a rl v t h e di ffe re nce l i e s i n t he c ri te ri a of a ce l l .
Th e re a re al so s om e P r otozoa , a nd t h e Bo o/m i tt
,
i n wh ich no n u cl e u s ha s
ce rta i nl y bee n de m ons t ra te d . Bu t i t i s us ua l t o co n s id e r t h e b od i e s o f s u ch
o rga ni s m s a s c el l s v
ne e rt h e l e s s , a nd it is s up
po s e d th a t s u ch ce l l s re p re s e n t
T HE M O RPH OLO G I CA L C O MP OS I T I O N OF PL ANT S .
21
’
I nt erp reti ng the f act s i n thi s g i m a nn er , we m a y r ec o n z e
th a t l a rge a m ou nt of t ru th whi ch the c ell d octr i ne c ont a i n s -

,
W i th ou t c om m i tti n g ou rs e lv es to the e r ror s i n v olv e d by a

sweepi n g a s s e r ti on of i t We a re en a bl e d to u n d ersta n d how
.
it h a pp en s th at org an i c stru ctu r es a r e u su a lly ce ll u l a r i n t he i r

c om p os i ti on at the s am e ti m e th a t th ey ar e not u nivers a lly
,
so . We a re sh own th a t whil e we m ay p rop erly c onti nu e to

r e g a r d t h e c ell a s the m or ph ol ogi c a l u n it w e m u s t c on st antly ,
b ear i n mi n d th a t it is s u ch only i n a qu a lifi e d s en s e .
181 .ggreg a t es of th e l ow es t ord er eac h f or m ed

T h ese a ,
Of phy s i ologi ca l u n i ts u n i ted i nto a g rou p th a t i s s t ru ct u ra lly

S i n gl e a n d c a n not be divid e d w i t h ou t d es t r u ct i on o f i ts
i ndivid u a lity m ay a s a b ov e i m pli ed exi s t a s i nd ep en d ent

, , ,
org a n i s m s T h e a ssu m p ti on to w hi c h we a re com mi tte d by

.
the h yp oth es i s of ev olu ti on t h a t su c h so c a ll e d u n i c e ll u l a r

,
- -
pl a nt s were a t fi rs t th e on ly ki n d s of pl an ts i s i n h a r m ony ,
wi th the f a ct th a t h a bit a ts not occu pi ed by pl ant s of high er

ord e r s c om m on ly c ont a i n th es e p r ot ophyt es i n g rea t abu nd
,
a n ce a n d g rea t v ar i ety T he v ar i ou s sp ec i es of P l eu r ococ

.
ca c ece of D cs m i di a c ew a n d D i a tom a ceae su pply e x a mpl es of

, , ,
m or ph ol ogic a l u n it s livi ng a n d p rop a g a ti n g s ep a r ate ly un d er ,
nu m er ou s m odifi ca ti ons o f for m a n d s tru ctu re Figu res 1 2 .

, ,
an d 3 r e p r es en t a few of th e c om m one s t typ e s

, .
Mostly s i m pl e pl a nt s a re too s m a ll to be i ndivid u a lly

,
a s t a ge o f dev el op m en t i n wh i ch th e n uc l e u s ha s not ye t b een v ve d

e ol ,
t h ough
th e
’
ch em ical s ub s t a nce n uc l e i n h a s b e e n form e d i n s om e o f t he m
P e rh a p s i t w i l l b e x
m o st corr e c t to s a y th a t, e cl u di ng t h e m i nu t e, n on
nu cl e a t ed orga ni s m s , a l l t h e h i gh e r org a ni sm s — Meta zoa a nd M etap l zyfa — a re
com po s ed th rough ou t of c el l s , or of t i s s u e s o ri gi na l l y c el l ul a r, or of m a t e r i a l s
v
whi ch h a e i n th e cours e o f de e l op m ent b een de ri e d from v v ce l l s . It m u s t,
h owe v er, be b orn e i n m i nd th a t, a ccording t o S u n dry l e a di ng bi ol ogi s ts , ce l l s
i n t he s tri ct s en s e a re p rodu ct s e i t h e r of t h e p ri m i ti v e
not th e im m edi a t e
fi s si ons or of s ub se q ue nt fi s s i on s ; bu t th a t th e m ul t i pl yi ng s o-c a l l e d c el l s a re
nucl ea te d m as s es of p r oto pl as m whi c h re m a i n conn e c te d by s tra nds of p roto
p l as m a nd whi ch ac q ui re l i m i t i ng m emb r a ne s b y a s ec onda ry p r oce ss So .
t h at , In t he view of Mr . A da m Se dg wi ck a n d oth ers, th e s u bs ta n ce of an

orga ni s m i s i n fa c t a. cont i nu ou s m a ss o f v a c uol a ted protopl a s m .
22 M O RPH OLO G I CA L DE VELO PM ENT .
visibl e with ou t the mic ros c op e . B u t, i n so me cases, th ese

v eg et a l a gg reg at es of the fi rst or d er grow to a pp reci a bl e
cs
i
s z es .I n th e myc eli u m of s om e f u ngi we h a v e s i ngl e c e lls ,
d ev el op ed i nto l ong b ran ch ed fil a m ent s or ra m i fi ed t u bu l es , ,
th a t are of cons id era bl e l ength s A n an a l ogou s stru ctu re ch ar .
a cter i z es c ert a i n t r ib es of A l g ae of w hi c h C odi u m a dh wr ens , ,
Fig 4 m ay serv e a s a n e x a mpl e I n B otrydi u m a noth er

.
,
.
,
a lg a Fig 5 we h a v e a s tru ct u r e whi c h i s d e s c r ib e d a s s imu

,
.
,
l a ti ng a high er pl a nt with root s t e m bu d a nd f ru it al l

, , , , ,
p rod u ced by the b ra n chi ng of a s i ngl e c ell A n d a m ong .
f u n gi the genu s Mu cor Fig 6 fu rni sh es an ex a m pl e of ,

.
,
a lli e d ki n d *
H ere th ou gh th e s i ze a tt a i ne d is m u ch g reater
.
,
th an th a t of m any or g an i s m s w hi ch a re m or ph ol ogi c a lly

c om p ou n d we a re c om p ell e d to con s id er th e m or ph ol ogi c a l
,
com p os iti on a s s i m pl e ; s i n c e th e wh ol e c a n n o m ore be
s ep a r ated i nt o mi nor wh ol es th an c an the b ranch ed v ascu l a r ,
I n fu r th er i l l us t ra t i on, Mr Ta ns l ey
. n a m es th e fa c t t h a t in th e g enu s
Ca ul zrp a w e h a v e
’
e xt re m e l y com p l i ca te d form s o fte n o f co n s i d e ra bl e s i z e
p rod u ce d i n t he sa m e wa v . T he va r i o u s s nec i c s s i m u l a te v ery p e f

r ec tl y th e
m em b e rs o f d i ffe r ent g ro u p s a n o n 'r

'
t he high e r p l a nts , s uch as Il ors a ta il s ,
Mos sc s , Ca ctuses , Coni fe rs a nd th e l ike .

THE M O RPH OLO G I CA L C O MP OS I T I ON OF P LA NTS .
23
Sy s t em im a l I n th es e c as es we h a v e c on s id er a bl e
of a n an .
b u lk a tt a i ned not by a nu m b er of a ggre ga t es of th e fi rs t

,
or d er b e i ng u n it ed i nto a n a g g re g a t e of t h e secon d or d e r bu t ,
by the con ti nu ou s g rowth of a n a ggreg a t e of the fi rs t or der .
t r a ns iti on to high er for m s b egi n s i n a v ery

T he
§ 1 82 .
u nobtr u s iv e m a nn er A m on g th es e a ggr e g at es of th e fi rs t
.
o r d er a n a pp r oa ch t ow a r d s th a t u n i on by whi c h a gg r e g a t e s
,
of th e s econ d or d er a r e p r od u c ed i s i n dic a t e d by m er e j u xt a
,
p os iti on P rot ophyt es m u l t iply r a pidly ; a n d th ei r ra pi d

.
mu ltipli ca ti on s om eti m es cau ses crow di ng When i n st ead .

,
of fl oa ti ng f r ee i n th e w a t e r th ey f or m a thi n fi l m on a m oi s t
,
s u r f a ce or a r e imb edd e d i n a c om m on m a t r ix of m u cil a ge ;

,
the m ech an i ca l Ob sta cl es to di s p ers i on resu lt i n a ki n d of

f eebl e i nteg ra ti on v a gu ely s h a d owi n g forth a c om bi ne d
,
g rou p S om ewh a t m ore d efi ni te c om bi n ati on i s Sh ow n u s by

.
su c h pl a nts a s P a l m e l l a bo tr yoi d es H e re th e m em b er s of a .
f a m ily of c ell s a r i si n g by the S p onta n eou s fi ss i on of a p a r ent

,
c ell rem a i n u n i te d by S l end er th r ea d s of th a t j elly lik e sub

,
-
st a nce whi ch env el op s th ei r su r f a ces I n som e D i a tom a cece .
s e v er a l i n divid u a ls i n st ea d of c ompl et ely sep a r a ti n g h old

, ,
t ogeth er by th ei r a n gl es ; a nd i n oth er D ia tom a cew a s the ,
B a ci l l a ri a a v ar i a bl e nu m be r of u n i ts c oh ere so S ligh tly th at

, ,
they a re conti nu a lly m ovi ng i n rel a ti on to one anoth er .
T hi s form a ti on of a gg reg a t es of th e sec on d or d er fa i n tly ,
i n di cat e d i n feebl e a nd v a r i a bl e u ni ons lik e the a b ove m ay ,
be t ra ce d th rou gh ph a ses of i n c rea s i n g p er m a nenc e a n d de

fi ni ten ess a s w e ll a s i ncrea s i ng ex tent
, I n th e y ea st —pl a nt .
,
Fig 7 w e h a v e c ells which m a y e xist s i ngly or j oi ne d i nto

.
, ,
g r ou p s of sev e r a l ; a nd w hi ch h av e t h ei r sh a p es sca rcely at

a l l m odifi e d by th e i r c onne xi on A m on g the D es m i dia cece it
.
,
h a pp ens i n m a ny cases th at th e two i n divid u a l s p rod u ce d by

divi s i on of a p a r ent i ndivid u a l p a rt a s s oon a s th ey a r e fu lly
-
,
for m e d ; bu t i n oth e r c a se s i n st ea d of p a rti n g th ey com p ose

,
a g rou p of two A lli e d ki n d s Sh ow u s how by s u b seq u ent

.
,
fi s s i on s of th e a dh erent i ndi vid u a ls a nd th ei r p rogeny th ere ,

res u lt l onger grou p s ; a n d i n s om e S p eci es a conti nu ou s threa d ,
o f t h e m i s th u s p r o d u ce d Fig s 8 9 1 1 exhibit th es e
. .
, , ,
s ev era l s ta ges Fig 1 0 rep r esents a S cen edes m us i n whi ch

. .
the i ndivid u a ti on of the g r ou p is m an ifest I ns t ea d of li near .
a ggr e g a ti on m a n y p rot o phyt es ill u s tr a te centra l a gg re g a ti on ;

,
a s sh ow n i n Fig s 12 13 14 15 . Oth er i ns t a n c es a re fu r
, , ,
.
n i s h e d by su c h f or m s a s t h e Gom a m p ec tora l e Fig 1 6 ( a

’
,
.
b ei n g th e fron t vi ew a nd b the e dge Vi ew ) an d the S a rci na

, ,
v en tr i cu l i
,
Fig 1 7 F u rth er we h a v e th a t s ph er i ca l m od e
. .
,
of a ggr eg a ti on of whi c h th e Vol vox g l oba tor f u rn i s h es a
f a mili a r i nst a nc e .
T hu s far h owe v er th e i n divid u a lity of th e s econd a ry a g

, ,
g rega te i s feebly p r on ou n ced : n ot S i m ply i n th e s en s e th a t

it is s m a ll ; bu t a l so i n the sens e th at the i ndivid u a li ti es of
the p r i m a ry a ggreg a t es a re v e ry li ttl e s u b or di n a t ed B ut on .
s e eki n g fu r th er we fi n d t r a n s i ti on s t ow a r d s for m s i n which

,
the c om p ou n d i ndivid u a lity i s m ore d omi na nt whil e th e ,
s i m pl e i n divid u a liti es a r e m or e ob s cu r e d Ob s c u r a ti on .
o f on e ki n d accomp a n i es m e r e i n c r eas e of s i ze i n th e se c on d
a ry a gg re g a te I n p r op orti on to th e gr ea t er nu m b er of the
.
m orphologi ca l un it s h eld t og eth er i n one m a ss b ecom e s th ei r ,
r e l a t iv e i n s i n ifi can ce a s i n divid u a l s
g W e se e thi s i n t h.e
i rreg ul a r ly s p re a di ng li ch en s th at for m p atch es on r ock s ;

-
a nd i n s u c h cree pi n g f u n gi a s g row i n fil m s or l a m i n ae on
d ec ayi n g w ood a n d th e b a r k of t rees I n th ese c as es h ow .

,
e v er the i nt eg ra ti on of the c omp onent c ells is of a n a lm ost

,
THE M O RPH OLO G I CA L C O MP OS I T I ON OF P L A NTS .
25
m ech a ni ca l ki nd T he a gg reg at e of th em i s sca r c ely m ore

.
i ndividu ated th an a l u mp of i norg a nic m att er : a s witne ss the

w ay i n w hi c h th e li ch en e x ten d s i ts c u rv e d e dg es i n thi s or
th a t di recti on a s the s u r f ace f a v ou rs ; or the w a y i n w hi ch
,
th e fu ng u s g r ow s r oun d a nd imb ed s t h e s h oots a nd l ea v es

th a t l i e i n i t s w ay j us t a s so mu ch pl ast i c c l a y m ight do
, .
T h ou gh h ere i n the a u g me nta ti on of m a s s we see a p r ogress

, ,
tow a r d s th e e v ol u ti on of a high er typ e w e h a v e a s yet non e ,
of th a t de fi ni tene s s requ i re d t o c on s t i tu t e a c om p ou n d u nit ,
or t ru e a gg re g a t e of t he sec on d or d e r A n oth er ki n d .
o f ob scu r a ti on of th e m or ph ol ogi ca l u n i ts i s b r ou gh t a b ou t ,
by th ei r m ore c om pl ete coa l es cen ce i nto the for m of s om e

s t r u c t u r e m a d e by t h ei r u n i on T hi s i s we ll e x e m plifi e d
.
a m on g th e C on fcr voi d ece an d C onju ga tes I n Fig 1 8 th ere . .

,
a re r e p res ent e d th e st a ge s of a g rowi ng M ou g coti a genufl exa ,
i n wh i c h thi s m er gi n g of th e s i m pl e i n divid u a li t i es i nt o th e
c om p oun d i n divid u a lity i s S h own i n th e hi st or y of a s i n gl e
,
pl a nt ; an d i n Figs 1 9 2 0 .2 2 2 3 a r e r ep r e s en t e d a
, , , ,
*
s er i es of s p ec i es f r o m thi s gr ou p a n d th a t of C l a dop hor a
,
in ,
w hi c h w e see a p r ogress i n g i nteg ra ti on Whil e i n th e l ower .

,
typ es th e p r i m itiv e s ph eroid a l f or m s of th e cell s a re sca r cely

,
a l te r e d i n th e high er typ es th e c e ll s a re s o fu se d togeth er

,
a s to con s t i tu t e c yli n d e r s divid e d by s ep t a H e r e h ow e v er .

, ,
It m ay be j
ob e cte d th a t i n Cla dqvhor a pa rt m e nts of
th e s e p a ra t e com
th e t h a l l us se v
e ra l l y cont ai n m a ny nu cl ei , m a k i ng i t dou bt fu l wh eth e r t h e y
de s cen d from u n i-nu cl e a t e cel l s . v

If, how e e r, th ey do not th ey s i m pl y i l l us
t ra te a n oth e r form of i nte g ra t i on.
th e i ndefi ni teness i s till g reat T h ere a r e no s p ecifi c li m its

s .
to the l ength of a ny th rea d thu s p r od u c e d an d th e re i s n on e ,
of th a t di f fere nti ati on of p a rt s requ i red to giv e a d e cid e d i n

di vi du a l i ty to th e W h ol e .
T o c on stit u te som ethi ng lik e a t ru e a ggreg ate of th e s e c on d

or d er ,
c a p a bl e of s er vi ng a s a com p ou nd u n it th a t m a y be
c om bi ned with oth ers lik e it s el f i nto s till high er a ggreg a t es ,
th ere mu st e xi st b oth m a ss and defi ni teness .
§ 1 83 . An
pp r oa ch t oward s pl a nt s whi ch u n it e th es e
a
ch ara cter s m ay be t race d i n su ch form s a s B a ngi a ci l i a ri s

, ,
Fig 24 T h e mu ltiplic a ti on of c ell s h er e t ak es pl a ce n ot i n

. .
,
a l on gi tu di n a l di r ec ti on on ly bu t a l s o i n ,
a t r a n s v er se di r ec ti on ; a nd th e t ra n s v er s e
m u l tiplic a ti on b ei n g g rea t er tow ar d s the

middl e of the f ron d th ere res u l ts a differ
,
en ce b et w ee n th e m iddl e a n d th e tw o ex
t rem i ti e s— a c h a r act er whi ch i n a feebl e ,
w a y u n it es a l l the p a rt s i nt o a wh ol e
, .
E v en this s light i n divid u a ti on i s h owev er , ,
v ery i n d efin it ely m ark ed ; s i nce a s S h own ,
by the figur es the l a t era l m u ltiplic a ti on

,
of cell s d oes not g o on i n a p re c i s e m a nner .
F rom som e s u ch typ e a s thi s th ere ap

p ear to a ri se th rou gh slight di fferenc es i n
,
th e m od es of gr ow th tw o c l ose ly alli e d ,
-
g rou p s of pl ant s h avi ng i n divid u a liti e s

,
s om ewh a t m ore p r onou n ce d I f w hil e ,

.
the c ell s mu ltiply l ongi tu di n a lly th ei r l a t era l m u ltipli ca ti on

,
g oes on i n one di recti on on ly th ere resu lt s a fl a t s u r fa ce a s

, ,
i n the g enu s Ul va ( S ea l e ttu c e ) or i n the u pp e r p a rt of th e

-
th a ll u s of E n ter om orp ha L i m a Fig 2 5 ; or wh ere the l at era l

, .
m u l tipli ca ti on i s l ess u n i for m i n i ts ra t e i n typ es lik e ,
Fig 2 6
. . Bu t w h er e th e l a t er a l m u ltipli c a ti on oc cu r s i n two
di recti ons t r ans v ers e to one anoth er a h oll ow f ron d m ay be
,
p rod uc ed— s om etim es i rreg ul a rly s pheroid a l a nd s om etimes ,

27
i rregu l a rly t u b u l a r ; a s i n E n ter om orp ha i ntes ti n a l i s Fig 2 7 , .
A nd oft en as i n E n ter om orp ha com pr essa Fig 2 8 a nd oth er

, ,
.
,
sp eci es thi s t u bu l ar f ron d b ecom es b ranch ed Fig s 29 and

, . .
3 0 a r e m a gn ifi e d p orti on s of s u ch f ron d s ,
s h ow i n g th e
s i m pl e c e ll u l a r a gg reg a ti on w hi c h a lli e s th e m w ith the
p re
c edi ng f orms .
I n the c om m on F u ci of ou r c oa st s oth er a nd s om ewh a t

,
high er st a ges of this i nt eg ra ti on a re di s pl ay ed We h av e

.
f ron d s p rese rvi ng s om ethi n g lik e c onst an t b rea dth s a nd

dividi ng di ch ot om ou sly with a pp roxim at e re gu l a r ity T h ou gh .
th e s u bdivi si on s so p r od u ce d a re not to be reg a r d e d a s

s e p a r a t e fr on d s , bu t on ly a s ext ens i on s of one f ron d th ey ,
f oresh a d ow a high er d eg ree of c om p osi ti on ; a nd by th e com

p a r a t i v e l y m e th odic w a y i n whi c h th e y a r e u n it e d giv e t o ,
th e a gg rega t e a m ore d e fi n it e a s w e ll a s a m ore c om pl ex i n

, ,
d i vi du a l i ty. M any of the high er li ch ens exhibit an

a na l og ou s a dv a n ce . Whil e i n th e l ow est li ch en s the di fferent
,
p a rts of th e th all u s a re h eld t ogeth er on ly by b ei ng a l l

a tta ch e d to th e s u pp ort i n g s u r f ace i n th e high er li ch en s th e
,
t h a ll u s i s so fa r i nt eg r a te d t h a t it c a n s u pp or t i tself by
a tt a ch m e nt t o s u c h su r fa ce a t one p oi n t only . A nd th en i n ,
still m ore d ev el op ed ki nd s we find the th a ll u s a s su mi ng a

,
28 M O RPH OLO G I CA L DE VE LO PM E NT .
di chot om ou s ly b r anch e d form a n d so g a i ni ng a m ore speci fic

-
,
c h arac ter a s w e ll a s g r ea t er S i z e .
Wh er e a s i n typ es lik e th ese the m orph ol ogic a l un its

, ,
S h ow a n i nh erent t en d ency to a rr a n ge th e m s e lv es i n a m anne r
whi ch i s s o far c on sta nt as to giv e ch arac te r i s ti c p r op ort i on s ,
w e m ay s ay th a t th ere i s a r ecogn i z a bl e c om p ou n d i ndivid u a l

i ty . Con s id er i n g the T h all ophytes w hi c h g row i n thi s w a y
a p a rt fr om th ei r ki n ship s a n d w h olly with re fe r enc e t o th ei r
,
m orphol ogic a l c omp os it i on we might not i na ptly d es c rib e ,
them a s p seu do f oli ar -

.
§ 1 84 . A noth er
m od e i n which a gg r eg a ti on i s s o c a rri e d
on a s to p r od u ce a c omp ou n d i n divid u a li ty of c on s id er abl e
defi ni ten ess i s v a r i ou s ly di s pl ay ed a m on g oth er fa mili e s of

,
A l ga '
. Wh en th e c ell s i ns tea d of m u ltiplyi ng l ongi tu di nally
,
a l on e ,
a n d i n st e a d of a l l m u l tiplyi n g l a te r a lly as w e ll a s
l ongi tu di na lly m u ltiply l at er ally on l y a t p a rti cu l ar pl aces

, ,
th ey p r od u ce b ranch ed s t ru ctu res .
I ndic a ti on s Of thi s m od e of a gg reg at i on occ u r a m ong th e

C onfervoi deae a s sh own i n Fig s 22 2 3
,
T h ou gh i n s om e of .
,
.
,
th e m ore d ev el op e d A l g az whi ch e xhibit th e ra m i fied a rra nge

m ent i n a high er d egr ee the c om p on en t cells a re lik e th os e
, ,
of th e l owe r A l g er u n i te d t og eth er en d t o e n d i n s u c h w a y
, ,
a s bu t littl e t o ob s c u r e th ei r sep a r a t e for m s a s i n C l a dop hom ,
H u tc hi nsi w Fig 3 1 ; th ey ne v erth el es s e vi n ce gr ea t er s ub

,
.
or di n a ti on to th e w h ol e of which th e y ar e p a rt s by a rra n gi ng ,
th em selves m or e m eth odi c a lly S till fu rt h er p ronou nce d .
b ecom es th e c omp ou n d i ndivid u a li ty wh en whil e the com ,
p onent cell s of th e b ra n ch es un it e com pl etely i nt o j oi nte d

c yli n d er s th e c omp onen t c ells of th e st e m for m a n a xis d i s
,
ti ngu i sh ed by i ts rel ativ e thi c kn ess a nd c ompl exity S u ch .
typ es of st ru ctu res a re i ndi cat ed by Figs 3 2 3 3 — fl g u res .

,
rep r esen ti ng s m a ll p orti ons of pl a nt s which a r e q u it e t re e
lik e i n th ei r enti re ou tlines On ex a mi ni n g Figs 3 4 3 5 3 6 . .

, , ,
whi ch S h ow the st ru ct u res of th e st ems i n th es e typ e s i t ,
will be s een too th at the c omponent c ells i n b ecomi ng m ore

, ,
T HE M O RPH OLO G I CA L C O MP OS I T I ON OF P L A NTS .
29
coh erent h av e u nd ergone ch a n ges of f or m whi c h ob scu re

,
thei r i ndividu a liti es m ore th an b efore N ot only are th ey

.
mu ch el onga t ed but th ey a re s o comp ress e d a s to be p ris

,
m atic rather th an cylin d rica l . T his st ru ctu re b es id e s di s

,
pl ayi ng i nt eg r a ti on of the m orph ol ogi ca l u n it s ca rr i e d on i n

two di r ecti on s i n s t ea d of one ; a nd b es id es di s pl a yi ng thi s
high er i nt eg ra ti on i n the g rea t er m erging of the i n divid u ali
ti es of the m or ph ol ogic a l u nits i n the general individ u a li ty ;
a ls o di s pl a y s it i n th e m ore p ronou n c e d s u bor di n a ti on of th e
b r anch es and b ranchl et s to the m a i n s t em .T hi s di ffer en ti a

ti on and cons olid a ti on of the st em b ri ngs al l the secon d ary
,
gr owth s i nt o m ore m ar k ed d ep end ence ; and s o r end er s the

i ndividu ality of the a ggr eg at e m ore d ecid ed .
We might not i n a pp r op ri a t ely c a ll this typ e of st ru ctu r e

p seu d a xi a l
-
. I t si m u l at es th at of the high er pl a nt s i n c er
t a i n su p er fi c i a l ch a ra ct er s
. We s ee i n it a p rim a ry a xis al ong
wh i ch d ev el op m en t m ay con ti nu e i nd efi n it ely and from
,
whi c h th er e bu d ou t l a t era lly s econ d a r y a x e s of lik e n a tu re

, , ,
b e ari ng lik e terti ary a x es ; a nd thi s i s a m od e of g rowth

with whi ch P h aenog a m s m a k e u s f a mili ar .
§ 1 8 5 S om e of the l arg er A l gae su pply ex a mpl es of an

.
i nt egr a ti on still m or e a dv a nced ; n ot si m ply i na smu ch a s

th ey u ni te m u ch g reater nu mb er s of m or ph ol ogi ca l u n its
30 M O RPH OLO G I CA L DE V ELO PM ENT .
i nto con ti nu ou s m a sses bu t a l s o i n a smu ch a s th ey c ombi ne

,
the p s eu d o foli ar s t ru ct u r e w i th the p seu d a xi a l st ru ct u r e

- -
.
Ou r own sh ores f u rni s h a n i nst a nce of t his i n the c om m on

’
L a m i na m a ; an d c ert a i n gig a ntic L a m i na ri a ccaz of th e A nt

a r ctic s e a s fu m i s h yet b e t ter i n s t a n ces
,
I n N ecrocys ti s th e .
ge rm d ev el op s a v ery l ong sl en d er st em whi ch ev entu a lly ,
exp an d s i n to a l arg e bl a dd er lik e or c yli n d ri c a l a i r v ess el ;

- -
a n d th e s u r fa ce of thi s b e a r s nu m e r ou s l e a f s h a p e d ex a n
p
-
s i ons . A n oth er ki n d L ess oni a fu s ces cens Fig 3 7 sh ow s u s a

, ,
.
,
m a ss ive s t em g row i ng u p th r ou gh w ater

m a ny feet d ee p— a st e m whi c h bi fu rca ti n g ,
a s i t a pp roa c h es th e s u r face fl a tten s ou t th e ,
e nd s of i ts s u bdivi s i on s i nto fron d s like

r ib a n d s T h es e h owe v er a re n ot t ru e f oli a r
.
, ,
a pp en d a g e s S i n ce th ey a r e m ere ly e x
,
p a nd e d
c onti nu ati on s of the s t em I n E gr egi a .
b r a nch es of the th all u s not only t a k e the

f orm of l eav es bu t th ese a re diffe renti at ed
,
i nt o sev era l c a t eg or i es i n a ccor d ance with a

divi s i on of l a b ou r I n any of th ese L a m i n .
a r i a c cce th e w h ol e pl an t g r ea t a s m a y be i ts ,
S i z e a n d m a d e u p th ou gh it s e e m s to be of
,
m any grou p s of m orph ol ogi c a l u n it s u n i te d ,
i nt o a com p ou n d g r ou p by th ei r m a rk ed s u b ordi na ti on to a
c onnecti ng m a s s i s n ev erth el ess a si ngl e th all u s w hi ch i s
, ,
“
a dd ed to by i nte rc a l a r y g row th at th e t ra ns iti on pl a ce at -
,
th e j u ncti on of th e s tem lik e a n d l ea f lik e p orti on s

-
T he -
.
a gg re g a t e i s s till a n a gg r e ga t e of th e s ec on d ord e r .
B u t a m ong c ert a i n of th e high es t A l gae w e do fi nd s om e ,
thi n g m or e th an thi s u n i on of th e p s eu d a xi a l w i th the -
p s eu do f oli ar stru ct u re I n a dditi on to p seu d a x es of c om

-
.
-
p a r a t i ve c om pl e xi ty ; a n d i n a dditi on to p seu d o f oli a th a t -
a r e lik e l e a v e s n ot only i n th e i r g ener a l sh a p es bu t i n h av

,
i n g m i d rib s a nd e v en v e i n s ; th e re a re the b e gi nni ng s of

-
a high er s t a g e of i nteg ra ti on Fig s 3 8 3 9 an d 4 0 sh ow

. .
, , ,
s om e of the st ep s I n Rhodym eni a p a l m a ta Fig 3 8 th e

.
,
.
,
T HE M O RPH OLO G I CA L C O MP OSIT I ON OF P L A NTS .
31
p arent -
fron d i s c om p a ra tiv ely i rre gu l ar i n for m a nd with ou t
,
a m i d rib
-
a nd a l on g with t hi s v e ry i m p er f e c t i n te g ra ti on ,
we see th at the s ec ond ary fron d s g rowi n g f rom th e edge s are

di st r ib u t e d v ery m u ch a t r a n d om a n d a re by n o m ean s
,
s p ec ifi c i n th e i r s h a p e s A c ons id era bl e a dv a n c e i s di s pl ay e d
.
by P hyl l op hora r u bens Fig 3 9 , . H ere the f ron d p r im a r y

.
, ,
s econ d a ry or te rti a ry b et ra ys s om e a pp roa ch tow ar d s r egu

, ,
l a ri ty i n both f or m a n d s i ze ; by whi c h as a l s o by its ,
p a rti ally d ev el op ed mid ri b th ere i s est a bli s h ed a m ore

- -
,
m arked i ndivid u a lity ; an d at the s a m e tim e the g rowt h of ,
th e s e c on d a r y fron d s no l onger oc c u r s anyw h ere on th e e dge ,
i n th e sa m e pl ane a s th e p a r ent fron d bu t f rom th e su rface

-
,
a t s p ec ific pl aces D el ess er i a s a n gu i n ea Fig 4 0 ill u str a t es a

.
,
.
,
mu ch m ore d efi n it e a rr angem en t of th e s a m e ki n d T he .
f rond s of thi s pl a nt qu it e r e gu l a rly S h a p ed h a v e th ei r p art s

, ,
d ecid edly su bor di na te d to th e w h ol e ; an d from th ei r mid

r ib s g row oth er fron d s w hi c h a r e j u s t lik e th e m E ac h of .
9
th ese fron d s i s a n organ i zed g rou p of th ose m orphol ogi ca l
u nit s whi c h w e di s ti ngu i s h a s a gg r e g a t es o f th e fi r s t or d er .
A nd i n this c ase two or m ore su ch a ggre ga t es of th e s econ d

,
32 M O RPH OLO G I CA L DE VE LO PM ENT .
or d er w ell i ndivid u ate d by th ei r forms and stru ctu res a re

, ,
u n i te d t og eth er ; a nd th e pl a n t c om p ose d of t h e m i s t h u s
ren d ere d i n s o fa r a n a gg re g a t e of t h e t hi r d or d er
, ,
.
J u st noti ng th a t i n c ert a i n of th e m ost d ev el op ed A l gae a s -

,
th e S a rgas s u m or comm on gu l f w ee d thi s t erti a ry d eg ree of

,
-
,
c omp os i ti on is fa r m or e c om pl ete ly di s pl a y e d s o a s to p ro ,
du c e a m ong T h allophyt e s a typ e of st ru ctu r e cl osely si m u l at

i ng th at of th e high er pl a nts l et u s now p a ss to th e c ons id era
,
ti on of th ese high er pl ant s .
1 86 . H a vi ng the
f c th e s oil for a su pp ort and th e
s u r a e of
a i r for a m edi u m te rres t r i a l pl a nt s a re m ech anic a lly ci rcu m

,
sta nced i n a m a nner w id e ly diff er en t from th a t i n w hich
a qu a tic pl a nt s a r e c i rcu m s t a nc e d I n s t ea d of b ei n g b u oy ed
.
u p by a s u rr ou n di ng fl u id of s p ec ifi c g r a vity eq u a l t o th e i r
own ,
th ey h a v e to erect th em selv es i nt o a ra re fl u id whi ch
yi eld s no a pp reci a bl e s u pp ort Fu rth er t h ey a re di s
.
,
S i m i l a rl j c on di t i on e d i n h a vi n g two s ou r c es of nu t r i m en t i n
r
pl a c e of one Un lik e the A l gae w hich d eriv e a l l th e m ate

’
,
.
r i a l s for t h e i r t i ss u es from th e w at er b a thi n g th ei r enti r e
su r f a c e s a n d u s e th ei r r oot s only for a tt a ch m ent m ost of th e

, ,
’
pl ants which c ov er the E a rth s su rfa c e a b sorb p art of th ei r ,
food th rou gh th ei r i m b edd ed r oots a n d p ar t th rou gh th ei r

e xp ose d l ea v es T h es e two m ark ed u nlik enes s e s i n the re l a
.
ti ons to su rrou ndi ng con di ti on s p rofou ndly affect the res pec ,
tiv e m od es of g row th We m u s t d u ly b ear th em i n mi nd

.
whil e st u dyi ng the fu rth er a dv a nc e of comp os iti on .
T h e c l as s of pl a n t s to w hich w e now t u rn— th a t of the

A r ch eg om a tw— is nea r ly rel ate d by i ts l ower m embe rs to th e
’
cl a sses a b ov e d ea lt w i th SO m u ch so th a t som e of the i nferi or ,
liverw ort s a re qu it e li ch eni form a nd a r e oft en m i st a k en for ,
li ch en s P a ss i ng ov er th es e l et u s rec om m enc e ou r sy nth es is

.
,
with s u ch m em b ers of the c l a s s a s r ep ea t th os e i ndic ati ons of

p rogres s t ow ar ds a high er com p os i ti on whi ch we h a v e j u s t Oh ,
s erv e d a m on g th e m ore d ev el op e d A l gae Th e J u ng er m a n

-
.
n ia c ew f u rn ish u s wi th a s er i es of typ es cl ea rly i n di c a ti n g the ,

M O RPH OLO G I CA L C O MP OS I T I ON P LAN TS
“
T HE OF .
33
t ran s iti on f rom an a ggr egat e of th e s econ d or d er to an a ggre

g a te of the thi r d or d er Fig s 4 1 a n d 4 2 i n dic a te th e s tr u e
. .
, ,
t u re a m ong the l owe st of thi s grou p H ere th ere is bu t a n i n .
com pl et e d e v e l opm ent of th e s e c on d or d e r of a gg re g a t e T he .
f ron d grows a s i rregu l ar ly a s the th all u s of a lich en : it i s i n

d e fi ni te i n Si ze a nd ou tli ne s p rea di n g hith er or t hithe r a s ,
the con diti on s f a v ou r Moreov er it l a ck s th e di fi erenti ati ons

.
,
r e qu i r e d t o s u b or di n a t e i ts p a rt s t o th e w h ol e : it i s u n i form ly
c ell u l ar h avi ng neither m i d r i b nor v e i ns ; a nd i t p u ts ou t

,
-
r ootl et s i n di ffe r ently from a l l p a rt s of i ts u n d er s u r fac e In -

.
Fi g 4 3 P el l i a ep i p hyl l a we h a v e a n a dv an ce on thi s typ e

.
, , .
T h er e i s h ere as s h own i n the t r a ns v er se secti on Fig 4 4 a

, ,
.
,
th i ck en i n g of the frond a l ong i ts c ent ra l p or ti on p rod u c i ng ,
s om ethi ng lik e an a pp roac h t owar ds a mid —ri b ; and from

thi s the rootl ets a re chi efly giv en off T he ou tli n e too is .
, ,
mu ch l es s i rre gu l ar ; wh enc e resu lt s g rea t er di sti nct ness of

A f u rth er s t ep i s di s pl a y e d i n M etzg em a
‘
th e i ndivid u a li ty .
f u r ca ta Fig
,
4 5 . T h e f r on.d o f t hi s pl a n t co mp a r a tiv ely w ell ,
i nt egr at ed by th e di st rib u ti on of i ts su b s t a nce a r ou n d a ‘
d ecid ed mid ri b and by i ts comp a ra tiv ely defi ni te ou tli n es

-
,
-
,
p rod u ces s econ d ary f rond s T h er e i s w h a t i s ca ll ed p rol i fer .
ou s g r owt h ; a n d occa s i on a lly a s s h ow n i n Fig 4 6 re p resen t ,

.
,
i ng an e n l a r ge d p orti on th e g rowth i s d ou bly p r oli fer ou s

,
In -
.
th ese c a ses h owev er th e t e rt i a r y a gg reg a te S O fa r a s it i s

, , ,
for m e d i s bu t v ery f eebly i n t eg ra t ed ; a n d i t s i nt eg r a ti on i s

,
bu t t em p or a r y F or not only do th es e y ou nger fron d s t h a t

.
bu d ou t from th e m i d r ib s Of old er fron d s d ev el op r ootl et s of

-
,
th ei r own ; bu t a s soon a s th ey a r e well gr own a n d a d equ a t ely

r oot ed th e y dis solv e th e i r c onnexi on s with the p arent f ronds
,
-
,
49
34 M O RPH OLO G I CA L DE VELO PM ENT .
a nd b ecom e qu i te i n d ep end ent . F rom th ese t rans i

ti ona l for m s we p a ss i n th e high er ,
J u ng er m a n ni a ccce, to
f orms comp os ed of m any f ron d s th at are p er m a nently un it ed

by a conti nu ou s st em A m ore d ev el op ed a gg reg a t e of the
.
-
thi r d or d er is th u s p rod u ced Bu t t h ou gh a l ong wi th i n

.
,
c rease d defin i ten ess i n the s econd a ry a gg rega t e s th er e i s h ere ,
a n i n t eg r a ti on of th e m so ex ten s iv e a n d s o regu l ar th a t th ey ,
a r e vi s ibly s u b or di n a t e d to th e w h ol e th ey f or m ; yet th e
s u b or di n at i on i s r ea lly v ery i n c om pl et e I n s om e i n st a nces .

,
as i n Ra du l a com p l a na ta Fig 4 7 th e l ea fl ets d ev el op root s

,
.
,
f rom th e i r u nd er s u r f a c es j u s t a s the p ri m i t iv e fron d d oes ;

,
a nd i n th e m a j or ity of th e g r ou p a s i n J ca pi ta ta ,
Fig 4 8 .
,
.
,
root s a r e giv en Of f a l l a l on g the conne cti n g s t em a t the s p ot s ,
wh ere th e l ea fl et s or frondl ets j oi n it : the resu lt b ei n g th a t

th ou gh th e connect e d fron dl ets for m a phy s i ca l wh ol e th ey ,
do n ot for m i n a ny d ec id e d m an ner a phy s i ol ogic a l wh ol e ;

, ,
si nce su c ce s s iv e porti ons of the u n it ed s eri es carry on th ei r ,
fu nc ti on s i n d e p end ently of the r es t Fi na lly the .

,
m ost d ev el op ed m emb ers of th e grou p wh eth er li nea lly de ,
s cend ed from the l es s d ev el op ed or from an ea rly typ e com

m on to th e two p r esen t u s w i th t e rti a ry a gg reg at es whi ch a re
,
phy si ol ogi c ally as w ell a s phy s i c a lly i ntegra ted *

N ot lyi n g .
Th e gr ea t m a s s of ea rl y ance st ra l t yp e s — pl a nt a nd a ni m a l — cons i s ti ng o f
s oft t i s s u e s , h a v e l e ft no r e m a i ns wh a te v e r, a nd we ha ve no rea s on t o s u pp o s e
th a t thos e whi ch l e ft rem a i ns fe l l wi th i n the di re ct a nce s tra l l i ne s of a ny
e xi sti ng form s Contra ri wi se w e h a ve rea s on t o s uppos e th a t th ey fe ll wi th

. ,
i n l i nes of e v ol u ti on ou t o f whi ch t he l i nes endi ng i n e x i s ti ng form s di v erged .
We m us t th erefore i nfer that the di ffi cul ti e s of a f

fi l i a t i on whi ch a ri s e i f we
35
rone lik e th e ki n d s th u s far d es c r ib e d bu t g rowi n g e rect

p , ,
t he s t e m a nd a tt a ch e d l ea fl et s b ec om e d ep end en t u p on a
s i ngl e root or g rou p of root s ; a n d b ei ng so p re v e nt e d f r o m
'
c arryi ng on th ei r f un cti ons s ep a ra t ely are m a de memb ers of ,
a com p ou n d i n divid u a l : th er e a r is es a d efi n i tel y est a bli s h e d -
a ggr e g a t e of th e thi r d d e g re e of c omp ositi on .
T he fa ct s a s a rra nge d i n the a b ov e ord er a re s u ggestiv e .
M i nu t e aggreg at es or c ells the g rou pi ng of wh i c h w e t r a c ed

, ,
i n § 1 8 2 s h owe d u s a na l ogou s ph as es of i n d efi nit e u n i on

, ,
whi ch a pp ea re d to l ea d th e way t owar d s d efinit e u ni on We .
s ee h e re a m on g c om p ou n d a gg reg a tes a s we s aw there ,
a m on g s impl e a gg r e g a t e s th e e st abli s hm en t of a sp ecific,
f or m a nd a S i ze th a t f a ll s withi n m od era t e limi ts of v ar i a

,
ti on T hi s p a s s a ge f rom l ess d e fi nit e ext ens i on to m ore

.
d efi nit e ext ens i on s eem s i n th e on e c a s e a s the oth er to be

, , ,
a cc om p a n i e d by th e r esu l t th a t g r owth ex c ee di n g a c ert a i n ,
r a t e end s i n th e f or m a ti on of a n ew a gg reg a t e r a th er th a n
, ,
a n e nl a r g em ent o f th e ol d A nd on the high er S t a ge as on .

,
th e l ow er thi s p roc ess i rregu l a r ly c a rr i ed ou t i n th e S impl er

, ,
types p rodu c es i n th em u ni on s th a t a r e bu t t em p or a ry ; w hil e

,
i n th e m ore d e v e l op e d typ es it p r oc ee d s i n a s y s t em a tic w ay

-
, ,
a n d end s i n th e p r od u c ti on of a p er m an en t a ggr eg a t e th a t is
d ou bly comp ou n d .
conte m v
p l a te di e rg ent typ es now e xi sti ng, w oul d not a ri s e i f w e h ad before
us a l l t he ea rl y i n te rm edi a t e t ype s Th e Ma m m a l i a differ i n s undry re s p ec ts
—Fi h
.
from a ll oth er k i n ds of Vert eb ra t e s es, Rep t il e s, Bi rds ; a nd if t h e

a bs en ce of h a i r, m a m m ae, a nd t w o o cci p i ta l condyl es , i n t h e s e oth e r v ert e
b ra te s we re ta k en t o i m pl y a funda m e nt al di s ti nc ti on, i t m igh t, i n th e a b s ence
of a ny k no wn fos s il l i nk s , b e i nfer re d th a t th e Mam m a li a bel on ge d t o a sep a
ra te phyl u m . But t h ese differen c e s a re not h el d to n e ga t i e v th e a s s um e d
rel a t i ons hi p . Si m il a rl y g pl ant s

a m on We m u s t not re e ct a n hyp othesi s
. j
re s p ect i ng a cert a i n s u p p os e d t yp e b ec au s e th e e x i s ti n t e s i t m ust ha ve
, g yp
b een a ki n t o p re s ent tra i t s whi ch i t co ul d not ha v e h a d . We a re j u st i fied in
a s s um i ng, w ith i n l i m i t s , hyp oth eti ca l type , unl ik e
a e xi s ting type s i n t ra i ts of
s om e i m porta n ce . Hence re s ul t s t h e a n s w e r to a c ri t i ci s m p a s s ed on th e
v
a b o e a rg um ent , th a t i t i m p l i e s rel a ti on s b e t wee n th e u nd e vel oped a nd dev el
for m s of th e J u ng er m
‘
O pe d a nm a ceae s uch as t h e fa cts do not S h ow u s . Thi s

j
ob ec t i on is m e t o n r em em b eri ng t h a t th e type s i n whi ch the suppose d tra ns i
ti on took pl a ce di s a ppea re d m yr i a ds of yea rs a go .
36 M O RPH OLO G I CA L D E VE LO PM ENT .
Mus t we th en conc lu d e th a t a s c ells or m orph ol og ic a l

,
u ni t s a r e i n t e g ra t e d i nt o a u ni t of a high er or d er whi c h w e
, ,
c a ll a th all u s or fron d ; so by th e i nt egra ti on of frond s th ere

, ,
i s ev olv e d a s t ru ct u r e s u c h a s th e a b ov e d eli ne at e d s p eci e s

-
p os sess ? Wh eth er this is th e i nt erp retati on to be giv en of

th ese pl an t s w e sh all b es t see wh en c onsid eri ng wh eth er it i s
,
th e i n ter p ret a ti on to be giv en of pl an t s w hich ra nk a bov e

th em . T hu s fa r we h a v e d ea lt only w i th the C r ypt oga m i a .
We h a v e now to d ea l wi th the P h an eroga mi a or P haenog a m i a .

CH A P T E R I I I .
TH E M O RP H OLO GI C A L C O M P O S I T I ON OF P L A N T S,
C ON TI N UE D .
1 87 . T H AT
dv a nc ed comp os iti on a rriv e d at i n the
a
A r ch egonia taz is c a r ri ed still fu rth er i n the Fl oweri ng P l ant s

, .
I n th ese m os t el ev a t ed v eg et a l forms a ggreg ati on of the thi r d

-
,
ord er is a lw a y s di s ti nc tly di s pl ay e d ; a nd a gg r e g a t e s of the
fou rth fifth s ixth &c or d ers are v ery c omm on

, , ,
.
, .
Ou r i nqu i ry i nt o the m orph ol ogy of th es e fl owering pl ants ,
m ay be a dv a nt a g eou s ly c omm enc e d by st u dying the d e v el op
m ent of Simpl e l ea v es int o c omp ou nd l ea v es I t is ea sy to .
t ra c e the t ra ns iti on a s w ell a s the c onditi ons und er whi c h it

,
occ u rs ; a n d t ra c i ng it w ill p rep are u s for u nd e rs t andi ng
how a n d wh en m et a m or ph oses still g rea t er i n d egree t a k e

, ,
pl a ce .
§ 1 88 . If
x mi ne a b ran ch of the c omm on b rambl e
we e a ,
wh en i n fl ower or a ft erwa rds we sh a ll not u nfrequ ently fi nd

,
a si m pl e or u ndivid e d l e a f a t the i ns erti on of one of th e

,
l a t er a l fl ower b ea ri ng a x es c omp osi ng the t ermi nal cl u s ter

-
,
of fl ow er s S om etim es this l eaf i s p ar ti a lly l ob e d ; s om et i m es

.
c l eft i nt o th ree s m a ll l ea fl ets L ow er d own on the s h oot if

.
,
it be a l a t era l one occu r l a rger l eav es c omp osed of th ree

, ,
l ea fl et s ; a nd in s om e of th es e two of the l ea fl ets m ay be ,
l ob ed m ore or l ess d eeply On the m ai n st em the l ea v es

.
,
u su a lly still l ar ger will be f oun d to h a v e fi ve l ea fl ets

,
S up .
p osi ng the pl ant to be a w ell g rown one, it will f urnish al l -
37 r
38 M O RPH OLO G I CA L DE VE LO PM E NT .
g ra d ati ons b etween th e s impl e v ery s m a ll l e a f and the l arge

, ,
c omp os it e l ea f,
c ont a i ni ng s om eti m es ev en sev en l ea fl ets .
Figs 5 0 to 6 4 rep res ent l ea di n g sta ges of th e tra n si ti on

.
,
.
Wh a t d et ermi ne s thi s t ransiti on ? Ob s erv ati on sh ows th a t

th e qu i nt u pl e l ea v es occu r wh ere the m ater i a l s for g rowth
a re s u ppli e d i n g reat es t ab und anc e ; th at the l eave s b ec om e
39
l es s a nd l ess c om p ou n d i n p rop orti on to th ei r rem otene s s

,
f rom the m a i n c u rren t s of s a p ; an d th at w h ere a n enti r e

a b s en c e of divi s i ons or l ob e s i s ob s er v e d i t i s on l ea v e s withi n
,
th e fl ower b u nch : at the pl a c e th a t i s wh er e the forces whi ch

-
, ,
c a u s e g row th a r e nea rly e qu ilib r a te d by the f orc e s w hi c h
opp os e g r owth ; a nd wh e re a s a c on s eq u enc e g a m og en esis i s

, ,
a b ou t to bese t i n A ddi ti on a l e vid ence th a t th e d eg ree

of nu t r iti on d e termi nes th e d eg ree of co mp os i t i on of the l ea f ,
i s f u rni sh e d by th e rel a tiv e s i zes of the l eav es N ot only on .

,
the a ver a ge is th e qu i n t u pl e l ea f mu c h l a rger i n its t ot a l a rea

,
th a n th e t ripl e l ea f ; bu t the com p onent l ea fl ets of the one ,
a r e u su a lly m u c h l a rg er t h a n th os e of th e oth er The lik e .
c ont ra sts a re s till m ore m ar k e d b etw een t ripl e l eaves a nd

s impl e l e a v es . T hi s c onn ec ti on of d ec re a s i ng s i ze w ith de
c reas i ng com p os iti on i s c ons pi cu ou s i n the ser i es of figu res
,
the di ff eren ces S h own b e i ng not nea rly s o gr e a t as m ay be

f requ ently ob ser v ed Con fi rm ati on m ay be d r awn f rom th e
.
f a ct th a t wh en th e l ea di n g s hoot i s b rok en or a rrest e d i n i ts

g rowth the s hoots it giv es off ( p rovid ed th ey a re given Off
,
a ft er th e i n j u ry ) a n d i nt o which its ch eck e d c u r r ent s of sa p

,
a r e th r own ,
p rod u ce l eaves of fi ve l ea fl et s wh ere ordi nar ily
l ea v e s of th ree l ea fl et s o ccu r Of cou r se i nc id ent a l ci rcu m
.
s t a nc es a s v a r i a ti on s i n th e a m ou nts of s u nshi ne or of r a i n
, , ,
or o f m a tt e r s u ppli e d to th e r oots a r e e v er p r od u c i ng c h a ng es
,
i n the s t a t e of the pl a nt a s a wh ol e ; a nd by th u s a ff ecti ng

the nu t r i ti on of i ts l ea f b u ds a t th e ti m es of th ei r f or m ati on
-
,
c au se i rre gu l ar iti es i n the rel a ti ons of s i ze an d c omp ositi on

a b ov e d es c r ib e d . B u t t a ki ng th ese c au s es i nto a cc ou nt it is ,
a b u n d a n tly m a n if es t th a t a l ea f bu d o f th e b ra mbl e w ill

-
d ev el op i nto a Si m pl e l ea f or i nto a l ea f c om p ou nd ed i n
di fferent d e grees a ccordi ng to th e q u anti ty of a s s imil a bl e
,
m a tt er b rou ght to it a t the tim e wh en the ru dim ents of i ts

s t ru ct u r e ar e b ei n g fix e d A nd on s tu dyi n g th e h a bits Of
.
oth er pl a nt s— o u obs er vi n g h ow a nnu a ls th a t h a v e c om p ou n d
l ea v es u su a lly b ea r si m pl e l ea v es at the ou t s et , wh en the

as simil ati ng su r f a ce i s bu t sm all and how wh en c omp ou nd ,
40 M O RPH OLO G I CA L DE V E LO PM EN T .
l ea ved pl ant s i n f u ll growt h bear s i m pl e l ea v es i n the mid s t

o f c o m p ou n d on es the rel a tiv e s m a ll n es s of s u ch simpl e
,
l ea v es Sh ow s th a t the b u d s f rom which they a ros e were ill

s u ppli e d wi th sa p ; it will cea s e to be d ou bt ed th a t a foli ar
o r gan m ay be m eta m orph ose d i nt o a g rou p of foli a r or g a n s ,
if fu rn i s h ed a t the right t im e wi th a qu a nti ty of m a tt er

, ,
g reat er th a n ca n be rea dily org ani zed rou n d a s i ngl e centre of

g rowth A n e x a m i na ti on of the t r a ns i ti ons th rou gh w hi ch a
.
c om p ou n d l ea f p asses i n t o a d ou bly —c om p ou n d l ea f a s s e en ,
i n th e v a r i ou s i nter m edi a te form s of l ea fl et s i n Fig 6 5 , will .
f u rth er en forc e this concl u s i on .
H ere we m ay a dv ant a geou sly not e, too, how s u ch c as es

the eal f st a
- lk u n d e r g o e s c o nc omit a n t ch a n g e s o f st r u ct u r e
.
I n th e b ram bl e l ea v es ab ov e d esc rib ed it b ec om es c omp ound

-
,
s im lt n
u a eou s ly with t h e l e a f — th e v e in s b e c o m e mid -
r ibs whi l e
ib
th e m i d r s - b ec om e p e ti o l e s
. M ore o v er t,
h e s e c on d a r y st a lk s
,
a nd still m ore the m ain st alks , b ear th orns S imil ar i n th ei

r
41
sh ap es an d a pp roachi ng i n th ei r si zes to thos e on the s tem

, ,
b es ides S imu l ati ng the s t em i n c ol ou r a nd te xtu re I n the .
p eti ol es of l a rge c omp ou n d l ea v es lik e t hos e of the c om ,
m on H era c l cu m we see s till m ore di s ti nctly b oth i nt er na l

,
a nd ext ern a l a pp r oxi m a ti on s i n c h a ra ct er to a x e s N or a r e .
ther e wanti ng pl ants wh ose l arge th ou gh simpl e l ea v es a re , , ,
held ou t fa r f rom the st ems by foot st a lk s th a t a re n ea r the -

,
en d s, s om etim e s s o lik e a x es th a t th e t ra ns v er s e s ecti on s of

'
the two are i ndi sti ngu ish a bl e as i nst a n c e the Ca l l a pa l a s tr ts .
One oth er f a ct resp ect i ng th e m odifi c a ti ons whi c h l ea v e s

u nd erg o s h ou ld be s et d ow n
,
N ot only m a y l e a f s t a lk s
.
-
a ssu m e t o a g rea t d eg ree th e c h a r a ct er s of st e m s wh en th e y ,
h a v e to di s ch arge the fu ncti ons of st em s by s u pp or ti ng m any ,
l ea ves or v ery l a rg e l ea v es ; bu t they m ay a s su m e th e ch a r ac

te rs of l ea v es wh en they h av e to u ndert ak e the f u ncti on s
,
of l ea v e s T h e A u s t ra li an A c a c i a s f u r ni s h a r em a r k a bl e
.
illu st r ati on of thi s A ca c i a s e l sew h ere f ou nd bear pi nnat e

.
l eav es ; bu t the m a j or ity of th ose f ou nd i n A u st r ali a hear

wh at a pp ea r to be s impl e l ea ves I t t u rns out h owev er th a t .
, ,
th ese a r e m ere ly l ea f s t a lks fl a ttened ou t i nt o foli ar sh ap es

-
th e l a m i nm of th e l ea v es b ei n g un d ev el op ed A n d th e p roof .
i s th a t i n y ou n g pl a nt s sh ow i ng th ei r ki n ships by th e i r
,
e m b ry on i c c h a r a c t er s th es e l e a f—lik e p eti ol e s b ea r t ru e l ea fl ets

,
a t th ei r en d s A m et a morph os is of lik e ki n d occu r s i n Oxa l is

.
bup l eu r i fol i a Fig 6 6 ,

T h e f a ct m os t d eservi ng of notic e
. .
,
h ow ev er i s th a t th es e l ea f
,
st a lks i n u su r pi ng the g en
,
e ra l a sp ect s a n d f u n cti on s
of l ea f bl a d e s h a v e to som e
-
, ,
e xt ent a ls o u s u r p e d
, th ei r
s t ru ct u r es : th ou gh th ei r v en a ti on i s n ot lik e th a t of th e l ea f
bl a d e s th ey repl ac e yet they h ave v ei ns and i n s om e ca ses

, ,
mid r ibs
-
.
Red u c ed to th ei r m ost g ener a l exp ress i on th e t ru ths a b ov e ,
sh ad ow ed f orth are th ese — T h a t g rou p of m orph ol ogi c a l

u nits or c ells , which we see int eg rat ed int o th e c omp ou nd
,
42 M O RPH OLO G I CAL D E V E LO PM E NT .
u ni t ca l ea f h a s i n ea ch high er pl ant a typic a l f orm

ll e d a , , , ,
du e to the s p eci a l a rr a ng em ent of th es e cell s a rou n d a mid

r i b a n d v e i ns I f th e m u ltipli c a ti on of m or ph ol ogi ca l u n i ts
.
,
a t th e ti m e w h en th e l ea f bu d i s t a ki ng on i ts m a i n ou t li nes
-
,
e xc ee d s a c er t a i n li m it th ese u ni ts b egi n to a rr a ng e th em
,
s e lv es rou n d s e c on d a ry cent res or li ne s of g r ow th i n s u ch , ,
wa y s a s to rep ea t i n p a rt or wh olly th e t ypi ca l for m : th e

, ,
l arger v ei ns b ecom e t ran s form ed i n t o i m p er fect m i d r ib s of -
p a rti a lly i nd ep end ent l ea v es ; or i nto c om pl ete mid rib s of -
qu it e sep ar ate l ea v es A n d as th ere goes on thi s t r ans iti on

, .
f rom a S i ngl e a gg rega t e of cell s to a gr ou p of su ch a ggr eg at es ,
th ere s i m u lt a neou s ly a ri s es by s i m il a r ly i n sens ibl e st ep s a , ,
di sti nct s t ru ct u re which su pp ort s the sev era l a ggreg at es thu s

p rod u c ed an d u ni tes th em i n t o a comp ou nd a gg reg at e T h es e
, .
ph enom ena sh ou ld be c ar efu lly s tu di ed ; Si nc e th ey giv e u s a

*
key to m ore i nv olv ed ph enom en a .
§ 1 8 9 T hu s far we h a v e d ea lt with l ea v es ordi narily so

.
c a lle d : b r i efly i ndi cati ng the h om ol ogi es b etween th e p arts

of th e s i m pl e an d th e c omp ou n d L et u s now t u rn to th e .
h omol ogi e s a m ong f oli ar org ans i n general T h es e h a v e b een .
There i s m u ch force i n th e c ri t i ci sm p as s ed on v
the a bo e p aragra ph, and
by i m pl ica ti on on s om e p re cedi ng p a r a gra ph s , t h a t th ough i n pl ants w h ic h

tend to p rodu ce c om p ound l e a v e s th e p roduc tion i s l argel y dependent on t he
s up pl y of nutri m ent , yet the u n q u a l i fie d s t a tem ent of th i s re l a t i on as a
ge n
e ra l one , is ne ga t i v ed b y t he e x i s te nce of p l a n ts whi ch b ear onl y s i m pl e
l ea v e s , h owe v e r m uch hi gh nut ri t i on ca us e s gro wth . B ut m ostl y v al i d th ough
t hi s j
i s , i t i s p rob a bl y not uni v ersa l l y v a l id
ob ec ti on . I am l ed to sa y t hi s by
wh a t occa s i ona l l y oc c urs i n owe r s The oweri ng fl . fl s tem of th e Hya cinth i s
s i ngl e
; b ut I h a e v s een a cul ti v a t ed Hya ci nth i n whi ch one o f th e fl o we rs
v
h ad de el oped i nto a l a t eral v idence was once
s p ik e . St i l l m o re s t riki ng e
s u ppl i e d t o m e by A gri m ony A l l s a m pl e s of t h i s p l a nt p rev i ous l y s een ha d

.
s i ngl e fl ower i ng s p ik es bu t s om e yea rs ag o I m e t wi th one e x t re m e l y l uxuri

, ,
a nt i a whi ch som e fl owers of th e p ri m i ti v e s pi k e we r e r e pl a ce d by l a tera l

,
s p i ke s ; a nd I a m not s u re th at s om e of t h es e a g a i n d i d not bea r l a t era l , ,
S p i ke s Now i f i n p l ant s wh i ch i n p rob a bl y m i l l i on s of ca s es h a v e thei r

.
, ,
floweri ng s tem s s i ngl e ex ces s i v e nutri ti on ch ange s certa i n of thei r fl o wers i nto
,
ne w s p i k es , i t is a re as ona bl e s up pos i t i on th a t i n l i ke m a nner pl a nts wh i ch

a re th ough t i n v a ri abl y t o b ea r onl y s i ngl e l e av es , wi l l , u nde r ki ndred condi a
ti on s , b e a r com pou nd l e a v e s .
T HE M ORPH OLO G I CA L C O MP OS I T I ON OF P L ANTS .
43
m a de fa mi li ar to r ea d ers of na tu ra l hi s t ory by p op u l ari zed

ou t l i nes of Th e M eta m orp hos i s of P l a n ts — a t i t l e by the ,
wa y which i s fa r too e x ten s iv e ; S i n ce th e ph enom en a t rea te d

,
of u n d e r i t for m bu t a s m a ll p orti on of th ose i t p rop erly

,
i nc l u d es .
P a ss i n g ov er c ert a i n v a gu e a n ticip a t i on s w hi ch h a v e b een

q u oted from anc i ent wr it ers a nd noti n g on ly th a t som e ,
c l e a r er r ecogn iti ons w er e r ea c h e d by J oa chim J u ng a H a m ,
bu rg p rofes s or i n th e middl e of the 1 7th centu ry ; we c om e

,
to th e T heori a Gen er a ti om s whi c h Wol ff p u bli sh e d i n 1 7 5 9

'
, ,
a n d i n whi ch h e giv es d efi n it e for ms to th e c once p t i on s th a t
h av e si nc e b ec om e cu rrent S p ecifyi ng the vi ew s of Wol ff

.
,
D r Ma s ter s w rit es
“
. A ft er s p ea ki ng of the ho m ol ogou s
na t u re of th e l ea v e s th e s ep a ls a nd p e t a l s an h om ol ogy
, ,
c ons equ ent on th ei r s i m il a rity of st ru ctu r e and id entity of

or igi n h e g oe s on to s t a t e th a t the p eric a rp i s m an i festly ‘
,
c om p osed of s ev er a l l ea v es a s i n the c a lyx w ith thi s di ffer

, ,
en ce on ly th a t the l ea ves whi ch a re m e rely pl a c ed i n cl os e

,
cont a ct i n th e c a lyx a r e h er e u n it e d t og eth er

, a Vi ew w hi c h
he Cor rob or a t e s by ref err i n g to the m a nner i n which m a n y

c a p su l es Op en and s ep ar ate i nt o th ei r l ea ves ‘
T he see d s .
’
,
too h e l ook s u p on a s cons isti n g of l ea v es i n c l ose c om bi n a

,
ti on H i s rea s ons for c onsid er i ng the p et al s and s t a m ens a s

.
h omologou s with l eav es a re b a sed u pon the s a m e f a ct s a s

,
th ose which l ed L i nnaeu s and m a ny y ears a fterwar d s Goeth e

, , , ,
I n a wor d sa y s Wol ff
‘ ’ ‘
t o th e s a m e c on c l u s i on . we s ee , ,
n othi n g i n th e w h ol e pl ant wh os e p a rt s at fi rs t s ight di ffer

,
so r em ar k a bly f rom ea ch oth er bu t l ea v e s a nd s te m to whi c h , ,
l a tt er the root is ref err ibl e I t app ears th a t Wol ff t oo

’ ‘
.
, ,
e nu n ci at e d th e now a ccept ed i nt erp ret a ti on of c om p ou n d

-
fru it s : b a s i n g it on th e s a m e e vid ence a s th a t S i n ce a ss igne d .
I n the essa y of Goeth e p u bli sh e d thi rty y ea rs a ft er t h ese

, ,
r e l at i on s a m on g th e p a rt s of fl oweri n g pl a nt s we r e t race d ou t
i n g rea t er d eta il bu t not i n s o ra di ca l a w a y ; for Goeth e did

,
n ot a s did Wol ff v er i fy h i s hyp oth esi s by di ssecti n g b u d s i n

, ,
th ei r early st a ges of d ev el op m en t Goeth e a pp ea rs to h a v e .

44 M O RPH OLO G I CAL D E VELO PM E NT .
a rr ived at hi s l i i n d ep en d ently
c on c u s on s B u t th a t th ey .
w ere origi na l w i th him a n d th a t he g a v e a m ore v a r i ou s ly

,
ill us t ra t ed exp os iti on of th em th an h ad b een giv en by Wol ff ,
d oes not en titl e hi m to an yt hi n g b ey on d a secon d a r y pl a ce ,
a m on g th os e wh o h a v e e s t a bli s h ed thi s i m p or t a nt g en er a li z a
ti on .
Were it not th a t th ese p a ges m ay be rea d by s om e to

wh om B i ol ogy i n a l l i ts di vi s i ons i s a new su bj ec t of s t u dy it
, , ,
w ou ld be n ee dl ess to n a m e th e evid ence on w hi ch thi s now

fa mili a r gener a li za t i on res ts For th e i n for m ati on o f s u ch
.
it will s uffic e to s ay th a t the fun d a m enta l ki nship e xi s ti ng

,
a m on g a l l th e f oli a r org a n s of a fl ower i n g pl a nt i s S h own by ,
the t r an s i t i on a l form s whi c h m ay be t ra c e d b et ween th e m ,
a n d by th e occ a s i ona l a s s u m pti on of on e a n oth e r s f or ms

’
.
“
Fl ora l l ea v es or b ra cts a re frequ ently only to be di sti n
, ,
g u i s h e d f r om o r di n a r y l ea v e s by t h e i r p os iti on a t th e b a s e o f
the fl ower ; a t oth er tim es th e b racts g ra d u a lly a ssu m e m ore
”
a nd m or e of th e a pp ea r a n ce of the se p a ls T he sep a l s or .
,
divi s i on s of th e c a lyx a r e not u nlik e u nd evel op ed l eav es

,
s om eti m es assu mi n g qu i te the s tru ct u r e of l ea v e s I n oth er .
c a ses ,
th ey a cqu i r e p a r ti a lly or wh olly the c ol ou rs of th e
p etal s— as i nd ee d the b r act s a n d u pp erm ost st em l eav es
, ,
-
oc c a s i on a lly do S imil a rly the p et a l s Sh ow th ei r a lli a nces to

.
,
th e foli a r or ga ns l ower d own on th e a xi s a n d to th ose high er ,
u p o n th e a xi s On th e one h a n d th ey m ay d ev e l op i nt o
.
,
or di n a ry l ea v es th a t a re g reen an d v ei ne d an d on the oth er ,
h a nd a s s o com m only seen i n d ou b l e fl owers th ey m ay hea r

, ,
a n th er s on th ei r e dg es A l l v a r i eti es of g ra d a ti on i nt o
.
n eighb ou ri n g foli ar org an s m a y be wit nesse d i n s t a m ens .
Fl a tt ened and ti nt ed i n v a ri ou s d eg rees th ey p ass i ns ens ibly ,
i nt o p etal s and th rou gh th em p rov e th ei r h om ol ogy with

,
l ea v es ; i nt o whi ch i nd eed th ey are t rans form ed i n fl ower s

, ,
th at be c om e wh olly f oli ac eou s T he styl e too i s occ asi onally .

, ,
ch ang ed i nto p et als or i nt o green l eafl ets ; a nd ev en the

ovu l es a re n ow a nd th en s een to t ak e on l ea f lik e f or m s -
.
T h u s we h av e cl ea r evid enc e th a t i n P h aenoga ms, al l the

TH E M O RPH OLO G I CA L C O MP OS I TI ON OF P L A NT S .
45
a pp end ag es of the a xi s a re h om ol ogu e s : th ey ar e al l m odifi e d

l ea v es .
Wol ff e st a bli sh ed a nd Goeth e fu rt h er ill u s t ra t ed anoth e r

, ,
genera l l a w of s tru ctu re i n fl oweri n g pl a nts E ac h l ea f .
c omm on ly conta i n s i n i ts a xil a bu d simil ar i n s t r uctu re to ,
th e t er mi na l bu d T his a xill a ry bu d m ay re m a i n u n d e
.
vel oped ; or it m a y d ev e l op i nt o a l a tera l sh oot lik e t h e
m a i n s h oot ; or it m ay d e vel op i nt o a fl ower I f a S h oot .
b eari ng l a t er a l fl owers be ex a m i ned it w ill be fou nd th a t the

,
i nt ernod e or sp a c e whi ch s ep ara t es eac h l ea f with i ts a xill a ry

,
fl ower f rom th e l ea f a n d a xill a ry fl ower a bov e it b ecom e s ,
gr a du a lly l ess t ow a r ds th e u pp er end of th e She e t I n s om e .
pl a nt s a s i n the fox gl ov e th e i nt ernod es constitu t e a

,
-
,
r e gu l a r ly dimi n i shi n g s e r i es
-
I n oth er pl a nt s the seri es th ey
.
,
for m su ddenly b egi n s to di m i ni s h s o r a pidly a s to b r i n g the ,
fl ower s i nt o a sh ort s pik e : i ns t a n ce th e com m on orchi s A n d .
a g a i n by s till m or e s u dd en dw a rfi ng of the i n t ernod e s

,
th e ,
fl ower s a re b rou ght i nto a cl u st er ; as th ey are i n the cow

sli p. On c ont e mpl a ti ng a c l ov er fl ower i n which this ,
cl u st eri ng h as b een c a rri ed so far as to p rodu c e a com

p a ct h ea d ; and on c ons id er i ng wh at m u st h a pp en i f by a ,
f u rth er arrest of a xi a l d ev el opm en t the foot s talks of the ,

-
fl or ets di s app ea r ; it will be s een th at th ere mu st r esu lt a

c row d of fl owers s eat ed c l os e t ogeth er on the en d of the a xi s
,
.
A n d i f a t the s am e t i m e th e i nt ern od es of th e u pp er st e m
, ,
l ea v es a l so rem ai n u nd ev el op ed th ese st em l ea v es will be

,
-
g rou p ed i nt o a c om m on i nv ol u cr e : we s h all h a v e a c om p os i te
fl ower s u ch a s the thi stl e H enc e to m odifi cati ons i n th e
,
.
,
d ev el opm ents of f oli ar or g ans h a v e to be a dd ed m odific a ti on s

,
i n the d ev el opm ent s of a xi a l or g an s C omp a r i son s di scl os e.
th e gr a d a ti on s th r ou gh whi ch a x es lik e th ei r a pp end a ges , ,
p a ss i nt o al l v ar i eti es of si ze p rop orti on and st ru ctu re A n d

, ,
.
we l ea rn th a t the occu r ren ce of th ese two ki n d s of m eta m or

ph osi s i n a l l conce iv abl e d egrees and combi n ati ons fu rn i sh es
, ,
u s with a p r oxi m a t e i nt erp ret a ti on of m or ph ol ogic al c om
os iti on i n P h aenog a ms
p .
46 M O RPH OLO G I CA L DE V ELO PM ENT .
I s ay a p roxi m a t e i nter p ret a ti on b e c au se th er e r em ai n to

,
be solv e d c ert a i n d eep er p robl em s ; one o f w hi ch a t on ce

p resents i tsel f to be d ea lt w i th u n d er th e p r esent h ea d .
L e a v e s p et a l s sta m ens &c b ei n g s h ow n t o be h om ol ogou s

, , , .
,
f oli ar or g a n s ; a nd the p art to w hi ch th ey a re a tt a ch ed ,
p rovi n g to be a n i n defi ni tel y ex ten d e d a xi s of g rowth or

-
,
a xi a l o r g a n ; w e a r e m et by th e qu e s ti on s — Wh a t i s a f oli a r
,
or g a n ? a n d Wh a t i s a n a xi a l or g a n ? T h e m orph ol ogi c a l com

p os i ti on of a P h aenog am i s u n d et erm i ned s o l on g a s we c an,
not sa y to w h a t l owe r s t ru c tu re s l ea v es a nd s h oots a re h om o
l ogons ; a n d how t h i s i nt egra ti on of th em origi nates T o .
th ese qu esti ons l et u S now a dd ress ou rs elv es .
1 90 1 -
. A l rea dy , i n 78 , re fer enc e
b een m ad e to th e
h as
occ a s i on a l d ev el op m e nt of foli a r or g an s i n t o a xi a l or g a n s
the sp e ci a l c a s e th er e d escr ib e d b ei n g th a t of a fox gl ov e i n -

,
which s om e of th e s ep als w ere repl ace d by fl ower bu d s -

.
T he ob ser v a ti on of th ese a nd som e a n a l ogou s m on stros iti es ,
r a isi ng th e su s pi c i on t h a t th e di s ti n cti on b et ween f oli a r
org a n s a n d a xi a l or g a ns is n ot a b s ol u t e l ed m e t o e x a mi ne
,
i nto the m a tter ; a nd the resu lt h as b een the d eep eni n g of

thi s s u s pi ci on i nt o a c onvi cti on P ar t of the evid ence i s giv en
.
i n A pp en dix A .
S om e t im e a ft er h a vi n g rea ch e d this c onvi c ti on I fou n d on ,
l ooki ng i nt o the li tera t u re of the s u bj ect th at a n a l ogou s i r

,
re g u l a r i ti e s h a d s u gg e st ed to oth er ob s e r v er s b eli ef s s imil a r ly

,
a t v a r i an c e with th e c u rrent m or ph ol ogi cal c ree d D i ffi .
c u l ti es i n sa ti s fa ctor ily d efi ni n g th e s e tw o e l em en t s h av e ,
s er v ed to sh a k e thi s c reed i n s om e mi nd s T o oth ers th e .

,
s t r a n ge l e a f lik e d e v e l op m ent s w hi c h a x es u n d e r go i n c er ta i n
-
pl ants h a v e a fford ed r ea s on s for d ou b ti ng the c on stancy of

,
thi s di sti ncti on which v eg et a l m orph ol ogi sts u su a lly d ra w .
A nd th os e not oth er w i se rend e re d sc ep ti ca l h a v e b een m a d e ,
t o h es it a t e by su ch c ases as th a t of the N ep au l b a rl ey i n -
,
which the gl u m e a foli ar org a n b ecom es d ev el op ed i nt o an

, ,
a xi s a nd b ea rs fl owe r s I n h i s ess a y
. Vege t abl e Morph
47
”
o ol gy : i ts H i stor y a nd P res ent Con diti on w h e n ce I h a v e ,
a l r e a dy q u ote d D r Ma s te r s i ndic a te s s u n d r y of th e g r ou n d s
,
.
for thi n ki n g th a t th e r e i s no imp a s s a bl e d em a r c a ti on b et w een

l ea f a nd s te m A m ong other di fficu lti es w hi ch m eet u s i f w e
.
a ssu m e t h a t t h e di s t i n c ti on i s a b s ol u t e on e is i m pli e d by thi s ,
q uesti on Wh at sh a ll we s a y to ca ses s u ch a s th ose

:
a ffor d e d by t h e l ea v es of Ga a r ea a n d T m chi l i a
'
w h e r e t he ,
l eav e s a fter a time a s su m e th e conditi on of b r an ch es a nd de

vel op y oun g l ea fl et s f ro m th e i r f r ee e xt r e miti es a p r oc e ss l ess ,
p erfectly s e en i n s om e of th e pi nn a t e l ea v e d ki nd s of B er beri s -
or Ma honi a to be f ou nd i n a lm ost e v e ry s h ru bb e r y ?
,
A c l a ss of fa ct s on whi c h it w ill be d es i r a bl e for u s h er e to

d well a m om ent be fore p roc eedi n g to d ea l with the m a tter
,
d ed u ctively i s p r esented by the Ca c ta cece I n thi s r em a r k

,
.
a bl e g rou p of pl a n ts d evi a t i n g i n s u c h v a r i e d w a y s from th e

,
or din a r y p h ze nog a m i c typ e w e fi n d m a n y highly i n s tr u c tiv e

,
m odifi ca ti ons of for m an d s t ru ctu re By cont empl ati n g th e .
ch a ng es h ere di s pl ay ed wi thi n the li m it s of a s i ngl e ord er ,
we s h a ll grea tly w id en ou r con cepti on o f th e p os sibiliti e s of

m et a m orph os i s i n the v eget a l ki ngd om t a k en as a wh ol e , .
T wo di fferent bu t s i m il a r ly —s i gn i fi cant t ru ths a re illu st r at e d

, , .
Fi rs t we a re S h own h ow of th ese tw o c om p onent s of a

, ,
fl oweri ng pl ant com m only rega r d ed a s p ri m or di a lly di s ti n

,
g u i s h e d on e m a y a s su m e th r ou gh ou t nu m er ou s S p ec i e s th e
, , ,
f un cti on s a n d to a g rea t d eg ree th e a pp ear ance o f th e oth er

, , .
S econd we a r e s h own h ow i n th e s a m e i ndivid u a l th er e

, , ,
m ay occu r a re m et a m orph os i s : th e u su r p e d fu n ct i on a n d
-
a pp ea r a n ce b e i n g m a i nt a i n e d i n on e p a rt of th e pl a nt w hil e
,
i n a n oth er p a rt th er e i s a r etu rn to the ordi na r y a pp eara nc e

a n d fu nct i on We w ill con sid er th ese two t ru th s s ep a
.
ra tel y . S om e of th e E a p hor bi a c ece whi ch S i m u l a t e ,
C a ctu ses Sh ow u s th e s t a g es t h r ou gh w h i c h su c h a b n or m a l
,
s t ru c t u res a re a rr iv e d a t I n E u p hor bi a s p l en dens th e l a ter a l

.
,
a x e s a r e c on s id e ra bly s w oll e n a t th e i r di s t a l e n d s s o a s oft en ,
to be cl u b S h a p e d : s till h ow ev er b ei n g c ov e red w i th b ar k
-
, ,
Se e B ritis h a nd F or eig n M edi co Chi ru rg i ca l Revi ew for J a nu ary, 1 862 .

of th e di na ry c ol ou r a nd s till b ea ri n g l ea v es B ut i n
or , .
ki nd red pl an ts as E up hor bi a n er zi fol i a th i s swelli ng of th e

’
, ,
l a tera l a x es i s c a rri ed to a fa r g rea ter ex tent ; an d at th e ,
s a m e ti m e a g r een col ou r a n d a fl es hy con s i s t enc e h a v e b ee n

,
a cqu i re d : the typic a l r el a ti ons n ev er th e l e s s b e i n g s till S h ow n
by the few l ea v es t h a t g row ou t of th es e s oft a nd s woll en

a x es . I n th e Ca cta ceaz whi c h are thu s resem bl ed by pl a nt s
,
n ot oth er wi s e a lli e d to th e m w e h a v e i n dic ati on s of a

,
p ar a ll e l t ra n s for m ati on S om e ki nd s not com m only b rou ght

.
,
t o E ngl an d b ear l ea v es ; bu t i n th e s p eci es m os t fa mili ar to

,
us ,
th e l ea v es are u n d ev e l op e d a n d th e a x es a s su m e th ei r
f u n cti ons P a s s i n g ov er th e m any v a r i eti es Of for m a nd
.
c ombi n a ti on w hi c h t h e se g r e en s u cc u l en t g r owt h s di s pl a y w e ,
h av e to note th at i n s om e g enera a s i n P hyl l oca ctus th ey , ,
b ecom e fl a tt ened ou t i nto f oli aceou s sh ap es h a vi n g mid r ibs ,

-
a nd s om ethi n g a pp roa chi ng to v ei n s S O th a t h ere a nd i n .

,
th e gen u s E p i p hyl l u m w hich h as thi s ch ara ct er s till m ore

,
m a rk ed the pl an t a pp ea r s to be com p osed of fl eshy l ea v es

,
g rowi ng one u p on another A nd th en i n Rhi psa l i s th e

.
, ,
s a m e p arts are so l ea f lik e th at a n u ncriti c al ob serv er

-
,
w ou ld reg ar d th em as l ea ves T h es e which a r e a xi a l or g ans

.
i n th ei r h om ol ogi es h a v e b ec om e f oli a r org ans i n th e i r

,
a n a l ogi es . Wh en i nst ea d of c om p ari ng th ese

,
st ra ngely m odi fi ed a x es i n different g enera of Ca ct u ses we

-
,
c om p ar e th em in th e s a m e i ndividu al w e m eet with tra ns for

m ati on s n o l ess s t r iki ng Wh ere a t ree lik e f or m is pro
.
-
du c ed by th e g row th of th es e foli a ceou s s h oot s on e on a noth er ; ,
a n d wh er e a s a con se qu enc e th e fi r st fo r m ed of th em b e c om e
, ,
-
th e m a i n s t em th a t act s a s s u pp ort t o s econ d a ry a n d t erti a ry

s tems ; th ey l os e th e i r gr een su ccu l ent ch ara ct er a cqu i re
, ,
b ar k a nd b ecom e woody I n resu m i ng the fu ncti ons of a x es

,
.
t h ey res u m e th e str uct u r es of a x es from w hi ch th ey h a d d evi ,
a te d . I n Fig 7 1 a r e sh own s om e of th e l ea f l ik e a x es of
.
-
Rhi ps a l t s r hom bea i n th ei r y ou n g sta t e ; whil e Fig 7 2 rep r e

'
se nts th e old es t p orti on of th e s a m e pl a nt i n w hich th e foli ,
a ceou s ch ara ct ers are qu it e oblit er a t ed a nd th ere h a s res u lt e d ,

T HE M O RPH OLO G I CAL C O MP OS I T I ON OF P L A NTS .
49
an or di nary st em s t ru ctu re
-
. One furth er f a ct i s to
be not e d A t the s a m e ti m e th at th ei r l ea f l ik e a pp e a ra n ce s
.
-
a re l os t ,th e a x e s a l s o l os e
th ei r s e p ara t e i n divid u a li
“
ti es A S th ey b ecom e s tem
.
like they a l so b ecom e i nt e

,
grat ed ; a nd th ey do thi s so
ef fect u a lly th a t th ei r or igi
na l p oi nt s of j un cti on at ,
fi r st s o st ron gly m a rk e d a re effa c ed and a c on s olid a t ed t ru nk

, ,
is p rod u c ed .
J oi ned with the f a cts p revi ou s ly sp ecifi e d these f a cts ,
h elp u s to conce iv e h ow i n the ev olu ti on of fl oweri ng pl ants

,
i n gener a l th e m orph ol ogic a l c om ponent s th a t were onc e

,
di s ti nct m ay b ecom e ext rem ely di s gu i se d We m ay r ati on

, .
a lly e xp ec t th a t d u r i n g s o l on g a c ou r s e of m odific a ti on ,
mu ch gr ea t er ch a ng es of f orm a n d m u ch m ore d ecide d f u s i on s

,
of p a r t s h a ve t a k en pl a ce S eei n g h ow i n a n i ndivid u a l
, .
,
pl ant the si ngl e l ea v es p as s i nt o comp ou nd l ea v es by the

, ,
d ev el opm ent of th e i r v e i ns i nt o mid r ib s whil e the i r p eti ol e s

-
b egi n to s im u l a te a x es ; a n d s eei n g th a t l eav es or di na r ily

e xhibiti n g defi ni tel y li m it e d d e v el opm e nt s
-
oc c a s i ona lly p r o
,
d u c e oth er l eav es f rom th ei r edges ; we a re l ed to s u sp ect the

p os s ibili ty of s till g reat er ch ang es i n foli a r org an s Wh en .
,
f u rth er we fi nd th at withi n the limits of one n atu r a l or d er

, ,
p eti ol es u su r p the f u ncti on s a nd a pp e aranc e s of l ea ves at the ,
s am e time th a t i n oth er ord er s a s i n Ra s cus l at er a l a x es so

, ,
simu l a te l ea v es th a t th ei r a xi a l na tu re wou ld by m os t not be

su s p ect ed did th ey not b ear fl owers on th ei r mid r ib s or
,
-
e dg e s ; a n d wh en a m on g C a ct u se s
,
we p er ce iv e th a t su ch
,
m et a m orph os e s an d re—m et a m orph os es t a k e pl a c e w ith gr ea t
f a cility ; ou r s u spi ci on th a t the m or phol ogi ca l el em ents of

P h aenog a m s a d m it of p rofou n d t r a n s f or m a ti on s i s d eep ened , .
A n d th en on di s c ov eri n g h ow f requ ent a re th e m onst ros iti e s

,
whi ch do not seem s ati s fa ct orily expli c abl e wi th ou t a dmitti n g

the d ev el opm ent of foli a r organs i nt o axi a l org a ns ; we be c om e
50
50 M O RPH OLO G I CA L DE V EL O PM EN T .
r ea dy t t i the hyp oth es is th at du r i ng the evol u ti on

to en e r a n
of t he ph aenog a mi c typ e th e di sti nc ti on b et wee n l ea v e s a nd

,
a x es h a s a r i s en by d e g r ees .
With ou r p reconcepti ons l oosene d by su ch f act s a n d ,
c arryi n g wi th u s the g en er a l id ea w hich s u ch fact s su gg est ,
l et u s n ow c ons id er i n w h a t wa y th e typic al st ru ct u re of a
fl oweri ng pl ant m ay be i nt erp ret ed .
§ 1 92 p.r o cTo
ee d m e th o di c a lly w e m u s t s ee k a c l u e to ,
the st ru ctu re s of P h a n erog a ms i n th e st ru ct u res of th ose ,
i nferi or pl a nts th a t a pp r oa ch t o th em— A r cheg on ta tce T he -

.
v ar i ou s divi si ons of thi s cl a ss p r es ent a l ong with su n d ry ,
ch ara ct er s w hi c h a lly them w ith T h a ll ophyt es oth er ch ar ac ,
t ers by w hi ch th e ph aenog a mic st ru ctu re i s sh a d owe d for th .
Whil e s om e of the i nferi or H epa ti cce or L iv erwort s sev era lly ,
c on s i s t of li ttl e m ore th a n a th a ll u s lik e f rond a m on g th e -

,
high er m em b er s of thi s grou p and still m ore a m ong the ,
Mosses a n d F ern s we fi n d a di sti nctly m ar k e d s t em

,
*
S om e .
A rch egoni a t e s ( or ra th er Rhi zoid s ) h a v e f oli a r exp ansi ons

th a t a re i n d efi n it e i n th ei r for ms ; a nd s om e h ave qu it e
defi ni tel y Sh a p e d l ea v es
-
Roots a re p oss ess e d by a l l the
.
m ore d ev el op ed gener a of th e c l as s ; bu t there a re oth er

g en era a s S p ha gn u m whi ch h av e no roots
, , H er e the .
f rond s a re f orm ed of on ly a s i ngl e l ayer of c ell s ; and

th ere a d ou bl e l ayer giv es th em a high er ch aract er— a di ff er
Sc hl ei den wh o ch oos e s to rega rd a s a n a x i s th a t wh i ch Mr Be rk el ey
, .
,
wi th m ore obv i ous t ruth ca ll s a m i d r ib s a ys

, The fl a t s te m Of th e L i v er
-
,
worts p res ent s m a ny v a ri eti e s cons i s ti ng fre q u entl y of one s i m pl e l a yer of

,
thi n w l e d c e s o i t e x hi bi ts i n i ts a x i s th e e l em ents of the ordi nary s te rn

”
- a l l l r , .
Thi s p a ss age e x em pl i fie s the whol l y gr a tui tous hypoth es es wh i ch m en w i l l

s om et i m e s hypoth e s es they di s lik e Schl ei den, w ith t he
’
e s po us e, to e s ca pe .
pos i ti v enes s cha ra ct eri s t ic of hi m , a sse rt s th e p ri m or di al di s ti ncti on bet we en
a x i a l orga ns a nd fol i a r org a n s In t he h igh er A rche goni a te s h e s ees a n

.
undeni a bl e s tem . In th e l owe r A rch egoni a t es cl ea rl y al l i e d to t he m by,
the i r fr uct ifi c ati on, th ere i s no s tru ct ure h a v i ng the rem otes t re se m bl a nce
v
'
“ "
to a s tem . B ut t o sa e h i s hypo th es i s , Sc hl e i den ca l l s th a t a fl at s tem ,
whi ch i s ob vi ou sl y a s t ruct ure i n wh i ch s te m a nd l eaf a re not di ffer

e nt i a ted . He i s th e m ore t o be bl a m ed for th is unphi l osophi cal as s um p t i on,
m erci l es s i nh i s unph i l osoph ica l
'
s i nce he i s s tri ct ure s on th e a s s um ptions of
o t h er bota ni sts .
T HE M O R P H OLO G I CA L C O MP OS I T I ON OF P LA NTS .
51
c
en e e xhibit ed be tween cl os ely a lli ed g ener a of one grou p th e ,
Mosses E qu a lly v a r i e d a re the d ev el opments of the f oli a r

.
or g a n s i n th e i r d e t a il ed s tru c t u r es : n ow be i n g w i t h ou t mid
r ibs or v ei ns ; now h a vi n g m i d r ib s bu t n o v e i ns ; n ow h a vi n g
-
both mid r ib s a nd v ei ns N or mu st we om it the s imil ar ly s i g

-
.
-
n i fi cant c i rcu mst a nce th a t w her ea s i n t h e l ower A r ch e gon i a t es

,
th e r ep rod u ctiv e el e m ent s a re imm erse d h ere a n d th ere in the

th a ll u s li k e f ron d th ey a r e i n the high e r or d ers s eate d i n
-
, , ,
well spe ci a li zed a n d qu it e di sti nct f ru c ti fyi ng or ga ns h a vi ng

-
,
a na l ogi es w ith th e fl owers of P h aen og a m s T hu s m a ny f a c t s .

,
i m ply th a t if the P h aenog ami c typ e i s to be ana lyzed at al l ,
w e mu st l ook a m ong the A r ch eg oni a t es for its m orph ol ogica l

c ompo nent s and the m anner of the i r i nte gr ati on
,
.
A l rea dy we h a v e see n a m on g th e l ow er C ryp toga m ia how , ,
as th ey be ca m e i n t e gr a t e d a nd d efi n it e ly limi ted a gg re g a t e s ,
a cq u i r e th e h a bit of b u ddi ng ou t o t h er a ggr eg a t e s on r e a ch ,
i ng ce r t a i n st a ges of g rowt h C ell s p rod u c e oth er c ells

.
en d og enou s ly or e x ogenou s ly ; an d f ron ds giv e origi n to
oth er f ron ds f rom th e i r edge s or s u r f a c es We h a v e s een too .

, ,
th at the new a ggr eg at es SO p rod u ce d wh ether of the fi rs t ,
ord er or the s ec on d or d er m a y e ith er s ep a r a t e or re m a i n

,
conn ect ed Fi ss ip a rou sly mu ltiplyi ng c ell s i n s om e c as es

.
-
p a rt com p any w hil e i n oth er ca ses they u nit e I nt o threa d s or

,
l a mi nae or m a s se s ; a nd f rond s or igi n a ti n g p rolif erou sly f rom

oth er fr on d s s om e tim e s wh en m a t u r e di sconn e ct th e ms e lv e s
,
f rom th ei r p arent s an d s om etim es conti nu e a tt a ch e d to th e m

, .
Wh eth er th ey do or do not p a rt i s c l ea r ly d etermi ned by ,
thei r nu t riti on I f the c on diti ons are s u ch th a t th ey can

.
se v er a lly th r iv e be tt er by se p a r a ti n g a ft e r a c ert a i n d e v el op
m ent i s r ea ch ed it wi ll be com e the i r h a bit th en to sep ar a t e ;

,
s i n ce na tu ra l s e l ecti on w ill fa v ou r th e p rop a g a ti on of th ose
whi ch s ep a rate m os t nearly a t th a t ti m e I f conv ersely it .

, ,
pr ofi ts th e speci es for the ce ll s or f ronds to c on ti nue l onger

'
a tt a ch e d whi ch it c an onl y do if the i r growth s an d Buhse

,
qu ent p owers of m u ltipli ca ti on a re thereby i nc rea sed it m u st ,
W en throu gh the conti nu al surviv a l of the fittest, tha t

'
,
52 M O RPH OLO G I CA L D E VE LO PM ENT .
l onger a tt achm en t will bec om e a n es tabli s h e d c h ar a cteri stic ;

and ,
by p er sist en c e i n this p rocess p erm an en t a tt ach m ent
,
will resu lt wh en p er m an ent a tt a ch m e nt is a dv ant a geou s .
T h a t di su ni on i s rea lly a c ons equ ence o f r el ativ e i nnu tr i ti on

a n d un i on a c on se qu e nc e o f r e l a tiv e nu triti on i s c l ear a ,
os ter i or i O n t h e on e h a n d th e s ep a r a ti on of th e new i n di
p .
,
v i du a l s wh eth er i n g er m s or a s d ev e l op e d a ggr eg a t es i s a
, ,
di ssolvi n g away of the con necti n g su b st a n ce ; and thi s i m pli es

th at the connecti ng s u b s t an ce h as c ease d to p erfor m its
f u n cti on a s a ch an nel of nu t r im ent On th e oth er h a nd
.
,
w h ere a s w e s ee a m on g P h aenog a ms th ere i s a bou t to t a k e

, ,
pl a ce a s ep ara ti on of new i ndivi d u al s i n the s h a p e of germs ,
a t the p oi n t wh er e th e nu t ri ti on i s th e l owe s t a s u dd en ,
i n crea s e of nu t r i ti on will c au se th e impe ndi ng sep ar ati on to

be a rr es t e d ; and the f ru ctifyi n g el em ent s rev e rti ng t owa rds
-
,
th e or di na ry form thereu pon d ev el op i n c onne xi on wi th the

,
p arent . T u rn i n g to th e A r ch egon i at es w e fi nd a m ong ,
th em m any i n di ca ti ons of thi s tr a ns i ti on f rom di s con ti nu ou s

d ev el opm ent to c on ti nu ou s d ev el opm ent T hu s th e L iv erwort s .
giv e or igi n to new pl a nt s by cell s whi ch th ey th row off f rom

th ei r su r f a ce s ; a s i nd eed we h a v e seen th at mu ch high er
, ,
” “
pl ants do “
A c c or di n g to B i sch off s a y s S chl e id en
.
, b oth ,
the c ell s of th e s t e m ( J u ng er m a nm a [ n ow L ap hocol ea ] bi den

’
ta l c ) a nd th ose of th e l ea v e s ( J cxs ec ta ) s ep a r a te t h e m selv es

.
a s p rop a g a tiv e c e ll s f rom th e pl a nt a n d i sol a ted c el l s sh oot

,
ou t a nd d ev el op w hil e s till c onne ct e d wi th th e p arent pl an t
i n t o s m a ll cellu l ar bodi es ( Metzg ert a fu rca ta ) w hi ch sep arat e

’
from th e pl a nt a n d g row i nt o n ew pl a nt s as i n Mm u m a ndro

'
, ,
e Moss e s ow i n th e way a bov e expl a i ned

”
gy n u m a m on g t h . N ,
th es e p rop a ga tiv e c ell s a nd p roli ferou s b u d s m ay c onti nu e ,
d ev el opi ng i n connexi on w ith the p a rent t o v a ri ou s d egrees

b efore s ep a rati n g ; or the b u d s whi ch are a bou t to be com e
f ru cti fyin g org an s m ay s i m il arly u nd er i nc rease d nu t riti on
, ,
d ev el op i nto y oun g f ron ds A s Si r W H ook er s ay s of the

. .
m a l e fru cti ficati on i n M etzg eri a fureuta I t h a s the a pp ear

,
a n ce of bei n g a y ou ng sh oot or i nnov a ti on ( for i n c ol our

THE M O RPH OLO G I CA L C O MP OS I T I ON OF P L A N TS .
53 ,
a nd t ext u re I c an pe rceiv e no differenc e ) roll ed u p int o

”
a s ph er ic a l figu r e On fi nding i n thi s s a m e pl an t th a t
.
,
s ometim es the p rolif er ou sly p rodu ce d f ron d bu d s ou t f rom -
it self anoth er f ron d b e for e s ep ar ati ng f rom the p arent ,
as s h ow n i n Fig 4 6 it b ecom e s cl ear

.
,
th a t thi s l on g conti nu ed connexi on m ay

-
r ea dily p a ss i nt o p erm a nent c onn exi on .
A nd wh en we s ee h ow e v en a m on g P h ae ,
nog a m s,
bu ds m ay eith er d et ac h th em
s elv es a s b ulbil s or r em a i n a tt ac h e d a n d
,
bec om e shoot s ; we ca n s ca rc e ly d ou bt
th a t a m ong i nfer i or pl ant s l ess d efi nit e i n ,
th ei r m od es o f organ i za ti on su ch t ra ns i ,
ti ons m u s t c onti nu a lly occu r .
L e t u s su pp os e th en th a t Fig 73 i s
, ,
.
the f ron d of som e p r i m itiv e A rc h egoni a t e ,
simil ar i n g ener a l ch ara ct er s to P el l i a

epi p hyl l a Fig 4 3 ; heari ng lik e it the
,
.
, ,
fru c ti fyi n g bu d s on its u pp er s u r f ace ,
a nd h a vi n g a slightly m ark e d mid r i b a n d

- -
rootl ets. A n d su pp os e th a t a s S h own , ,
a se c ond a r y f r on d i s p roli ferou s ly p r o
du c ed from th e mid —ri b a nd c onti nu es ,
a tt a ch e d to it . E vid en tly the ordi n a r y di s

conti nu ou s d ev e l op m en t can th u s b e c om e ,
a c on ti nu ou s d ev e l op m ent on ly o n c on ,
diti on th at th ere i s an ad equ a t e su pply ,
to the s ec ond a ry fr ond of su c h m a t eri a ls ,
a s a r e f u r n i s h e d by th e r ootl e t s : th e r e
m a i ni n g m a t er i a l s b ei n g obt ai n a bl e by
its el f from th e a i r H enc e th at p orti on of
.
,
the mid r i b lyi n g be t ween th e se c on d ary

-
f ron d a n d the chi e f rootl ets h a vi n g i ts fu ncti on i ncrea se d , ,
will in c rea s e i n b u lk A n a dditi on a l c on s equ en ce will be a

.
g rea t er con c ent ra ti on of th e rootl ets— th er e wi ll be ext ra

growt h of th os e which ar e m ost servic ea bly pl a c ed Obs er v e .
,
54 M O RPH OLO G I CA L D E V ELO PM E N T .
ne xt th at
, the st ru ct u re s o a r i si ng is likely to be m a i n
t a i n ed . S u c h a v a r i a ti on i m plyi ng a s it d oes c i r cu m st a nc e s
, ,
e s p e ci a lly f a v ou r a bl e to th e gr owt h of th e pl a n t w ill giv e ,
to the pl a nt e xt r a c h an c es of l ea vi n g d es c end a nts ; s i nc e

the a rea of f ron d s u pp ort e d by a giv en a r ea of th e soil ,
b ei ng great er th an i n oth er i ndivid u a l s th ere m ay be a ,
g reat er p rodu cti on of sp ore s A nd then a m ong th e m ore

.
,
nu m er ou s d escend a nts thu s s ecu re d by it th e v a r i at i on w il l ,
give a dv a nt a ges to th os e i n whi ch it recu rs S u ch a m od e .
of g r owth h a vi n g ,
i n this m a nn er b ec om e es t a blish ed l et
, ,
u s a sk w h a t i s n e xt lik ely to resu lt I f i t b ecom es th e

.
h a bit of the p r im ary f ron d to b ear a s econ d ary f rond f rom i ts

mid ri b this s econ d a ry f rond c omp os ed of phy s i ol ogi c a l
-
, ,
u n its of th e s a m e ki nd will i nh er it th e h a bit ; a nd s u pp os i ng

,
th at th e s u pply of mi nera l m a tt ers obt ai ned by the rootl et s

s uffic es for the fu ll d evel opm ent of th e s econ d ary f rond th ere ,
i s a lik elih ood th a t th e growt h from it of a t erti a ry f rond ,
will b ec om e a n h a bit u a l ch a r a ct eristic of the v a ri ety A l ong .
with the est ablishm ent of su ch a t erti ary f rond a s Sh own i n ,
Fig 7 4 th ere mu s t a ri se a f u rth er d ev el opm ent of mid ri b

.
,
-
i n th e p r i m a ry f rond a s w ell a s i n the s econ d a ry f rond— a

,
d evel op m ent whi ch mu s t b ri ng wi th it a g rea t er int egra ti on

of the two ; w hil e , s im u lt a neou sly e xt r a g rowth will t a k e
,
pl a ce i n su ch of th e r ootl et s a s are m ost di rectly c onnect e d

wi th thi s m a i n ch ann el of ci rcu l ati on With ou t fu rth er ex .
pl ana ti on it will be s een on i ns p ecti ng Figs 7 5 a n d 7 6

, .
,
th a t th ere m a y i n thi s m a nn er resu lt an i nt eg ra t e d s er i e s of

f ronds pl a c ed a lt erna t ely on opp os it e s id es of a c onnecti ng
,
v a s cu l ar s t ru ctu re T h a t thi s c onnecti ng v a s cu l a r s tru ctu re

.
w ill a s s h own i n th e figu re s b ecom e m or e di s ti nc t f rom the

, ,
f oli ar su r f a c es a s th ese mu ltiply is no u nwarran t ed assu m p

,
ti on ; for we h a v e seen i n com p ou nd l e a v e d pl a nt s h ow u n d er

-
, ,
a n a l ogou s con diti ons mid rib s b ecom e d ev el op ed i nt o s ep a

,
-
r a t e su pp orti n g p a rt s whi ch a cqu i re s om e of th e ch a ra ct er s

,
of a x e s whil e a ss u mi ng th e i r fu n cti on s A n d now

.
m ar k how cl ear ly the st ru ctu re thu s bu ilt u p by i nt eg rati on

T HE M O RPH OLO G I CA L C O MP OS IT I ON OF P L A NTS .
55
of p roliferou sly gr ow i ng frond s cor res p on d s wi th th e s t ru e

-
,
t u re of the m ore d ev el op e d J u ng er m a nn i a ceaz E a ch of the .
fron d s su ccess iv ely p rodu ced re p e a ti n g th e c h a ra cter s of i ts ,
p a rent wi ll b ea r roots ; a n d will hear th em i n h om ol og ou s

,
pl a ces as s hown F u rth er the u n i te d mid r ib s h avi ng bu t

, .
,
-
v ery littl e r igidi ty w ill be u na bl e to m a i nt a i n a n erect p osi

,
ti on H ence th ere will resu l t the re cu m b ent conti nu ou s ly

.
,
root ed st em whi c h th es e typ es e xhibit : a n e mb ry o ph aen o g a m

,
h a vi ng the wea kn ess of an e mb ry o *

.
A natu r a l c onc omit a n t of th e m od e of growt h h ere de

s c r ibe d is th a t the st e m w hil e it i nc rea s es l ongitu di na lly
, , ,
i nc reas es sc a rcely at a l l t ra ns v ers ely : h en c e the ol d n a m e

A crog ens Cl ea r ly the t r a ns v ers e d e v el op m ent of a st em i s
.
the correl a tiv e p a rtly of i ts f un cti on a s a c h a nn e l of ci rcu l a

,
ti on a nd p a rtly of i ts fu ncti on a s a m ech a n i ca l s u pp ort

, .
T h a t a n a xi s m a y lift i t s a tt a ch e d l ea v es i nt o the a i r impli es ,
thi ck nes s and s olidity p r op orti on at e to the m a ss of su ch

l eav es ; and a n i n crea s e of i ts s a p v ess el s a l so p rop orti onat e -
,
t o th e m a s s of s u ch l ea v es i s n eces s it a t e d w hen the root s a re ,
al l a t one end a n d the l ea v es a t t he o th er B u t i n the .
g en era lity of A c rogens th ese c onditi ons u nd er which a ri ses , ,
the nec ess ity for t r an s v ers e growth of the a xis a re a b sent ,
wh olly or i n grea t p a rt T he st em h a bit u a lly c reep s be l ow

.
the s u r f a c e or li es p ron e u p on th e su r f a c e ; an d wh er e it
,
grows i n a v ert i ca l or i n cli ne d di recti on d oes thi s by atta ch ,
i ng it sel f to a v erti c a l or i n cli ne d obj ect Moreo v er th rowi n g .

,
ou t r o otl et s a s it m os tly d o es
,
a t int e r v a l s th rou gh ou t i t s ,
l ength it i s not c a lle d u p on i n a ny con s id era bl e d e gr ee to

, ,
t r an s fer n u t ritiv e m at er i a l s f rom one of i ts en d s to the oth er .
To th i s i nterp reta t i on i t i s j
ob e ct ed t h at
“
th e m ore de v el oped J u ng er .
m a nni a cem
”
do not a ppe a r t o ha v e a ri se n fr om th e l ow e r form s of J u ng er
m a nni a oeae — th a t i s to sa y, from s u ch l owe r form s as a re no w e xi st ing . It
m a y, howe v er, b e conten ded x cl ude th e i nterp reta ti on
th a t th i s fa c t does no t e
gi v en ; s i nce th e hi gh e r form s m a y wel l h a v e bee n ev ol v ed not from any of ,
th e l o wer form s w e now know bu t from l ow er form s whi ch h a v e b e co m e

,
e xt i nc t .Thi s i nde ed i s th e i m p l i ca ti on of the e v ol ut i ona ry p roce s s a s

, ,
poin te d o ut i n th e note t o Ch a p I . . I f th en w e a s s um e som e ea rl y t ype of

i nte rm edi a te s t ruct ure , th e e xpl ana ti on m ay not i m probabl y hol d .
56 M O RPH OLO G I CAL D E VE LO PM ENT .
H enc e th is p ecu li ar ity which gives the i r nam e to the A c rogens ,
n ow c a ll ed A rc h eg on i a t es i s a n a t u ra l a c c om p a n im en t o f t he
,
l ow d eg ree of s p ec i a li z a ti on reach e d i n th em A n d th a t i t i s .
a n i n c id en t a l a n d n ot a ne c ess a r y p ecu li a r i ty i s d em on s t r ate d ,
by two conv erse f a cts On th e one h a n d i n th ose high er.

,
A crogen s whi c h lik e the t ree fer ns li ft l a rge m a ss es of

,
-
,
foli a g e i nto th e a i r th er e i s j u s t a s d ec id e d a t r an s v ers e e x

,
p a ns i on of th e a xi s a s i n dic otyl e d onou s t ree s On the ot her .
h a nd in th ose D i cotyl ed on s whi ch lik e the com m on D odd er

, , ,
g ai n s u pp ort a nd nu t rim en t from the s u r f a ces over whi ch

t h ey c ree p th ere is n o m ore l a t era l exp an si on of th e a xis
,
th an i s h a bitu a l am ong A crogens or A rch egoni ates Con .
cl u di ng a s we a re thu s f u lly j u stifi e d i n d oi ng th a t the

, ,
l a tera l exp a ns i on a ccom p an yi ng l on gi tu di na l ex tens i on whi c h ,
i s a gene r a l c h a ra cte r i s tic of P h anerog a ms a s di sti ngu i s he d

f rom A rch egoni ates i s nothi ng m or e th an a concomit a nt of
,
th ei r u s u a lly v erti ca l g row th ; l et u s now g o on to c ons id er

-
h ow v e r ti c a l g rowth origi n at es a n d wh a t a re th e st ru ct u ra l ,
ch anges it i nv olv es .
§ 1 9 3 P l ants d ep en d for th ei r p rosp erity m ai nly on ai r

.
a nd light : th ey d w i n dl e wh ere th ey a re s m oth ered a nd th r iv e

,
wh ere th ey c an exp and th ei r l ea v e s i nt o free s p a c e an d su n

s hi ne T hose ki n d s whi ch a ssu m e p rone p os iti on s c onse
.
,
q u ently l a b ou r un d er di sa dv a nt a ges i n be i ng h abitu a lly i nt er

fered with by one a n oth e r— th ey a re m u t u a lly sh a d ed a nd
I am in deb te d to D r Hook er for p oi nti ng out furth er facts s up port i n
.
g
thi s v iew . I n hi s F l or a A nta rcti ca , h e d es cri be s the genus L esson i a ( s ee
Fi g . a nd es p e cia l l y L . ov a ta , a s h a vi ng a m ode of
g rowt h s i m u l a ti ng th a t
of th e di c ot yl e donou s t re e s , not onl y
i n genera l form but i n i nt ern a l s truc
t ure . Th e ta l l v erti ca l s te m th i ckens a s i t gro ws , by t h e p e ri od i ca l a ddi t i on
of l a yers t o i ts p eri ph e ry v . Tha t e e n Tha l l ophytes s houl d thus , under
c erta i n condi t i ons , p re s en t a t ra ns v ers el -i nc rea si n xi s s how s th a t there
y g a ,
is nothi ng a b s ol ut el y ch a ra c te ri s t i c of P h an eroga m s in t he i r h a bi t of
s te m -
thi cke ni ng . Mr Ta nsl ey gi v es
. me furth e r v e rific a ti on b y t he s ta te
“
m e nt t h a t it is a l so now c ert ai n t h a t m em b ers of the E qu is et i new a nd
L ym p odi new , as well a s so m e Fe rn s whi ch fl ou ri s he d i n Ca rbonife rou s ti m es ,

had seconda ry th i ck eni ng i n t hei r s te m s q uite com pa ra bl e to tha t o f m odern
”
Di cotyl edonous tree s .
T HE MORP H OL OGI CA L C OMP OSI TION OF P L A NT S .
57 ,
mu tu a lly i n j u red S u c h of th em h owev er a s h a pp en by

.
, , ,
v a ri ati ons i n m od e of g rowth to r i s e higher th an others

, ,
a re m or e lik e ly t o fl ou r i s h a n d l ea v e of fs p r i ng t h an oth er s .
T h a t i s to s ay n a tu ra l se l ecti on w ill fa v ou r the m or e u p r ight

,
growi ng f orm s I ndivid u al s with s t ru ctu res whi c h li ft th e m

.
a bo v e th e re s t are th e fit tes t for th e con diti ons ; a n d by the

,
c onti nu a l s u rviv a l of th e fittest su ch s tru ct u res mu st b ecom e

,
es t a bli s h e d
. T h ere a re two es senti a lly di ff erent w ay s i n
-
whi ch the i nt eg ra ted s eri es of f ron ds a bove d escr ib ed m ay be ,
m odifi ed so a s t o acqu i re the stiffness need fu l for m a i nt a i ni ng

p erp en dicu l a rity We will c onsider th em s ep ara t ely
. .
A thi n l ay er of s u b sta n c e g a i n s g rea tly i n p ow er of res ist

i ng a t ra ns v erse st ra i n if it i s b ent rou n d s o a s to for m a
,
tu b e : wit ness th e di fference b etween the pli ability of a s hee t

of p a p er w h e n ou t s p rea d a nd th e r igidity of th e s a m e s h ee t
,
of p a p er w h en roll e d u p E ngi neer s c onst a ntly recogn i ze

.
thi s t ru th i n d evi s i ng a ppli a n ces by w hich the g rea tes t

,
st rength s h a ll be obt ai ne d a t the sm a ll est c os t of m at eri al ;

a n d a m on g or g a n is m s w e s ee th a t na t u ra l s el ecti on h a bit
,
u a l l y e st a blish e s s t ru ct u res c on f or mi ng to the s a m e p r i nc ipl e ,
wh erev er light nes s and s ti ffness a re to be c om bi ned T he .
cyli nd ri ca l b one s of m a mm a ls and bi rds and the h oll ow ,
sh a fts of f ea th er s a re ex a mpl es T he l ower pl ants too, fur

,
.
,
ni s h c a ses wh er e the st rength

need fu l for m a i nt a i n i n g a n
u p r ight p os i t i on i s a cqu i r e d
,
by thi s rolli n g u p of a fl at
th a ll u s or f rond I n Fig 7 7
. .
we h a v e an A l g a whi ch a p
p r oache s t owa r d s a t u bu l a r
di st r ibu ti on of s u b st ance ;
an d which h a s a c on se qu ent
r igidity . S u n d r y c om m on
f orm s of lich en h avi ng the,
th a llu s fold ed i n t o a b r a nch ed tu b e still m ore d ecid e dly dis

,
pl ay the c onnexi on b et ween this st ructu ra l arrangem ent

58 MORP H OL OGICA L D E VE L OP ME NT .
a nd this m ech an ic a l a dv a nt age A nd f rom the p articu l ar

.
c l a s s of pl an t s we a r e h e re d ea li n g with— the A r ch eg on i a tes
a typ e is s h own i n Fig 7 8 Ri el l a h eli cop hyl l a s i m il arly c h a r

.
, ,
ac teri z e d by a thi n f ron d th a t is m a d e sti ff enou gh to st a n d ,
by a n i ncu r vi ng which th ou gh it d oes not p r od u c e a h oll ow

,
c yli n d er
,
p rod u c es a ki nd red f orm I f th en a s we h ave .
, ,
s een na tu r a l s el ecti on or su r viv a l of th e fitt es t will f a v ou r

,
s u c h a m on g th es e r e c u mb en t A rch eg on i ate s a s a re en a bl e d ,
by v a ri ati on s i n thei r s t ru ctu res to m a i nt a i n r ai s ed p os tu res ;

,
it will f a vou r the f or m a ti on of f rond s th at c u r v e r ou nd u p on

th em s elves a nd c u rv e rou n d u p on the f ron d s g rowi ng ou t
,
of th em . Wh a t now will be the r esu lt sh ou ld s u ch a

, ,
m odifi cati on t a k e pl a c e i n the g rou p of p roliferou s fronds

r e p r esent e d i n Fig 7 6 ? . C l early the r esu lt will be a
,
s tru ct u re lik e th a t s h own i n Fig 7 9 An d i f thi s i nrolli ng

. .
bec omes more c ompl et e a form lik e J a ng er m a nni a c ordi fol ia

, ,
r ep r es ente d i n Fig 8 0 w ill .

,
be p rodu ce d .
Wh en the su cc es s iv e
f rond s a re thu s fold ed rou n d
so com pl et ely th a t th ei r
oppos it e edg es m eet these ,
Opp os it e edges will be apt to

u n it e : not th at th ey will
g row t ogether a fter be i ng

f orm ed but th at th ey w il l ,
79 - d eve l op i n c onnexi on ; or ,
”
i n b ot a nic a l l angu a ge will be c om e
,
a d nate T h at f oli a r
s u r f a c es whi ch i n th e i r e mb ry on i c s t a t e a re i n c l os e c ont a ct
, , ,
o ft en j oi n i n t o one i s a fa m ili ar f a ct
,
I t i s h a bit u a lly so
.
w ith s ep a l s or divi s i ons of the c a lyx I n a l l c a mp an u l at e

.
fl owers it i s so w ith p et a l s A nd i n s om e t rib es of pl ants

.
it i s s o with s t am ens We a r e th ere for e w ell w a rrant ed i n

.
i nferr i n g th a t u nd er the c onditi ons a bov e d escrib ed th e su c

, ,
cess i ve f rond s or l ea fl et s w ill by u n i on of th e i r r em ot e e dg es

, ,
fi rst at th ei r po i nts of origi n a nd a fterwar ds high er u p ,

THE MORP H OL OGI CAL COMP OSITION OF P L A NTS .
59
form sheaths i nsert ed one withi n anoth er and i ncl u di ng the ,
a xis . T his in cu rvi ng of the su ccess iv e f rond s ,
endi n g i n the f or m a ti on of sh ea th s m ay be a cc om p an i e d by
,
diff erent s ets of m odific a ti ons S u pp os i ng Fig 8 1 to be a

. .
t rans v erse s ecti on of su ch typ e ( a be i ng the m i d r i b a nd -

,
b t he exp a ns i on o f a n o ld er f ron d ; w hil e 0 i s a y ou nger f ron d

p roliferou s ly d evelop ed withi n it ) th ere m ay be gi n two di
,
v ergent ki nds of ch anges l ea di ng to two cont ras t e d s t ru e

,
t u res I f while f ron d conti nu es to grow ou t of f rond the

.
, ,
s eri es of u ni ted mid r ibs conti nu es to be the ch annel of ci rcu

-
l a ti on be tween the u pp er m ost f ronds and the roots —i f a s a ,
c ons eq u enc e the c omp ou n d mid ri b or ru dim ent a ry a xis con

,
-
, ,
t i nu es to i nc rea s e in s i ze l a t era lly ; th ere will a ris e the s eri es

of t ra ns iti ona l f or m s rep r es ent e d by the t ra nsv ers e s ecti ons
8 2, 8 3 8 4 8 5 ; endi ng in the p rod u cti on of a s olid

, ,
v ywh ere wr app ed roun d by the f oli a r s u r f a ce of th e

e er
frond a s a n ou ter l a y er or s h ea th
, Bu t i f on th e oth er
.
,
h a nd ci rcu m st a nces fa v ou r a form of pl an t wh i ch m a i nt a i ns

,
i ts u p rightn ess a t the s m a ll es t cos t of su bs ta nce— i f th e

v a scu l ar b u ndl e s of each s u ccee di ng mid r i b i n st ea d of r e
-
,
m ai ni n g con cen t ra t ed bec om e di s t ribu te d a l l rou n d the t u be

,
60 MOR P H OL OGI CA L D E V E L O P ME NT .
for me d by the i n fold ed frond ; th en th e s t ruc tu re ev entu a lly

reac h ed th rou gh the t ran s i ti ona l form s 8 6 8 7 8 8 8 9 will
, , , , ,
*
be a h oll ow cyli n d er . A n d now ob serv e h ow th e
t w o s t ru ct u res thu s p r od u c ed corresp on d wi th tw o ki n d s of
,
Mon ocotyl e dons Fig 9 0 r ep resen t s a s p eci es of D en drobi u m

. .
,
i n which w e s ee cl early h ow ea c h l ea f i s bu t a c onti nu a ti on

o f th e e xt erna l l a y e r o f a s olid a xi s — a sh ea t h su c h a s wou ld
r esu lt from th e i n f old e d edg e s of a f r on d b ec om i n g a d n a te ;
a nd on e x a mi ni n g h ow the s h ea th o f ea c h l ea f i n cl u d e s th e
one a bov e it a n d h ow the s u cc es siv e sh ea ths i n cl u d e the a xi s

, ,
it will be m anif est th a t the rel a ti on s of p a rts a re j u st s u ch

.
as e xist i n th e u n it e d s er i es of f r on d s s h ow n i n Fig 7 9 — th e .
s u ccess iv e nod es a nswer i ng to the su cces siv e p oi nts of or igin

of th e f ron d s Conv ers ely th e stem of a gr a s s Fig 9 1 dis
.
, ,
.
,
pl a ys j u st su cl ti ons o f p a r t s a s w ou ld resu lt f rom the de

h re a ,
vel op m ent of the typ e s h own i n Fig 7 9 i f i n st e a d of th e m i d

.
,
r ib s thi ck en i n g i nto a s olid a xis th e m a tt er c omp os i ng th e m

,
b eca m e e v enly di s t rib u t ed r ou n d th e f oli a r s u r fa c es a t th e ,
s a m e ti m e th a t the i n c u r v e d e dg es of th e foli ar su r f a c es
u n it e d
. T h e a rra ngem ents o f th e t u b u l ar a xi s a nd i ts a p
p enda g es thus r esu lti n g ar e still m ore i n st ru ctive th an thos e
, ,
See note a t the end of th e cha pte r.

THE MORP H OL OGICA L COMP OSIT ION OF P L A NTS .
61
of th e so lid a xi s . v For whil e,

m ore cl ea rly th a n i n the
e en
D endr obi u m we see a t the poi n t b a c onti nu ity of st ru ct u re

, ,
b etween the s u bstance of the a xi s b el ow th e node a n d th e ,
s u b sta n c e of th e s h ea th a bo v e th e nod e : w e s ee th a t t hi s
s h ea th i nst ea d of h a vi ng i t s ed g es u n i te d a s i n D endr obi u m

, ,
h a s th e m s i m ply ov erl a ppi n g s o a s to f or m a n i n com pl ete

,
h oll ow cyli nd er w hi c h m ay be t ak en off a n d u n roll ed ;

a nd w e s ee th a t w er e th e ov er l a ppi ng e dg es of t hi s s h e ath
u n it e d a l l th e w ay f ro m th e n od e a t o the nod e 6 it wou ld ,
c on s tit u t e a t ub u l a r a xi s lik e th a t whi c h p r ec e d es it or lik e

,
th a t which it i nclu d es A nd th en givi ng a n u nexp ect e d

.
,
concl u s iveness to th e ar gu m ent it t urn s ou t th a t i n one ,
f a mily of gr a ss es th e ov erl a ppi ng edges of th e sh ea th s do

,
u n i te : th u s fu rn i s h i n g u s w i t h a d em on s t ra ti on th a t tu b u l a r
s t ru c t u r es a r e p rod u c e d by the i ncu r vi n g a n d j oi n i n g of
f oli ar s u r f a c e s ; a nd th at so h oll ow a x e s m ay be i nt erp rete d

,
as a bov e w i th ou t m a ki ng any a ssu mpti on u n wa rr a n te d by

,
f a ct. One fu r th er corres p on d enc e b et w een th e typ e

thu s id eally cons tru ct ed a nd the m on ocotyl ed onou s typ e, ,
m u s t be not ed I f a s a l re a dy p oi nt e d ou t the t ra n sv ers e

.
, ,
g rowt h of a n a xi s ar i ses wh en the a xi s com es to be a ch anne l

of c i r c u l a ti on b e t we en a l l the roots a t one o f i ts e xt r e miti e s
an d a l l th e l e a v es a t th e oth er ; a n d if thi s l a t e r a l b u lgi n g
m u st i ncr ea s e a s f a st a s the qu a ntity of foli a ge to be b rou gh t

i n com m u n i c a ti on w i th th e root s i n c rea s es— e s p ec i a lly i f su c h
foli a ge h a s a t th e s a m e tim e to be r a i s ed high a b ov e the
ea r th s s u r fa ce ; w h a t m u s t h a pp e n to a pl an t c ons t ru c t e d i n
’
the m a nner j u s t d es c r ib ed ? T he eld er f rond s or f oli a r or ga n s ,
en s h ea thi n g th e y ou nger ones a s w ell a s th e i n cipi ent ,
a xi s s e r vi n g a s a b on d of u n i on a re a t fi rs t of s u ch c i r cu m
,
f er en ce on ly a s s u ffi ces to i n cl ose th ese u nd ev el op ed p arts .
Wh a t then will t a k e pl a ce w h en th e i ncl osed p a rt s g row

, ,
wh en the a xi s thick ens whil e it e long a t es ? E vid ently the

ea r li e s t for m e d
-
s h e a th s n ot b e i n g l ar g e en ou gh for th e
,
s we lli ng a xi s mu st b u rs t ; a nd evid ently e ac h of the l a t er

,
for m e d sh ea th s m u s t i n i ts tu r n do the lik e ,

T h ere m u s t,
.
62 MORP H OL OGI CA L D E VE L OP ME NT .
s lt a g ra du a l exfol i ati on of th e s u cces siv e sh eath s lik e

'
re u ,
th at i n di ca te d a s begi nni n g i n the a b ove figu r e of D cndr o

bi u m ; whi ch at a s h ows the bu d of the u n d ev e l op e d p a rt s
, ,
j u st vi s ibl e a b ov e the e nwra ppi n g sh eaths whil e at b a n d c , , ,
it s h ows the o ld er s h ea th s i n p rocess of b ei n g s plit op en .
T h at i s to s ay th e re m u s t res u lt th e m od e of grow th which

,
h elp ed to giv e the na m e E ndog ens to thi s cl a ss .
The oth er way i n w hi ch an i nt egra t e d ser i es of fron ds

m ay a c qu i re the r igidity n ee d fu l for m a i nt a i n i n g a n erect
p os iti on h a s next to be c ons id ered I f the s u cc essiv e f ronds

,
.
do not a cqu i re su c h h a bit of c u r li n g as m ay be t a k en a d

v a nt a ge of by na tu ra l se l ecti on s o as to p rodu ce the requ i s ite
,
s ti fin ess ; t h en th e on ly way i n which th e r e qu i s it e sti f

,
f nes s
a pp ear s p rod u c ibl e i s by the thi ck en i n g a n d h ar d eni ng of
,
th e fu s ed s er i e s of mid r ibs T he i ncipi ent a xi s w ill not i n

-
.
,
t h i s c a s e be i ncl osed by th e roll ed u p f ron d s ; bu t will con

,
-
t i nu e exp ose d S u rviv a l of th e fitt es t w ill f a v ou r th e g ene sis

.
o f a typ e i n whi c h th os e porti on s of th e su cc ess iv e mid r ibs

,
-
th a t enter i nt o the c onti nu ou s bond be c om e m ore bu lky th an ,
th e di s eng a ge d p orti on s of th e mid r ibs : the i ndivid u als -
w hich th riv e a nd h a v e th e bes t ch an ces of l ea vi ng off s p ri n g ,
b ei ng by the hypoth es i s i ndivid u a ls h a vi ng a x es s tiff

, ,
e n ou gh t o ra i s e th e i r foli a g e a bov e th a t o f th e i r f e ll ows .
A t the s a m e ti m e u nd e r the sa m e i nfl u ence s th er e will ten d

, ,
to re su lt a n e l on g a ti on of those porti ons of th e mid r ib s -

,
whi c h b ecom e p arts of the i ncipi ent a xi s ; seei ng th at it will

p rofit the pl a nt to h a v e i ts l ea v es s o far rem ov ed from one
a noth er a s to p rev ent m u tu a l i n t er fer enc es H enc e f rom th e
’
, .
,
r ecu mb ent typ e th ere w ill e v olv e by i ndi rect eq u ilib ra ti on

,
s u ch m odific a ti on s a s a re sh own i n Figs 9 2 9 3 9 4 ; .

, ,
the fi rst of whi ch i s a slight a dv an c e on the id ea l typ e

rep resent e d i n Fig 7 6 a risi n g i n th e way d es cr ibe d ; a n d
.
,
the oth ers of whi ch are a ct u a l pl a nt s— H a p l om i tm u m Hooker i

’
a n d P l a i ochi l a d e ci i ens T h u s th e high er A r ch e goni a t e s

g p .
s h ow u s h ow a l on g w ith a n a ss u mpti on o f th e u p r ight a tti

,
t u d e th ere d oes go on a s we see th ere m u st go on a separa

, , ,
‘
T HE MORP H OL OGICA L C OMP OS IT ION OF P L A NTS .
63
ti on of the l ea f p rodu ci ng p art s f rom th e root p rod u ci ng

- -
p a rts ; a g reat er d ev e l op m en t of tha t connecti ng por ti on of

the su c c es siv e frond s by which th ey a re k ept i n com m u ni c a
,
ti on with the roots and r a is ed a bov e the grou n d ; and a con

,
se qu ent i ncrea s ed di f ferenti a ti on of su c h connecti ng p orti on

f rom th e p arts a tt a ch ed to it A nd this l a t era l b u lgi ng of
.
the a xi s di rec tly or i ndi r ectly c on sequ ent on i ts fu n cti on s a s

,
a su pp ort a nd a c h annel b ei ng h ere u nres t ra i ned by the

,
ea r ly form e d f r on d s f o ld ed rou n d it
-
g oes on with ou t the
,
bu rs ti ng of th es e H enc e a ri s es a l ea di ng ch ar a ct er of wh a t
.
is c a ll ed ex og enou s g rowt h— a g row th which is h owev er s till , ,
h a bitu a lly a cc omp ani ed by exf oli ati on i n fl a sk s of the ou ter

, ,
m os t l ayers con ti nu a lly b ei ng cra ck ed a n d s plit by the a ccu

,
m u l a ti on of l a y er s withi n th e m A n d n ow if w e ex
.
a mi ne pl a nts of the ex og enou s typ e w e fin d a m on g th e m m a ny

,
di s pl ayi ng th e st a ges of this m et a m orph os i s I n Fig 9 5 is . .

,
s h own a for m i n w hich the con ti nu ity of th e a xi s w ith th e
mid ri b of the l eaf is m anifest— a c onti nu ity th at i s con

-
,
s pi cu ou s i n th e c omm on thi stl e H er e th e foli a r exp a nsi on

.
,
ru nn i n g s om e di st an c e d ow n th e a xi s m a k es the i ncl u d ed
,
p orti on of the a xi s a p art of i ts mid ri b ; j u st as in the id ea l

-
typ es a bov e d ra wn By the g reater g rowth of the int ernod es ,

.
64 MORP H OL OGI CA L D E V E L OP ME NT .
whi ch v ery v ar i a bl e not on ly i n di fferent pl ants bu t i n

a re ,
the s a m e pl a nt th ere resu lt s a m odific ati on lik e th a t de

,
l i neat e d i n Fig 9 6 A n d th en i n su ch f or m s a s Fig 9 7 th er e

. .
, .
,
is sh own the arra ngem en t th a t ar is es w h en by m ore r a pid ,
d ev el opm ent of the p roxim a l end of the m i d ri b th e di sta l -

,
p art of the foli ar su r f a c e is s ep a r at e d f rom the p a rt which

e m b r a c es th e a xi s : th e w i ng s of th e mid r i b s till s e rvi ng -
,
h owever to c onnect the two p orti ons of th e f oli ar s u r f a ce

, .
S u c h a s ep a r a ti on i s a s p oi n t e d ou t i n § 1 8 8 a n h a bit u a l
, ,
oc c u r r en c e ; a n d i n s om e c ompou n d l ea v e s a n a ct u a l tea r i ng
,
of th e i nt er v enou s ti ssu e i s c a u se d by e xt r a growth of th e

-
mid ri b Modifi cati ons lik e this a nd th e f u rth er one i n Fig

-
.
, .
9 8 we m a y e xp ec t to be est a bli sh e d by s u rviv a l of th e fitt es t

, ,
a m on g th os e pl an ts w hi c h p r od u ce c on sid era bl e m a s s es of
l ea ves ; s i nce th e d e vel opm ent of mid r ib s i nto footst a lk s by

-
,
th rowi n g the l ea v es fu rth er awa y from th e a x es will dimi ni s h ,
the sh a di n g of th e l ea v es on e by an oth er
,
A n d th en a m ong
.
,
pl a nt s of bu shy grow th i n which the a ss i m il ati ng su r fa ce s

,
’
b e come still m ore li a bl e to i nt er cept one anoth er s light ,
n a t u r a l s el e c t i on will c on ti nu e to giv e an a dv a nta g e t o th ose
which c a rr y th e i r a ss i m il a ti n g s u r f aces a t th e ends of the

p eti ol es and do not devel op a ss imil a ti ng su r f aces c l ose to the
,
a xi s wh ere th ey a r e m os t sh a d e d
,
Wh en ce w ill resu lt a d i s
.
a pp ea ra n c e of th e stip u l es a nd t he f o li a r f r i ng e s o f t he m i d
r ib s ; e n di n g i n the p r od u c ti on of th e or di n a r y s t a lk e d l ea f ,
Fig 9 9 which i s ch a ract eri s ti c of t rees Meanwhil e the a xis

.
,
.
,
thick ens i n p roporti on to th e nu mbe r of l eaves it has to

c ar ry and to pu t i n comm u nic a ti on w ith the r oots ; a nd so
,
T HE MOR P H OL OGICA L COMP OSITION OF P L A NT S .
65
there c om es to be a m ore m ar k ed t
con ra s t be tween it a nd th e
*
p eti ol es s ev era lly c a rryi ng a l ea f
,
ea c h .
1 94 . Wh en ,
i n th e cou r s e of
p roc es s a b ov e sk et ch e d th e
ou t t h e r e h a s a ri s en su ch c om m u n i ty of nu tr i ti on a m on g th e
,
frond s t h u s i nt e gr a te d i nto a seri es th a t the y ou n ger on es ,
a re a id e d by m a t er i a ls whi c h th e old er on e s h a v e e l a bor a t e d ;
th e y ou nger f ronds will be gi n to sh ow a t ea r li er a n d ea rli er ,
p er i od s of d ev el opm en t the s t ru ct u res a bou t to or igi nate

,
from th em A bu n d a nt nu t r iti on w ill a bb re vi a t e the i nt er v a l s

.
between the s u ccess ive prol i fica ti ons ; so th a t ev ent u a lly ,
whil e eac h fron d i s ye t imp e r fe ctly form ed the ru dim en t of ,
th e n ext will b e gi n to sh ow it se l f A l l e mb ry ol ogy j u s tifi es .
this i nference The ana l ogi es it fu rni s h es l ea d u s to e xp ect

.
th at wh en thi s seri a l a rr angem ent be com es org an ic the ,
growi ng p a rt of the se ri es w ill s h ow the g enera l rel a ti ons of

the forth com i n g p art s whil e th e y a r e v ery sm a ll a n d u n
,
s p ec i a li z e d Wh a t will i n su ch ca s e be the a pp ear anc es th ey

.
a ssu m e ? We s h a ll h a v e no di fficu lty i n p erceivi n g wh a t it

w il l be if w e t a k e a for m lik e th a t s h own i n Fig 9 2 a n d
, .
,
d wa r f i ts sev era l p a rt s at the s a m e tim e th a t we genera li ze

t h em .Figs 1 00 1 01 1 02 and 1 03 will sh ow the res u lt ;
.
, , , ,
an d in Fig . 1 04 , whi c h i s th e bu d of a di cotyl e dbn, we s ee
h ow l
c ea r i s th e
ph ol ogi c a l c or re s p on d en ce : a b ei n g the
m or
r u di m ent of a foli ar or g an b e gi n n i n g t o t a k e sh a p e
; b b ei ng
the a l m ost form l es s r u di m ent of th e ne xt foli ar or ga n ; an d
Sin ce thi s p ara gr a ph w a s put i n typ e [ th i s re fers to t he fi rs t edi ti on

],
I ha v e obs erved t h at in s om e v a ri e ti es o f Ci ne r a ri a , a s p rob a bl y i n oth er
p l a nts , a sin gl e i ndi v i du al fu r ni s h e s a l l t h e se form s of l ea v e s— a l l gra da t i ons
b e twee n uns ti p ul a t ed l ea v es on v
l ong p e ti ol e s , a nd l ea es t h at e m b ra ce the
a x is . It m ay be ad de d t h a t th e v
di s tri but i on of t hes e a ri ous form s i s ui te q
i n harm ony w i th th e ra t i ona l e a b o e v gi ven .
51
6 bei ng the qu it e u n di fferenti a t e d p a rt when ce the ru dim ents

-
of s u b s equ e nt f oli ar org a n s a r e to a r i s e.
A n d now we a r e p re p are d for e nt er i ng on a s till re m a i n i ng

-
qu esti on res p ecti n g th e s t ru ct u re of P h aenoga m s —w h a t i s th e

o r igi n of a xill a r y b u d s ? A s the s ynth es i s a t p resent s t a nd s ,
it d oes not a ccoun t for th ese ; bu t on l ooki ng a little m ore

cl os ely i nt o the m a tt er we sh a ll fi nd th a t the a xill ar y b u ds
,
a r e i nt erp r et a bl e i n th e s a m e m a nner a s th e t er mi na l b u d s .
S o to i nt er p ret th e m h owe v er we m u s t r etu rn to th a t pro

, ,
c es s of p r oliferou s g rowth w ith which we set ou t for the pu r ,
p ose of ob servin g som e fa cts not b efore n a med D el es s eria .
hyp ogl oss u m Fig 1 05 rep resents a s eaw eed Of the sa m e genu s
,
.
,
a s one ou tli ned i n Fig 4 0; bu t of a s p ec i es i n w hi c h pr o

.
l i fer ou s g rowth i s c a rr i ed mu ch f u rth er H ere n ot only d oes

.
,
the p r im a ry f ron d bu d ou t m a n y s ec on d a r y fron ds f rom i ts

mid ri b bu t m os t of th e s ec ond ary f ron d s s i m il a rly bu d ou t
-
s ev era l t erti ary f ron ds ; and ev en by s om e of the terti ary
f rond s thi s p r ol i fi c ati on i s r ep eat ed

,
. B es id e s b ei ng s h own
t h a t th e b u ddi n g ou t of sev era l f ron d s f rom on e f ron d m ay ,
b ec om e h abitu a l ; we are a ls o sh ow n th a t it m ay b ec ome a

T HE MORP H OL OGI CA L COMP OSIT ION OF P L A NTS .
67
h abit i nh erit ed by the f rond s so p rod u ce d and a ls o by the ,
f ronds they p rodu c e : the m a ni fes t ati on of the t end ency

be i ng p rob a bly limit ed only by f ailu re Of nu t r iti on T h at .
u nd er fit c on diti ons a n a na l og ou s m od e of g r owth w ill occu r
i n frond s of the a c rogenic typ e l ik e th os e w e set ou t w ith

, ,
i s s h own by the c a se Of M etzg em a fu r euta Fig s 4 5 4 6 i n

’
, , ,
.
whi c h s u ch com p ou n d p rol i fi cati on i s p arti a lly di s pl a y ed .
L e t u s s u ppos e th en th a t the f ron d a Fig 1 06 p rod u ce s

, , ,
.
,
not on ly a s i ngl e s ec on d ar y f ron d b bu t a l s o a noth er s u ch ,

'
second ary f rond b L e t u s s u pp os e f u rther th at the f rond
.
, ,
b is in lik e m a nner d ou bly p rolife rou s : p rod u ci n g b oth 0

'
L a s tly l et u s s u pp os e th a t i n th e s ec on d f ron d b
’
a nd c .
,
’
whi c h a p rod u ces a s w ell a s i n the s econ d f ron d c w hich b
,
p rod u ces the d ou bly p roliferou s h abit i s m ani fested

,
-
If .
,
n ow t hi s h a bit gr ow s or g a n i c— i f i t be com es a s it n a t u r a lly

, ,
will bec ome the c h aract er i stic of a pl ant of l ux u ri ant g rowth

, ,
t he u nfoldi ng p art s Of w hich c an be fe d by the u n f old e d

p art s ; it will h a pp en with e ach l a t era l s er i es a s w ith the ,
m a i n s er i es th a t its su c ces s iv e c om p onents w ill b egi n to

,
s h ow th em s elv e s a t earli er a n d ea r li er s t a g es Of d e v el opm ent .
An d i n the s am e way th a t by dwa rfi ng a nd g enera li zi ng

,
the origi na l seri es we a rr iv e at a st ru ct ure lik e th a t Of th e

,
t ermi na l bu d ; by dwarfi ng and g ener ali zi ng a l a t era l s er i es ,
a s sh own i n Figs 1 07— 1 1 0 w e a rr iv e a t a st ru c tu re a ns wer

.
,
ing in n atu re and p ositi on to the a xill ary bu d .
F act s confi rm i ng th es e i nt er p r et ati ons a r e a ffor d e d by the

st ru ctu re an d di str ibu t i on o f bu d s Th e ph aenog a m ic a xi s i n
.
i ts p r im or di a l for m b e in g a n i n t egr a t ed s er i e s of f oli a ; a n d

,
the d ev e l op m ent of t h at p a rt by whi ch th es e f oli a are

h eld t ogeth er at cons id er abl e di st a n ce s f rom one a noth er ,
t aking pl a ce a ft erward s ; it is i n fera bl e f rom the g enera l

MORP H OL OGI CA L D E V E L OP ME NT
“
68 .
p ri ncipl es of em b ry ol ogy th at i n i ts ru dim ent ary st a ges the

, ,
phaenog amic sh oot will h av e its foli a r p a rt s m ore cl early

m a rk ed ou t th a n i ts a xi a l p a rt s T hi s w e s ee i n e v ery bu d
. .
E v ery bu d c on si st s of th e ru dim en t s of l ea v es p a ck e d to
geth er wi th ou t any a pp reci abl e i nt ernod a l s p a ce s ; an d th e
i nt ernod a l s p a c e s be gi n to i n c rea se w i th ra pidi ty only w h en ,
th e foli ar or ga n s h a v e b een con sid e r a bly d ev e l op ed More .
ov e r w h ere nu t riti on f a ll s sh ort

,
a n d a r re s t Of d e v el opm ent
,
t a k es pl a ce— th at i s wh ere a fl ower i s form ed— the i nter

,
nod es r em a i n u n d ev e l op ed : the u n f oldi ng c ea ses b ef ore

the l at er—acqu i re d c h a r a ct er s of th e ph aen oga mic sh ee t
a r e a ssu m e d L a s tly a s th e hyp oth es i s l ea d s u s to e xp ect
.
, ,
a xill a r y bu d s m a k e th e i r a pp e a r a n ce s l a t er th a n th e foli ar
or g an s whi c h th ey a ccomp a n y ; a nd w h e r e a s a t th e e n d s of ,
s h oot s th es e foli ar org an s s how f a il u re Of chl orophyll th e

, ,
a xill ary b u d s a r e not p rod u c e d a t a l l T h a t th es e a re i n

.
fer a bl e t ra i t s of s tru ctu re will be m a ni fes t on i ns p ecti ng

,
Fig s 1 06— 1 1 0; a nd on Ob s ervi ng fi rst th at the d ou bly

.
, ,
p rolif erou s t en d ency of whi ch th e a xill ary bu d is a resu lt ,
impli e s a b u n d ant nu t r iti on ; a n d on ob ser vi ng next th at th e , ,
or igi n a l pl a ce of secon d a ry p rol i fi ca ti on is s u ch th a t th e f oli a r

,
s u r f a c e on which it occ u rs mu st gr ow to s om e ext e nt be fore

,
the bu d a pp e ar s .
On th u s l ooki ng a t th e m a tt er— ou c ont empl a ti n g a f resh

the id ea l t yp e s h ow n i n Fig 1 06 a nd n oti ng how by th e
.
, ,
c onditi ons of th e c a s e th e secon d a ry p rol i fi ca ti on s m u s t c ea s e

,
b efore th a t p rim a ry prol i fi c a ti on whi ch p rod u c es th e m a i n

a xi s ; w e a r e en a bl e d to rec on c il e a l l th e ph e nom e na of a xil
l ary gem m a ti on We s ee h a r m ony a m ong the s ev era l f act s

.
fi r s t th a t the a xill a ry bu d b ec om e s a l a t era l l ea f—b ea rin g

, ,
a xis i f th er e i s a b u n d a n t m a t e ri a l for g r owt h ; se c ond th a t ,
i ts d ev el opm ent i s a rres t ed or it b ecom es a fl ower b ea r i n g

,
-
a xis , if the s u pply Of s a p i s bu t m od era t e ; thi rd th a t it i s ,
a b se nt wh en th e nu t r i t i on i s fa ili n g We a re no l onger
.
c om m itt ed to the g ra t u it ou s a s s u m pti on th at i n th e ph m no ,
g a mi c typ e th ere m u s t exi s t an a xill ary bu d to ea ch f oli a r

,
T HE MORP H OL OGI CA L COMP OSIT I ON OF P L A NT S .
69
or gan ; bu t we a re l ed to
c oncl u d e ct pri ori th at which we , ,
fin d at p os teri ori th a t a xill a ry b u d s a re a s n orm a lly a b sent

, ,
i n fl ower s as th ey a re norm a lly p resen t l ower d own the a xi s .
A nd then to com pl et e the a rg u m ent we are p rep ared for th e

, ,
c oroll a ry th a t a xi ll a ry prol i fi ca ti on m a y n a t u ra lly a r is e ev en
a t the en d s Of a x es s h ou ld th e f a ili n g nu t r i ti on whi c h

,
c a u ses the dwarfing Of the foli ar organs to form a fl ower be ,
su dd enly c h a nged i nt o su c h high nu t r i ti on as to t ran s form

the components of th e fl ower i nt o a pp end a ges th a t a re green ,
if not oth erw i se l ea f l i k e— a conditi on un der whi ch only this

-
,
phenomenon is p rov ed to occ ur .
§ 1 95 . m orOn e
e q u es ti on p res ents it self wh en we con ,
t ra st the early s t a ges of d ev el opm en t i n the two cl a s ses Of

P haenog a m s ; an d a fu rth er a n swer s u ppli e d by the hyp o ,
th es i s gives to the hyp oth es i s a f u rther p rob a bility I t i s

, .
ch a ra cteri sti c of a m onocotyl ed on to h a v e a s i ngl e s ee d l ea f

,
-
or c otyl e d on ; an d it is ch a r a c t er i s tic Of a dic otyl ed on to ,
h a v e a t l ea st two c otyl ed ons i f not m or e th an two T h at i s

, .
to say the m onocotyl ed onou s m od e of ger mi na ti on ev ery

,
wh ere c oexi s t s with th e en d og en ou s m od e Of gr owth ; a nd

a l ong with th e e x ogenou s m od e of g rowt h th er e a l wa ys goe s ,
e i the r a di cotyl ed onou s or p olycotyl ed onou s ger mi na ti on .
Why is thi s ? S u ch correl ati ons c an not be a cc identa l

c ann ot be m e ani ngl es s A t ru e th eory of th e p hae nog a m ic
.
typ es in thei r origi n an d divergen ce sh ou ld a cc ou nt for th e ,
conne xi on of th es e t ra its L et u s see wh eth er th e foregoi ng

.
theory d oes thi s .
The high er pl ants lik e the high er a n im al s bequ ea th to

, ,
thei r offs p r i ng m ore or l ess of nu t rim ent a nd s t ructu re .
S u p er i or or g a nisms of e ith er ki ngd om do not a s do a l l ,
n f
i e i or o g anism s c as t Off th ei r p rogen y i n th e sh a p e Of
r r ,
-
mi nu t e p orti on s of p rot opl a s m u nor g a niz ed and with ou t,
s toc k s Of m a te r i a l for th e m t o or g an i z e ; bu t th ey e ith er
d ep os it a l on g with the g erm s th ey c a st off cert a i n qu antiti es ,
of a lb u mi noid s u b s t an c e to be a pp r op r i a t e d by th e m w hil e th ey
d ev el op th em s el ves or e l s e th ey conti nu e to s u pply s u ch

,
s u b st a n c e whil e th e germ s p a rti ally d e vel op th em se lv es b ef ore

t h e i r d et ach m ent A m on g pl an ts thi s con stitu t es one di s
.
ti n cti on b etween s ee d s an d s p ores E very s ee d cont ai n s a

.
s t ore Of foo d to s er v e th e y ou ng pl a n t d u r i ng th e fi rs t s t a g es
o f i ts i n d e p e n d en t li fe ; a n d u s u a lly too b ef ore the see d i s

, ,
d et a ch ed the y ou ng pl a nt i s s o fa r a dva nced i n s tru ct u re

, ,
th at it b ea rs to the a tta ch e d s tock of nu trim ent mu ch th e

s a m e rel ati on th a t th e young fi sh b ear s to the a ppe nd ed yelk
bag a t the ti m e Of l ea vi ng th e egg S om etim es i nd eed the
.
, ,
d ev e l op m ent of chl orophyll give s the s eed l eav es a b right -
g reen whil e the s ee d is s till c on t a i ned i n the p a rent

,
pod . T hi s e ar ly org a ni za ti on of th e ph aen oga m

m u s t be s u pp ose d ru d ely to i ndi cat e the typ e ou t Of whi ch
the ph m nog a m i c typ e a ros e O n th e for egoi ng hypoth es is
.
,
the s eed l ea v es th ere fore re p resen t the p r im ordi a l f rond s ;

-
w hich i n d ee d th ey s imu l a t e i n th e i r s i m pl e c ell u l a r u n

, , , ,
v ei ne d s t ru ctu res A n d th e q u es ti on h ere to be as k e d i s

.
do the di ffere nt rel a ti ons Of th e p a r t s i n y ou n g m on ocotyl ed ons

a nd di c otyl e d ons c orr e s p on d w i t h th e diffe r en t rel a ti ons Of
th e p r im or di a l f rond s impli e d by th e en d ogenou s a n d the

,
e x og enou s m od es Of g rowth ? We sh all fi nd th a t th ey do .
S t arti ng a s b e for e w i th th e p roli ferou s for m s h own i n

, ,
Fig 1 1 1 it i s cl ea r th a t i f th e s t ren gth requ i red for m a i n

.
,
t a i ni ng the v ertic a l attitu d e i s Obt a i ne d by th e rolli ng u p Of

,
the f ron d s th e p ri m a ry fron d w ill m ore a n d m ore c onc ea l th e

,
s econd a ry frond w ithin it A t th e s a me ti m e the second ary

.
,
frond m u st c onti nu e to be d ep end ent on the fi rs t for i ts nu t r i

t i on ; a nd b ei n g p rodu ced w ithi n th e fi rst mu st be p re v ente d
, ,
by d e fectiv e su pply Of light an d a i r f rom ev er b ecomi ng syn

,
ch ron ou s i n i ts d ev e l op m en t with th e fi rs t H ence thi s .

,
i n foldi ng w hi ch l e a d s to the end ogenou s m od e of g rowth

°
,
impli es th a t th er e m u s t a l wa ys c onti nu e s u ch p re emi nenc e -
of th e fi rst for m e d fron d or i ts rep res en t a tiv e a s to m ak e the

-
,
germi nati on m onoc otyl ed on ou s Figs 1 1 1 to 1 1 5 sh ow the

. .
,
t ransiti ona l f or ms th a t wou ld res u lt f rom the infoldi ng of

TH E MORP H OL OGICA L COMP OS IT ION OF P L A NT S .
71
the f ronds I n Fig 1 1 6 ( a v ertic a l se cti on of the for m repr e

. .
s e nt e d i n Fig 1 1 5 ) a re e xhibit e d th e rel a ti on s of t h e su cce s

.
sive fron d s to ea ch othe r T he m odifi ed rel a ti ons th a t wou ld

.
res u lt if th e nu t r iti on Of the e m b r y o a d m itt e d of a nt i c ip a t ory

,
d eve l op m ent of the su cce ssiv e f rond s a re s h own i n Fig 1 1 7

“
,
.
A nd h ow rea dily th e s t ru ct u re m ay p ass i nt o th a t Of the

m onocotyl ed onou s ger m, w ill be s een on i nsp ectin g Fig 1 1 8 .
whi ch is a v ertic a l s ecti on Of an a ct u a l m onocotyl ed on at an

early st a g e— the i nc om pl et e li ne s a t th e l e ft of i ts r oot i n di ,
ca ti ng its c onn exi on with th e se e d Cont r ar iwi se

*
.
,
Since t hes e figures were put on th e bl ock , i t h as oc curr ed to m e th a t the

rel a t i ons w oul d b e s t il l cl ea rer were th e
, pri m a ry frond re pre sente d a s not
t aki ng p a rt i n th es e proce ss e s of m odi fi ca ti on, whi ch h a v e be en de sc rib ed a s
gi v ing ri se to the e rect form ; a s , i ndeed , t he rooti ng of i ts u nde r s u rfa ce
will pre v ent i t from d oi ng i n a ny con si de ra bl e de g re e In s uch cas e,

. e a ch o f
the Fi gs . 1 1 1 to 1 1 7 , sh oul d ha v e a h ori zonta l rooted frond at i t s ba s e,

hom ol ogous wi th th e pro e m b ryo am ong A c rogens
-
Thi s p ri m a ry frond
.
woul d then m ore m a nife s tl y s ta nd i n the sa m e rel a ti on to th e res t, as th e

72 MOR P HOL OGI CA L D E VE L OP ME NT .
where the s t rength requ i red for m a i nt a i ni n g a n u p right a tti

t u d e i s n ot Obt a i ne d by th e rolli ng u p of t he fron d s bu t by ,
the s t rength en i ng of th e conti nu ou s mid ri b th e s econ d -

,
fron d s o far from b ei ng l es s fa v ou ra bly ci rcu m s t a nce d th a n

,
th e fi rs t b ecom es i n s om e res p ec ts e v en m or e f a v ou r a bly

,
c i r c u m s t anc e d : b e i n g a b ov e th e oth er it g e ts a gr e a t er s h a r e
,
o f light a nd it i s l e s s r e s t r ic te d by s u r rou n di n g Ob s t a cl e s
, .
T h ere i s nothi ng t h ere fore to p re v ent it f rom r a pidly g a i n i ng

, ,
a n eq u a lity w i th th e fi rs t A n d if we a ssu m e a s th e t ru th s of
.
,
e mb ry ol ogy e ntitl e us to do a n i nc r eas i n g t en d enc y t ow a r ds

,
a n ti c ip a ti on i n th e d ev e l op m e nt o f s u b s eq u en t f ron d s— i f
.
w e a ssu m e t h a t h ere a s i n oth er c ase s s t ru ct u res which w er e

, ,
o r igi n a lly p ro d u ce d i n su cce s s i on w ill if t h e nu tr i t i on a ll ow s

,
a n d no m ec h a n i c a l d e pe n d ence hi n d e r s c om e to be p r od u ce d ,
s im u lt a neou s ly ; th ere i s n othi ng to p rev ent the p ass age

o f th e typ e re p r esen t e d i n Fig 1 1 1 i nt o th a t rep resent e d
.
,
i n Fig 1 22 . Or ra th er th ere i s ev eryt hi ng to f a c i l it a te it ;

.
,
see i ng th a t na t u r a l s e l e cti on will conti nu a lly f a v ou r the pr o
d u cti on Of a f or m i n whi ch the s econd f rond g rows i n su ch

way a s not to s h a d e the fi rst an d i n s u ch way as a ll ows the ,
a xis r ea dily to a s s u m e a v erti c a l p os iti on .
T h u s th en i s i nt erp ret a bl e th e u n iv ersa l connexi on be

, ,
t ween m onoc otyl ed onou s germi na ti on a nd en d og en ou s gr owth ;

a s w ell a s t h e simil a rly u n iv ers a l c o nn exi on be t wee n ex oge
-
n ou s g ro wt h a nd t he d ev el opm en t o f t w o or m or e c otyl ed ons .
T h a t it expl a i ns th es e fu nd a m ent a l rel a ti on s a dd s v ery ,
g rea tly to the p rob a bility Of the hyp oth es i s .
§ 1 96 . Whil ethi s m anner enabl ed to dis cern

we a re in
the ki nship th a t exi st s b et ween th e high er v eget a l typ es
th em selv es a s well a s b et ween th em and the l ower typ es ; we
,
cotyl e don doe s to th e pl um ul e —both by p os i t ion, a nd a s a s up pl i er of nu tri

m ent . Fig 1 1 7a , which I
. am ena bl e d to a d d, s ho ws th a t thi s woul d c om
p l ete th e i nte rp re ta t ion Of the di co tyl ed onous s e ri e s , i t i s needful to a dd no

.
f urt h er e xpl a na t i on th a n t ha t the diffe re nce i n h a b i t of g ro wth, wi ll pe rm i t

t h e s ec ond frond to root i t s el f as w el l as the fi rs t ; a nd s o to bec om e an a ddi
t i ona l s o urce of nut ri m ent, s i m i l a rl y ci rcum s t a nced to th e firs t and e qual wi th i t

.
T HE MOR P H OL OGICA L COMP OS ITION OF P L A NT S . ,
73
a re a t the m e time s u ppli ed wi th a r ati ona l e Of th ose t ru ths

sa
whi ch v ege ta l m or ph ol ogi st s h a v e est a bli s h e d T h ose h omo .
l ogi es whi ch Wol ff i ndi ca ted i n thei r c hi ef ou tli nes a nd

Goe th e foll owe d ou t i n d et a il h a ve a new m ea ni ng giv en to
,
th e m wh en we reg ar d the phaenog a mic a xi s a s h avi ng be en

e v olv e d i n th e wa y d escr ib e d F orm i ng the m odifie d con
.
c epti on which we a re h er e l ed to do res p ecti ng th e u n it s Of ,
which a fl oweri ng pl a nt is c omp osed we a re no l ong er l eft ,
with ou t a n a nswer to th e q u es ti on— Wh a t i s a n a xis ? A nd

we a re h elp e d to u nd e rs t a n d th e na t u r a l n ess of th os e cor
r es p ond enc es w hich th e su ccess iv e m em be r s Of e a c h s h oot
di s pl a y L et u s gl an ce at the f a cts f rom ou r p resent s t and

.
p oin t
’
Th e u nit Of c omp ositi on of a P h aenog a m i s su ch p orti on of ,
a s h ee t a s a ns wer s to one of the p r i m ordi a l fron d s T his .
p orti on i s neith er one of th e f oli a r a pp end a ges nor one of th e

i nt ernod e s ; bu t it c ons i st s Of a foli a r a pp end a ge tog eth er
wi th the p rec edi ng i n t ernod e i ncl u di ng the a xill a ry bu d
,
wh ere thi s is d ev elop ed T he p a rt s i nt ercept e d by the d otte d

.
li nes i n Fig 1 2 3 c ons titu t e s u c h a s egm ent ; a nd the tru e

.
,
hom ol ogy is be tween thi s a n d a ny oth er f oli ar org an wi th the

p orti on Of the a xi s b el ow it A nd now Ob s er v e h ow wh en we
.
,
ta k e thi s for th e u nit Of c ompos iti on th e m et a m orph os es ,
whi c h the ph ze nog a m i c a xi s di s pl a y s a re i n f era bl e f rom kn own

,
l aw s of d ev el op m ent .E mb ry ol ogy t ea ch es u s th a t a rres t

of d ev e l op m en t s h ow s its e lf fi r st i n th e a b s en c e of th os e p art s
th a t h a v e a r i s en l at est i n the c ou r s e of ev olu ti on ; th a t if

d e fect Of nu t riti on c au s es a n earli er a rr est p arts th a t a re of ,
m ore a nc i ent or igi n a b ort ; and th a t the p art a l one p rod u ce d

wh en the su pply of m at er i a l s f a il s near th e ou t set is th e p rim ,
or di a l p a r t . We mu st i nfer th erefore th a t i n ea ch seg

, ,
m ent of a P h aen og am the f oli ar org a n whi ch ans w ers to th e

, ,
p r i m or di a l f ron d will be th e m ost c on s t ant el em en t ; and

,
th a t th e i nt ern od e a nd th e a xill a r y bu d w ill be su cc ess iv ely ,
l es s c onst ant T his we fi nd A l ong w ith a s m all er size of

. .
f oli ar s u r f a c e implyi ng l ower nu t riti on it i s u s u a l to see a ,

74
. MOR P H OL OGICA L D E VE L OP ME NT .
mu ch dimini sh e d i nt ern od e a nd a l es s p ronou nce d a xi ll ary

- -
bu d a s i n Fig 1 2 4
,
On a pp roachi n g the fl ow er the
. .
,
a xill a ry bu d di sa pp ea r s ; a nd th e s e gm en t is re d u c e d to a
s m a ll foli a r s u rfa c e with a n i n t ernod e whi ch i s i n m os t

,
4
a. W a
s
c a s es v ery sh ort if not a bs ent a s in 1 2 5 a nd 1 2 6 I n the ,

.
fl ower i tse lf a xill ary b u ds a n d i nt ernode s a re b oth w an t

,
i ng : th ere r em a i ns on ly a f oli a r su r fa c e which ,
th ou gh oft en l arger th an the imm edi at ely p recedi ng foli ar

s u r f a c e sh ows f a ili ng nu t riti on by abs enc e Of chlorophyll
,
.
A n d th en i n th e qu it e t ermi n a l or ga n s of fru cti fi ca ti on ( 1 29 )

‘
we h a v e th e f oli a r p ar t it s elf redu c ed to a m ere ru dim en t .
T h ou gh th ese p r ogress iv e d eg enera ti ons a re by no m eans

re gu l ar b e i ng i n m any c a s e s v a r i ed by a d a pt a ti on s to p a r
,
ti cu l ar requ i rem ent s yet it c annot I think be qu esti oned

, , , ,
th at the genera l rel ati on s a re a s d es c rib ed and th at th ey a re ,
s u ch a s th e hyp oth es i s l ea d s u s to e xp ect N or a re .
we with ou t a ki nd red expl ana ti on Of c ert a i n rem a i ni ng t ra it s

Of foli ar org ans i n th e i r l eas t d evel op ed forms P et a l s
-
.
,
s t a m en s pistils &c , b es id es r emi ndi ng u s Of the p r im or di a l

, ,
.
f ronds by th ei r dimi ni sh ed si zes and by the want of th os e ,
s ev e r a l s u ppl em ent a ry p a rts which the p r ec edi n g s egm ents
p osses s a ls o remi nd u s of th em by thei r hi stologic a l ch ara c

,
t er s : they c onsist Of simpl e c ellu l ar ti ssu e sc ar cely a t al l ,
di fferenti at ed T he f ru cti fyi ng c ell s too which h ere m ak e

.
, ,
th ei r a pp earanc e are b orn e i n w ays lik e th os e i n whi ch the

,
l ower A crog ens b ear th em — at the edge of the f rond or at ,
the end of a p ed u ncl e or imm ers ed i n th e gen er a l s u b s t an ce ;

,
a s in Figs 1 2 8 a nd 1 2 9
. N ay it might ev en be s a id th a t
.
,
T HE MORP H OL OGICA L COMP OSIT ION OF P L A NT S .
75
the l
co ou rs a s su m e d by thes e t er mi n a l foli a c a ll to m i n d the ,
pl a nts ou t of whi ch we con cl u d e th a t P h aenoga m s h a ve been

ev olv e d for it is s a id of the f rond s of th e J u ng er m a nni a c ew
, ,
th a t th ou gh u nd er c ert a i n ci r cu mst an c e s of a p u re gree n

“
, ,
they a re i ncli ned to be s h a d e d with red p u rpl e chocol a te or , , ,

”
oth er ti nts .
A s th u s u nd ers t oo d th en the h om ol og i es a m ong th e p a rts

, ,
of th e ph aenog a m i c a xi s a re i nt er p ret a bl e not a s du e to a ,
nee dl e ss a dh es i on to s om e typica l f or m or f u lfilm en t Of a p re
d et erm i ned pl an ; bu t a s the i nevit a bl e c onseq u enc es of the

mod e i n whi ch the ph aenog am i c a xis origi nat es .
1 97 . A nd it rem a i ns only to ob s erv e i n con fi rm a

now ,
ti on of the foregoi n g sy nth es is th a t it at once e xpl a i ns for u s

,
v a ri ou s i rregu l a riti es Wh en we s ee l ea v es s om eti m es p ro

.
duci ng l ea fl et s f rom th ei r edge s or e xt re miti es we recogn i ze ,
i n th e a nom a ly a resu mpti on Of a n origi na l m od e Of gr owt h

fron ds f requ ently do thi s Wh en we l ea rn th at a fl ow eri ng
.
pl ant a s the D r os em i nter m edia h as be en k nown to d evel op

, ,
a y ou n g pl a nt f rom th e s u r f a c e Of one of its l ea v es we ar e a t ,
onc e remi n d e d of th e p r oli f erou s growths a n d f ru ctifyi ng
or g a n s i n th e L iv erwort s T h e occ a s i on a l p rod u cti on of bu l

.
bil s by P h aenoga m s c eas es to be s o s u rp ri s i n g w h en we fi nd

,
it to be h a bitu a l a m ong th e i nfer i or A crog ens a nd wh en we ,
s ee t h a t it i s bu t a re p etiti on on a high er st a g e Of t h a t sel f

, ,
d et ach m ent whi c h is c omm on a m ong p r oliferou s ly p rod u ced -
fron ds N or a r e we a ny l onger with ou t a s ol u ti on of th a t

.
t ra ns form ati on Of foli ar or gan s i nt o axi a l orga ns which not ,
u nc omm on ly t a k e s pl a c e H ow thi s l as t i rre gu l arity of de

.
ve l O m e nt is to be a c cou nt e d for we w ill h e re p a u s e a m om ent

p ,
to c on s id er L e t u s fi rst gl a nce a t ou r da t a
. .
The f or m Of e v ery or g a ni s m we h av e s een m u st d ep en d , ,
on the s t ru ctu res Of i ts phy s i ol ogi ca l

[ or cons ti tuti ona l ]
u n i ts . A n y g r ou p of su ch u n it s will t en d to a rran ge i tse lf
i nto the compl et e or g an i s m i f u ncont roll e d a n d pl a ce d i n fi t
,
conditi on s H ence the d e v e l opm en t Of fertili zed germ s ; a nd

.
76 MORP H OL OGI C A L D E V E L OP ME NT .
h en ce the d ev e l op m ent of thos e s el f d eta ch ed c ell s whi ch-
c h a r a c t e r i z e s om e pl a nts C on v erse ly phy s i ol ogi c a l u n i ts

.
,
w hi c h for m a s m a ll g rou p i n v olv e d i n a l a rg er grou p an d a r e ,
su bj ec t to a l l th e for c es of the l a r ge r g rou p w ill beco m e s u b ,
or di na t e i n th e i r s t ru ct u r a l a rr a n gem e nt s to th e l a r ge r gr ou p
—w ill be c o ordi na ted i n t o a p a rt of th e m a j or w h ol e i n

-
,
s te a d Of co ordi n ati ng th em selves i nto a mi nor wh ol e T his

-
.
a ntith es i s w ill be cl e a rly u n d er st ood on r em e mb eri n g h ow ,
on t h e o n e h a n d a s m a ll d e t ach e d p a r t Of a hyd r a s oon

,
m ou ld s it se lf in t o th e s h a p e of a n enti re hyd ra ; a nd h ow ,
on th e oth e r h a n d th e ce ll u l a r m ass th a t b u d s ou t i n pl ac e
,
’
Of a l ob ster s l os t cl a w gr a d u a lly a ssu m es th e form of a cl aw
,
— h as i ts p ar ts s o m ou ld e d a s to c o mpl et e the st ru c tu re Of
the or g a n i s m : a resu lt whi ch w e c a n not bu t a s c rib e to the

f orces which the rest of the org a ni s m ex ert s u p on it Con .
s eq u ently ,a m on g pl a n t s w e m ay e xp ect th a t w h eth er any

,
p orti on of p rot opl a sm m ou lds i tself i nto the typi ca l for m

a rou n d a n a xi s of i ts own or i s m ou ld e d i nt o a p a rt su b or
,
di m at e to anoth er a xi s w ill d ep end on the rel ative m a s s of

,
i ts phy s i ol ogic a l u nit s— the a c cu m u l a ti on Of th e m th a t h a s

t a k en pl ace b e fore the a ssu m p ti on of a ny s t ructu ra l arrange
m en t . A few ill u st ra ti on s w ill m a k e cl ea r th e v al idity of
thi s i nference I n the com p oun d l ea f Fig 6 5 the sev era l
.
,
.
,
l a t era l g row th s a b c d are m ani festly h om ologous ; a n d

, , , ,
on c omp a r i n g a nu mb er of s u c h l ea v es t og eth er it will be ,
s een th at one of th ese l a t era l gr owt h s m ay a ssu m e any de

g ree of c ompl exity a c cor di ng to the d eg ree of i ts nu t riti on
,
.
E v e ry f ern l ea f ex e mplifi es th e s a m e general t ru th s till bet

-
t er . Wh eth er e ach su b f rond rem a i ns an u n d ev el op ed wi ng

-
o f th e m a i n f ron d or w h eth e r it org a n i zes it s el f i nt o a g r ou p

,
o f fron dl ets borne by a s ec on d ar y r i b or wh eth er

, goi n g ,
fu rt h er a s it often d oes it giv es r i se to t erti ary ribs be a r

, ,
i ng frondl ets i s d et ermi ne d by th e s u pply of m at er i a l s for

,
growth ; si nc e s u ch high er d ev el opm ent s a re m ost m a rk ed

a t p oi nt s wh er e th e nu t riti on i s g r ea t es t ; n a m ely ne xt th e ,
t
s e m . B u t th e c l e ar e s t e vid e nc e i s a ff or d e d a m ong th e A l yce ,
T HE MORP H OL OGI CA L C OMP OSITI ON OF P L A NT S .
77
which not d r awi ng nu t ri m en t f rom roots h a v e th ei r p arts

, ,
mu ch l ess m u t u a lly d ep end ent ; and are th erefore c a p a bl e of

s h ow i ng m or e c l ea r ly h ow a ny p a r t m ay re m a i n an a pp en d
,
a g e or m ay b e c om e th e p ar en t of a pp en d
a g es ,
a c co r di n g t o ci rc u m st an ce s I n th e .
a nne x e d Fig 1 3 0 rep resenti n g a b r a nch Of

.
,
P ti l ota p l u m os a we s ee how a w i n g grow s

,
i nt o a wi ng b ear i ng b r a nc h if its nu t riti on

-
p as se s a c ert a i n p oi nt T his for m so s t rik .

,
i ngl y lik e th a t of th e fea th ery c ry sta lli z at i ons

of m a ny i nor g an ic s u b s t a nc e s i m pli es th at , ,
as i n s u ch c ry st a lli za ti ons the simplicity ,
or c om pl e xity of s t ru ctu re at a ny pl ace

d ep end s on the qu anti ty of m a tter th a t ha s to
*
be a rra nge d a t th a t pl a c e i n a giv en tim e .
H ence th en w e a r e not with ou t an i nt erp ret a ti on of th ose

, ,
o v er d ev e l op m en t s w hi ch th e ph aenog a mic a xi s oc c a s i on a lly

-
u n d er g oe s Fig 1 04 r ep resent s th e phaenog a mi c bu d i n its

. .
,
r u di m ent a r y st a t e T h e l at era l p r oc es s b whi c h ordi n a r ily

.
,
bec om es a foli ar a pp end a ge diff er s v ery littl e f rom the ,
t erm i na l p rocess 0 w hi ch 1 8 to be c om e an a xis— d iffer s

,
m a i n ly i n h a vi n g a t thi s p er i od wh e n i ts for m
, i s be i n g
d et erm i ned a s m a ll er bu lk
,
I f whil e thu s u n differenti a t ed .
,
i ts nu t r iti on rem a i ns i nfer i or to th a t of th e t erm i na l p roc ess ,
i t b e c om e s m ou ld e d i n t o a p a r t th a t i s su bo r di na t e to the
g en er a l a xi s B u t if as som etim es h a pp ens th ere i s s u ppli e d
.
, ,
to it su c h an a b u n d a nc e of th e m a t er i a l s n eedfu l for growth ,
th a t it b ecom es a s l a rg e a s the t er mi na l p rocess ; th en we

How th e e l e m ent of t i m e m od i fi es th e res ul t, is sh o wn by th e fa m i li a r
fa ct t h a t c rys ta l s ra pi dl y form ed a re s m a l l , a nd b ecom e re l a t i vel y l a rge when
l e ft t o form m ore s l ow l y . If th e q ua nti ty Of m ol ecul es co nt a i n ed in a s ol u t i on
is i v el y g rea t , s o th a t th e m u tu a l p ol a ri ti e s
l
re a t Of th e m ol ecu l es cr o wde d
togeth er i n e v e ry pl a ce th r ough ou t the sol uti on a re i nt ense , t here a ri s e s a
cry s t a l l i ne a g re a ti on a round
g g l oca l a xes ; wh e r ea s , i n prop ort i on as th e l oca l
a c ti on of m ol ecu l es on one a n o th e r is re nde re d l e s s i ntens e by th e i r wide r
d i s pe rs i on, they
b ecom e rel a t i e l y m o re v s u bord i na te to t h e fo rce s e x e r te d on
th em by th e l a rge r a ggr eg at e s o f m ol e c ul es th a t a re at gre a t e r di s ta nce s , and
t hus a re l eft t o a r ra nge th em s el es v round fe wer a xes i nto l arge r crys ta l s .

78 MORP H OL OGICA L D E VE L OP ME NT “
m ay n a u ra t lly
xpe ct it to b egi n m ou ldi ng it self rou nd eu
e -
a xis of i ts own : a f oli ar org a n w ill be r epl a ced by a n a xi a l
or g a n A n d this resu lt w ill be es p eci a lly li a bl e to occu r

.
,
wh en the gr owth Of the a xis h a s b een p revi ou s ly u nd ergoi ng

th a t arres t whi ch l ea ds to the form a ti on of a fl ower ; th at i s
wh en from d efect of m at eri al s the t er mi na l p roce ss has
, ,
a lm os t c ea se d t o i n cr ea s e a nd w h en s om e concu rren ce of
,
f a vou ra bl e c au se s b ri ngs a su dd en a ccess of s a p which reach es

the la tera l p roc esses b e fore it rea ch es the t ermi nal p roc ess *
.
1 98 . T he
genera l c on cl u si on to whi c h th e se v ar i ou s li nes
of e vid en c e c onv er g e i s th en t h a t th e s h oot Of a fl ower i n g
, , ,
pl ant i s an a ggr e ga t e of th e thi r d d egr ee of c omp os iti on .
'
T a ki ng a s a ggr eg a tes of th e fi rs t order those s m a ll p orti on s ,
of p rotopl a sm w hi c h or di n ar ily a ss u m e the f or ms u nd er
whi c h th ey a re k nown a s c ell s ; a nd c on s id e r i ng a s a gg reg a t e s

O f th e s ec on d or d er th e s e a ss embl a ge s o f s u ch cel l s whi ch
, ,
i n the l ower c rypt og a m s c omp ose th e v a r i ou s ki nd s of th a l

,
l u s ; th en t h a t st ru ct u re com m on to the high er c rypt oga ms

,
a n d to ph aenog a ms i n whi c h we fi nd a seri e s o f s u c h gr ou p s

,
Of cells bou n d u p i nt o a c onti nu ou s w h ol e m u s t be reg ard e d ,
a s a n a ggr eg a t e of th e th i r d or d er T he i n fer enc e d ra wn .
f rom ana ly s i s and v erifi ed by a s yn th esi s w hi ch corres p onds

,
i n a rem a r k a bl e m an n er w i th the f acts i s th a t th ose com ,
p ou nd p a rts whi ch i n Monocotyl ed ons an d D i cotyl ed ons a re

,
ca ll ed a x es h a v e r ea lly a r i s en by i n te g ra ti on Of s u ch s i m pl e
,
p a rt s a s i n l ower pl a nt s are c a ll e d frond s H ere on a high er .

,
It is j
ob ect ed t ha t t he s e t ra n s form a ti ons s h oul d be m uch com m on e r t h a n
th ey a re , w e re th e y c a u s ed sol el y by th e v a ri a t ions Of nu tri ti on d es c ribe d .
Th e r epl y i s th a t th ey a re com v
p a ra ti el y r a re i n u ncul ti a te d pl a nt s , where v
s uch v a ri a ti ons a re not fre q u ent Th e o cc urr en ce o f th e m i s ch i e y a m ong
. fl
c ul ti v a t ed p l a nt s whi ch b e i ng ,
a rt i fic i a l l y m a nu red , a re s pec i a l l y l i ab l e to
im m e nse a cc es s i ons O f nu tr i m ent , ca us e d now by s u dd en s upp li es o f fe rt i l i zi ng
m a tt ers , a nd no w by s u dden a ri
r v al Of th e roo t s at s uch m a t te rs a l re a dy
d e pos i ted i n t h e I t i s to t h es e grea t cha ng es

s oi l . of nu t ri t i on, es pe c i a l l y a pt
to t ak e p l a ce i n ga rden s , t h a t th e s e m ons tro s i t i es a re a sc ri be d ; and it seem s
to m e tha t t hey a re as frequent a s m ay be exp ec wd .

T HE MOR P HOL OGICA L COMP OS ITI ON OF P L A NTS .
79
l evel a pp e ars to h av e t a k en pl a ce a r ep etiti on of the p roc ess

,
a l rea dy Ob s erv e d on l ower l e v e ls T he form a ti on of those

.
s m a ll grou p s Of phy s i ol ogi ca l u nits whi ch c omp ose t he l owes t

p rot ophyt es is itsel f a p rocess Of i nt egr a ti on ; a nd the con
,
s olid a ti on of s u ch g rou p s i nt o defi ni tel y c i rcu m sc rib ed and

-
c oh erent c e ll s or m or ph ol ogi c a l u n it s i s a c om pl eti n g o f th e

’
p roc ess . I n th ose coa l es c en ce s by which m an y su c h c ells

a re j oi ne d i nt o th r ea d s,
a nd di s c s a nd s olid or fl a ttened
,
Ou t m a ss es ,we s ee th es e m orph ol ogic a l u nits a ggreg a ti ng

i nt o u nit s of a c omp ou n d ki nd : the di fferen t ph ases of the
t r ansiti on b ei ng ex emplifi e d by g rou p s Of v a ri ou s s i zes ,
v ar i ou s d eg rees Of c oh es i on and v a r i ou s d eg rees of defi ni te

,
ness . A nd now we fi nd e vid enc e s of a lik e p r oc es s on a

l a rger s c a le : th e c omp ou n d g rou ps a re a ga i n c omp ou nd ed .
Moreov er a s be fore th ere a re not w an ti n g typ es of org an i

, ,
za ti on by whi c h the st a ges Of thi s high er i n t e g ra ti on a re
sh a d owe d forth . F ro m f rond s th a t occas i ona lly p rodu ce

oth er f r on ds fro m th ei r su r f a ces we p a s s to th ose th a t
,
h a bitu ally p rod u c e th em ; f rom those th at do so i n a n i n

defi n ite m anner to th os e th a t do s o i n a d e fi nit e m anner ;
,
a n d f rom th ose th a t do s o s i ngly to th os e th a t do so d ou bly

,
a nd t r iply th rou gh s u ccess iv e g enera ti on s Of f ron d s E v en .
withi n th e limit s of a s u b c l as s we fi n d g rad a ti ons b etween

-
,
fronds i rregu l a r ly p r oli ferou s a nd g rou ps of s u ch f ronds

,
u n it e d i nt o a r e gu l a r s er i es
.
N or d oes the p roc es s end h er e T h e fl oweri n g pl ant is

.
r a r ely u n i a xi a l — i t is n e a r ly a l w a ys m u lti a xi a l F rom its .
p r i m a ry sh oot th ere g row ou t s econ d ary s h oots Of lik e ki nd .
T h ou gh oc c a s i on a lly a m on g P h ae noga m s a nd f re qu en tly

‘
a m on g th e high er C r ypt og a m s th e g erm s of n ew a x es d et a c h

,
them se lv es u n d er th e for m of bu lbil s a n d d e v el op sep ar a tely

,
i nst ea d of i n conn exi on with th e p a r ent a xi s ; yet i n m os t -
P h aenog a m s th e ge rm of ea c h new a xi s m a i n t a i n s i ts c on
nexi on with th e p ar en t a xi s : w h en ce re su lt s a g rou p Of a x e s
— an a ggreg at e of th e f ou rth ord er E v ery t ree by the pro
.
,
d u cti on of b ran ch ou t of b r anch sh ow s u s thi s i nt egrati on

,
rep e a t e d ov er a n d ov e r a g a i n ; f or m i n g a n a gg re g a t e h a vi ng a
d egree of c omp os iti on too compl ex to be any l onger d efi ned .
N O — A c r iti c i s m p assed on th e genera l a r gu m ent s et

[ TE
f orth i n the f oregoi ng s ecti ons ru ns a s foll ows — I h a v e

,
“
a l rea dy p oi n t e d ou t th a t th e p rocess of e v ol u ti on by w hi c h
you b eli ev e th e L iv e rwort s w ith a disti nct a xis a n d a pp e nd

a ge s to h a v e be e n p rod u c e d f rom th e th a ll oid f or m s i s n ot
f ou nd e d on sou n d evid ence e i th er i n c omp a ra tiv e m orphol ogy

o r d e v el op m en t . Bu t e v en if w e a d m i t th a t su ch a n i nte
g ra t i on o f a p r olif ero u s ly p
-
r od u ce d c ol on y might h a v e giv en
r i s e to th e l ea fy J u ng er m a nni a ceae ther e a re ev en m ore

,
w eighty obj ecti on s to th e su ppos i ti on th at the s a m e p rocess

p rod u c ed the s h oot st ru ctu res of the fl owe r ing pl ants I n the .
fi rst pl ace th e fl owe ri n g pl ant b ody is n ot hom ol ogous wi th

-
th e li verwor t p l a n t body si nce th ey rep r es en t diffe rent genera

-
,
ti on s. T he liv erwor t pl a nt b ody or ga m etop hy te i e the

-
,
. .
,
g ener a ti on b e a ri ng s exu a l org ans is h omol ogou s w ith the

,
p roth a llu s Of f erns and oth er P terid ophyt es and i n th e ,
Flower i ng P l ant s wi th red u ced s t ru ct u r es cont a i ned w ithi n

the s p ores ( e mb ry o s a c a nd p oll en g ra i n ) bu t s till givi ng r i se
- -
t o s ex u a l c ell s
. The liv erw or t spor e c ap su l e a nd i ts a ccessory
-
p arts ( i n f a ct ev ery thi ng p rod u ced from the fertili zed eg g ) i s

h om ol ogou s with th e s p orogon i u m of the m osses a nd as m ost , ,
b ot a ni sts thi nk with the l ea fy pl ant b ody of P t eridophyt es

,
-
a nd P h a n erog a ms . T hi s genera ti on i s c a ll e d the s p or op hyte

a n d f rom th e sp or es whi ch it p r od u ces a re d e v el op e d th e
g a m et ophyt es of th e next g enera ti on T h es e gener ali zati ons.
w ere fi rs t e st a bli sh e d by H ofm ei st er a n d a l l su b sequ ent ,
w ork h a s t end e d to es t a bli s h th em m ore fi r m ly T he .
on ly d ou btfu l qu esti on i s ( a n d th e d ou bt i s m a i n ly I thi n k ,

,
p ecu li a r to my self c ert ai nly not b ei ng sh ared by the m a j ori ty

,
of b ot an ist s
) wh eth er the sp orophyt e Of Mos s es and L iv e r
f orts i s rea lly h omol ogou s wi th th at of P t erid ophytes and
T HE MORP H OL OGICAL COMP OS IT ION OF P L A NT S .
81
P h a nerog a m s, wh eth er it m ay not ra th er be reg ar d e d a s a

p a ra ll el d e v el opm ent a l on g a noth er li ne of d es cent from th e
Green A lgae .
H en ce we m u s t l ook for th e or igi n of th e s h oot s t ru c t u r e -
of fl ow er i n g pl a n t s i n th e spor ophyt es of th e P t er id ophyt e s ,
from whi c h gr ou p th ere i s n o r ea s on to d ou bt th at th e

p h a nerog a ms h a v e a r i s en i n d es c ent T he v a rio u s gr ou ps of .
P ter id ophyt es v a ry m u ch i n th e org an i za ti on of th ese sh oot

s y s tem s as a m en t a l gl a nc e a t the type s e xhi bit e d by th e
,
F e rns H orse t a il s C lu b m os ses Op hi og l oss a ceaz a nd th e i so

,
-
,
-
, ,
l a te d I s oet es wi ll con vi n ce y ou a t onc e I t m a y be th a t .
som e of th e s e g rou p s a r e i n d ep en d en t i n d esc en t t e th a t th e , . .

,
P teri dop hyta a r e polyp hyl eti c a nd th e cu r rent hyp oth es i s

,
w i t h r eg a r d t o the ph a n er og a m s i s th a t th ey h a v e a r i s en by
t wo i f not th ree s ep ar a t e li nes of d e sc ent f rom difi er ent
, ,
grou p s of P t erid ophyt es ( thi s is i ndi ca t e d i n th e cl a ss i fi catory

di a gr a m on p 3 7 7 of vol I ) I sh ou ld not h owev er c a re to
. . .
, ,
p i n my f a i th to th ese or to a ny su c h li ne s Of an cest ry S till .
I thi n k w e mu s t l ook for the an c es t or s of the Fl ower i n g

P l ant s a m on g th e P t er id ophyt e s a nd th e l att er a l w a y s
,
h a v e a good disti ncti on b etween a xis a nd a pp end a ges T he .
p roblem of th e ev ol u ti on of th ese di ffer enti a t ed s p orophytic

s h oot s i s u n d ou bt e dly th e g rea t ou t s t a n di ng p r obl em of
m orph ol ogy Va ri ou s a tt empt s h a v e b een m a d e to s olv e it
.
,
of w hich p r ob a bly th e m ost imp ort a nt is th e th eor y of P r of s .
Bown a n d C a m pb e ll wh o d er iv e th e P t e r id ophy tes fr om s om e

,
L iverw ort lik e A n th ocer os bu t the s p or oph y te of c ou r se

,
fr om th e s p or ophyti c p ort i on of th e pl an t ( n ot m u ch m or e
th an a s p ore ca p su l e ) th e p roth a ll u s of th e F ern r ep res en ti n g
-
,
the v eget a tiv e th a ll u s of A n th o c eros I a m n ot wh olly con .
vi nced by th ese u n d ou b te dly i n g en i ou s hyp oth es es i n su pp ort ,

-
of whi c h a n i m m e ns e a m ou n t of fa ct s h a v e b een c oll ecte d ;
bu t my p os i ti on w ou ld I k now s i m ply pu t u s t o ign or a n ce

‘
, ,
a gai n on t hi s qu es t i on .
I h a v e di scu ss e d thi s a t s om e l eng th i n or d er t o b r i n g

ou t cl e ar ly th e i m m en se dif fi cu l ty of c on st ru ctin g a w e ll
52
82 M OR P H OL OGICA L D E VE L OP ME N T .
g rou nd ed th eor y of the or igi n of th e di fferenti a t ed s h oot

s y s te m of th e high e r pl a n t I con fess I d on t thi n k i t c an
.
’
be d on e a t a l l w i th the m a ter i a l s a t p r esent a t ou r di sp os a l .
Of c ou rs e i t i s j u st p os s ibl e t o s u pp os e th at s om e a nces tra l

s p or ophyt e h a d t h e s t r uc tu r e of a p r oli fe r ou s t h a ll oid liv e r
w or t g am e t ophyte a nd th at f rom it was ev olv e d th e ph a nero

,
g a mic s h oot i n th e wa ys you s u gges t T hi s giv es us abs o .
l u tel y n o c l u e h owev e r to a ny P ter id ophytic s h oot whi c h

, , ,
ou ght t o be i nt er m edi a t e ( m or e or l ess

) b etween th e hyp o
th eti c a l ancest or and th e P h an erog am and i s fu rt h er m ore , ,
a s fa r a s I ca n see n ot s u pp or ted by an a t om of evid en ce of

,
a ny ki n d . I t i s tru e th a t y ou r th eor y fi ts i n well wi th th e

ph enom ena exhibit e d by ph a n erog a mic sh oots th em s elv es ,
bu t thi s fa ct you will see m u s t l ose mu ch of i ts signific a nc e

if the hyp oth es i s l acks f ou nd ati on .
“
With reg a r d t o y ou r m eth od of expl a i ni ng the fu nd a
’ ‘ ’
m ent a l ch ara ct ers Of E x ogen s and E nd ogen s thi s of
‘
,
c ou rse is p ar t of the s a m e hyp oth es i s ; bu t I m ay p oi nt ou t

th a t si nc e Von Mohl a n d S an i o b etween 1 8 5 5 an d 1 8 6 5, ,
sh owe d ( 1 ) th a t the gr owth a t the s t em a p ex of a m ono

c otyl ed on w as n ot end ogenou s a nd ( 2 ) th at th e thi ck eni ng
,
‘
r in g n ear th e a p e x of a di cotyl e d on w as n ot to be con fu s ed ,
a s h a d b een d on e u p till th en w ith the r i n g O f s econ da ry

,
m er i s tem or tr u e ca m bi u m whi ch a ros e l ower d ow n a nd only

, ,
in w oody or p racti ca lly woody s t em th e t erm s E x ogen and ,
‘
E nd ogen
’
h av e n ecess arily fall en i nt o di su s e s i nce th ey ,
imply a fa l se c onc epti on Of wh at h a pp en s Both m on ocotyl .
‘ ’
e d on s a nd di c otyl ed on s h a v e a thi ck eni n g r i ng which ,
giv es r i se to the p r i m ary v ascu l ar cyli nd er of the st em .
Wh en th e st em is of c on s id era bl e thi ckn es s as i n P a l m s &c , ,

.
,
it grows by th e a ctiv e c ell divi s i on Of i ts ou ter l ayer s s o th a t

-
,
b oth c l a sses a re e x ogenou s i n thi s sen se ; whil e the a dditi on

o f a c ent rif u g a l z on e of second ary w oo d is c onfi ne d to c ert a i n
D i cotyl e d on s ( T rees s h ru b s
, ,
T he di sti ncti on b et ween the em b ry os m or eov er is not , ,
a b s ol u t e
. T he si ngl e cotyl ed on i s u su ally t er mi na l i n m ono
T HE MORP H OL OGIC A L COMP OS ITI O N OF P L A NT S .
83
c otyl ed ons bu t not a l ways ( D i os cora cece h av e l at era l cotyl

,
e d ons ) ,
bu t th e pl u m u l e m a y p u s h th rou gh it ( Gra sses ) or
m a k e i ts exit sid ewa ys ( P a l m s ) or be for m ed a t th e s id e
,
( A l i s m a ) a n d D i c o tyl ed ons v er y simil a r ly .
T h e oc cu rren c e of compl et ely sh ea thi ng l ea v es i n g ra s s es

is p erh a ps c orrel a t e d with the a bs en c e Of c a m bi u m bu t ,
g rass es a r e an a be rr an t typ e a mong m onocotyl ed ons and ,
s ec ond a r y thi c k eni n g is on ly f ou n d i n v ery few g en er a of
thi s c l a ss so th a t th e c orrel ati on is so to s p ea k neg ativ e and

, , ,
i ndi rect I t is cl ea r t h at the gr eat er p a rt of the dis

cu ss i on w ill h a v e to be re w r itt en -
.
F or th e rea son s a ss igned i n the p refa ce I c annot u n d ert a k e

t o re writ e th e di scu ssi on a s su gg es t e d
-
,
I t m u s t s t a n d for
.
wh a t it i s worth A l l I can do is h ere to i ncl u d e a l ong w ith

.
it th e foregoi ng criti ci s m s .
I m ay h ow ev er i n di cat e th e li ne of d e fenc e I sh ou ld t a k e
, ,
w ere I to g o a g a i n i n to the m a tt er T he Obj ec ti ons a re b a sed

.
on th e s t r u ct u r e o f e xi sti n g L iv erw orts an d P h aenog a m s Bu t .
I h av e a l rea dy r e ferre d to th e p rob a bili ty— o r i n d eed th e , ,
c ert a i n ty— th at i n con for m ity w ith th e genera l p r i n cipl e set

f orth i n the n ot e t o C h a pt er I we m u s t c oncl u d e th a t the
,
e ar ly typ es of L iv er w orts ou t o f w hi c h th e P h aenog a m s a re
s u pp ose d to h a v e ev olv ed a s well a s the early typ es of

,
P h aenog a m s i n whi ch the st a ges of ev ol u ti on were p re sent ed ,
no l on g er e xi s t We m u st i n fer th at for m s s impl er th an

.
a ny now kn own a n d m or e i n t er m edi a t e i n t h e i r t ra i ts w er e

, ,
the for m s concerne d ; a n d if so it m a y be h eld th a t th e ,
i n con gru iti es w i th the hyp oth es is whi c h a re p r es ent e d by

e xi s ti n g form s do not neg a tiv e it
, T he s cepti c i sm m y c r iti c
.
hi m sel f exp res s es r es p ecti n g th e c u r rent i nt er p reta ti on i s a

p arti a l j u stific ati on of thi s vi ew Mor eov er hi s a d m i ss i on
.
,
t h a t th e th eor y set forth “

fi ts i n well w i th th e ph enom en a
”
e xhibite d by ph a n er og a mi c s h oot s m u s t I t hi n k be r e g ar d e d
, , ,
a s weighty evid e n ce On th e E v ol u ti on hyp oth es i s we a re

.
-
oblige d to su pp os e th a t th e Mon ocotyl e d on s a n d D i cotyl e d on s
r es p e ctiv ely a r ose by i nt e g r a t i on of fron ds

; a n d i f to the
84 MORP H O L OGI C A L D E V E L OP ME NT .
qu esti on a ft er wh a t m anner th e i nt egr a ti on t ook pl a c e th ere ,
i s a n hyp oth es i s w hi c h ren d ers it c om p reh ens ibl e a nd a gr e es ,
b oth wi th th e s tru ctu re s of the two ki nds of s h oot s an d the

s t ru c t u res o f th e t wo ki nd s of s eed s a s w e ll a s w ith v a r i ou s
,
o f th e oth er ph enom en a th e two typ es p res ent it h a s s t ron g ,
c l a i m s for a c cept an c e .
Recons id er ati on su gg ests th e foll owi ng rem a r ks .
1 A lt ern a ti on of g enera ti ons i s a m ea ns of fu rth er i n g

.
m u ltipli c at i on T o be e ffectiv e e ach m e m b er of e i th er genera

.
t i on m u st be a s e l f su pp orti n g c ent re of g rowth or di ffu s i on

-
or b oth H en c e i f a s i n th e L iv erw ort s on e of th e s o c a ll e d

.
, ,
-
a lt ern a ti n g g en e r a t i on s i s n ot i n d ep en d en t bu t a p er m an ent
,
g r owth on the oth er— a p a r a s it e— i t is a mi su se of word s

t o c a ll the a rra n gem ent A l tern a ti on of gen era ti ons ( S i n ce .
this w as w r itt en I h a v e fou n d th at S i r E dw ard Fry t ak es the

s a me vi ew H e a pp r ovi ngly qu ot es P r ofess or B ower wh o
.
,
th e a lt ern ati on of g ener ati ons is not a n a cc u rat e

“
s a y s th a t
”
st at em ent of f a ct s or a u s e fu l ana l ogy )
2 T h e a lt er na ti ng Of s ex u a l a nd non s ex u a l p roc esses is
.
-
not fu nd am ent a lly di sti n ctiv e ; for a s sh own by

,
s u n d ry
A r ch egon i a t es it i s a n i nc ons t a nt t ra it a n d a s sh own by
, ,
K l ebs expe r i m ent s on Va u cher ia th e c onditi on s m ay be

’
,
v ar i ed s o a s to d eterm i ne its occu rrenc e or non occu rrenc e -

.
N ay th e s a m e i n divid u a l m ay rep rod u c e i n eith er w ay

,
.
3 S till m or e s ign ifi ca nt i s th e fa c t th a t i n s om e of th e
.
m ar i ne T h a ll ophyt es th er e i s a p roc es s lik e th a t w hich i n a

,
m oss or a fer n i s c on s id er e d a n a l tern a ti on of g en er at i ons ,
wh erea s i n oth er s a s th e B rown Wra c k ( F u cu s ) ea ch gen era

, ,
t i on i s sex u a l T hu s th e p r esence or a b s enc e of thi s m od e Of

.
g enes i s cann ot be a ca rdi n a l di sti ncti on .
4 Wi th th ese fa c ts b e fore u s it i s n ot only a rea s on abl e

.
,
s u pp os i ti on bu t a high ly p r ob a bl e s u pp os iti on th a t th e r e h a v e
,
e xi st e d pl a nt s of th e L iv e r wort typ e i n w hi ch th e so c all e d -
a l tern ati on of g en er a ti on s did n ot t a k e pl a c e I f s o n ea rly a l l

.
,
the f oregoi ng Obj ecti on s to my hyp oth esis f a ll to the g rou nd ]

C H A P TE R I V
TH E M O RP H O L O G I CA L C O M P O S I T IO N OF A N IMAL S .
§ 1 9 9 WH A T was s a id i n § 1 8 0 resp ec ti ng the u ltim a t e

.
,
st ru ctu re Of org a ni s ms h old s m ore m a nif es tly of a nim a ls

,
th a n of pl a nts T h at th rou gh ou t the v eg et a l kingdom the

.
c e ll i s th e m orph ol ogi c a l u nit i s a p rop os i t i on a d m i tti ng of a

,
b ett er d ef ence th a n the p rop os iti on th a t the c ell i s the m or

,
phol og i c a l u n it th rou gh ou t th e a n i m a l ki ngd om T he qu ali .
fi cati ons with w hi ch a s we s a w th e c ell d oct r i ne m u s t be

, ,
-
t a k en a re qu alifi cati ons th ru s t u p on u s m ore es pe ci a lly by

,
the f a ct s w hi c h zool ogi st s h a v e b rou ght to ligh t I t i s a m ong .
the P r otozoa th at th ere occu r nu m e rou s c a ses of vit al a ctivity

di s pl a y ed by s p ec k s of p rot opl a sm ; a n d from the mi nu t e
a na t omy of a l l c r e a t u re s a b ov e th e s e a r e d r a wn the nu m er ,
ou s p r oof s th a t n on c e ll u l ar ti ssu e s m a y a r is e by di rect m et a

-
morph os i s of mix ed colloid a l s u bst anc e s *

.
Si nce th i s p a ra ra ph
g wa s pub l i sh ed i n 1 8 6 5 , m uch h a s b een l ea rned c on
ce rni ng c el l s tru c tu re , a s
-
i s s h own i n Ch a pte r VI ‘l of P a rt Whil e s om e I .
as se rt t h a t the re e xi s t p orti on s Of li v ing p rotopl a s m w i th ou t n ucl e i oth e rs .
as s e rt th a t a n u cl eu s is i n e ve ry c as e pres ent , a nd th at wh e re i t doe s n ot ex i s t

in a de fini t e a gg r e ga ted for m i t e xists in a d i s p e rs e d form . As r e m a rk e d in
th e ch a pte r na m e d t he e v ide nce i s s om ew ha t s t ra i ne d to j us t ify th i s dogm a .
”
W ords a re ta ken 1n the i r nonina tural sens e s , i f one which connote s an i ndi
vidu a l i zed bo dy i s a p p l i ed t o t he wi del y diffu s ed c om ponents o f s uch a b ody ;
-
a nd v
th i s pe r e rti ng ofp rope r m ea ni ng s l ea ds to of wha t m a yO bs cura t i on
pe rha ps be an e ss ent i a l t ruth A s a rgued i n the cha pte r na m ed

. 74 e, 74f ) ,
nu cl e a r m atte r i s , as shown by i t s chem i ca l ch a ra cte r , a n e x trem el y unstabl e
s ub s ta nc e
, the m ol e cul ar ch anges of whi ch, perpetual l y goi ng on, ini ti a te
85
86 MOR P H OL OGICA L DE V E L OP ME NT .
Ou r v ey of m orphol ogi ca l c om p os i ti on th rou gh ou t the

su r
a ni m a l ki ngd om m u s t th er e for e b egi n w ith t h os e u n differ

,
en ti a ted a gg re g a te s Of phy s i ol ogi ca l u ni t s [ or c on s t i tu t i on a l
u nit s ] ou t of whi c h are form e d w h a t w e c a ll w i th c on s id er

, ,
a bl e li cen s e m or ph ol ogic a l un it s
,
.
s i on of th e P r otozoa di sti ng ui sh e d a s
§ th
2 00 I n
a t
. divi
Rhi zop oda a r e p r esent e d u n d er v a r i ou s m odific a ti ons th ese
, , ,
m i nu t e p ort i on s of livi n g org a n ic m a tt er s o littl e di fferent i ,
a t e d if n ot p os i tiv e ly u n di f
,
fer ent i a te d th a t a n i m a l i n divid u ,
a lity ca n sc a r c e ly be c l a i m e d for th em Fig s 1 3 1 1 3 2 an d . .

, ,
1 3 3 rep resent c er t a i n n ea r ly a lli ed typ es of th e s e— A m eaba ,

,
-
A cti nop hrys , a nd L i eber ku hm

vi scid j elly or s arcode
a . Th e ,
c om p a r a bl e i n i ts phy s i c a l p r op erti es t o w hit e of egg ou t of ,
w hi c h one of t h es e c r ea tu res i s m a i n ly for m ed s h ow s u s i n ,
v ar i ou s w a y s the f eebl enes s w i th whi ch the c om p onen t

,
phy s i ol ogi c al u n it s are i nteg ra te d— sh ow s u s thi s by its v ery

s light c oh e s i on by th e e xt r em e i n de fi ni ten ess a n d m u t a bility
,
o f i ts form a n d by th e a b s en c e Of a li m i ti n g m e mb ra ne
, .
I t is no l on ger h eld e v en by u n qu a lifi ed a dh er ent s of the

ce ll d oct r i n e th a t th e A m oeba h a s a n i n v e s tm ent
-
I ts ou t er .
s u r f a c e c omp a r e d to the film w hich for m s on th e s u r fa ce of

,
p a st e d oe s not p r ev en t the t aki n g of s olid p a rticl es i nt o th e

,
m a ss of th e b ody a n d d oes not i n s u ch ki n d r e d f or ms as

, ,
Fig 1 3 3 p rev ent the p seu d op odi a f rom c oal es ci ng wh en th ey

.
,
m eet H ence it c annot p rop er ly h av e the na me of a cell wall

.
-
.
A consid e r a bl e p or ti on of th e b ody h owev er i n D i / fi g

u i a , , ,
s h ock s ,p roduci ng c ha nges al l a round I n th e e a rl i e r s t a ge s o f c ell -e v ol uti on

.
t hi s u ns ta bl e s u b s ta nc e i s di s p ers ed th rough out t h e cyt opl a sm ; whe r eas i n

th e m ore a dv a nce d s t a g e s i t i s ga th e re d t oge th e r i n one m a ss I f so, i ns te ad .
o f s ayi ng th e re i s a di s p e rs e d nucl e us we s houl d s ay th ere a re th e m a teri a l s
o f a nucl e us not ye t i nteg ra t e d

. .
T HE MORP H OL OGICA L C OMP OSIT I ON OF A NIMA L S .
87
Fig 1 3 4 h as a d enser c oa ti ng form ed of a ggl u ti nated forei gn

.
,
p a rti cl es ; so th a t the p rot ru si on of th e p seu d op odi a i s limit ed

to one p a rt of it A n d i n the s oli ta ry F ora m i ni fer a lik e
.
,
Grom ia the s a r cod e is c ov ere d ov er m os t of i ts su r f a c e by a

,
d elica t e c a lc a reou s sh ell pi erc e d with mi nu t e h o l es th rou gh

, ,
which the s l en d er p s eu d op odi a a r e th ru st T he .
Grega ri na e xhibi t s a n a dv a nc e i n i nt e gr a ti on a n d a conse ,
qu ent g rea ter de fin i teness Fig s 1 3 5 a nd 1 3 6 ex emplifyi ng . .

,
thi s typ e s how the c om pl et e m em b ran e i n whi ch the su b

,
s ta n c e of th e crea tu re is con t a i n ed H ere th ere h a s a r i se n.
w h a t m a y be p rop er ly ca ll e d a ce ll : u n d er i ts s olit a ry for m

t hi s a n i m a l i s t ru ly u ni ce ll u l a r I ts em b ry ol ogy h a s c on
.
-
s i der a bl e s ign ific a n c e A ft er p a ss i ng th rou gh a cert a i n qu i

.
“ ”
e s c e nt ,
e n cy s te d st a t e i ts i nteri or b rea k s u p i nt o s m a ll
,
porti ons whi ch a fter th e i r e xit assu m e for m s lik e th a t of

, , ,
the A m oeba ; a n d f r o m thi s y ou ng c on diti on i n whi c h th ey

a r e u ndi ffere nti a t e d t h ey p a ss i nto th a t a d u lt c onditi on i n
,
which th ey h a v e li m iti n g m em b ra nes I f thi s d ev el op m en t .
of the i n divid u a l Gr eg o ri na typifi es th e m o d e o f ev ol u ti on o f
the s p eci es it yi eld s fu r th er s u pp or t t o th e b eli ef th a t f ra g

, ,
m ent s Of s a rcod e exi st ed ea rli er th a n a ny of the s t ru c tu res

whi ch a r e c a ll e d cell s A m on g a ggreg a tes of th e fi r st
.
or d er ,
th ere a r e s om e mu ch m ore highly d ev e l op ed T hese .
a r e th e I nfus or i a c ons tit u ti n g th e m os t nu m e r ou s of the

,
P r otozoa i n s p ec i e s a s i n i n divid u a l s
,
Figs 1 3 7 1 3 8 and . .
, ,
1 3 9 a re e x a mpl es
,
I n t h e m w e fi n d a l on g w i th g r ea t er
.
,
defi n i ten es s a c on s id e r a bl e h et e rogen eity

,
T h e s a rc od e of .
which the b ody c ons i st s h a s a n i n d u r a t e d ou t er l ay e r b ea ri ng

, ,
c ili a a nd som etim es s pin e s ; th er e i s a n op en i n g s ervi n g a s
m ou th a p er m anen t oesoph a gu s a nd a c a vity or c a vi ti es

, , ,
t em p ora rily form ed i n the i nt eri or of th e s ar c od e to s erv e a s ,
one or m or e s tom a ch s ; a n d th er e is a c omp a r a t iv ely s p ec ific
a r r an g e m ent of th es e a nd v a r i ou s mi nor p arts .
T h u s i n th e a ni m a l ki n gd om a s i n th e v eg et a l ki ngd om , ,
ther e exi sts a cl a s s of mi nu t e f or ms h avi ng thi s p ecu li ar ity ,
th a t no one of th e m i s s ep ar abl e i nt o a nu mb er of vi s ibl e

co mp on ent s h om ol ogou s with one a noth er— no on e of th em

c a n be res olv e d i nto mi n or i n divid u a liti e s I ts p roxim a t e.
u n i ts a re th os e p h y s i ol ogi ca l u n i ts of whi c h w e c oncl u d e e v e r y
or g an i s m c ons i s t s T h e a gg regate i s a n a gg re g at e of the fi rs t

.
or d e r .
201 . Am on g pl ants
f ou n d typ es i ndi cati n g a t ra ns i
are
t i on from a gg re g a t e s of
th e fi rs t or d e r to a gg r eg a tes Of th e
s econ d o r d er ; a n d a m ong a n i m a l s we fi nd a na l o g ou s typ es .
Bu t the s ta ge s of p rog r es si ng i nt eg r a ti on a r e n ot h ere so di s

ti n ct T he rea s on p rob a bly i s th a t th e s i m pl est a ni m a l s
.
, ,
h a vi ng i ndividu a liti e s m u ch l es s m a rk ed th an those of the

s impl e s t pl a n ts do n ot a f for d u s the s a m e fa ciliti e s for Obser
,
v at i on I n p rop orti on a s the limi ts of th e mi n or i n divid uali

.
t i es a re i nd efi ni te t he for m at i on of m a j or i n divid u a li ti es ou t
,
of th e m na tu ra lly l e a v es l e ss c ons pic u ou s t r a c es

,
.
B e thi s a s i t m a y h ow ev er i n su ch typ es of P r otozoa a s

, ,
the comp ou n d Ra di ol a ri a we fi nd th a t th ou gh th ere i s r ea son

,
t o reg a r d th e a gg reg a t e a s a n a gg reg a t e of th e s econ d ord er ,
yet its di vi s i bi l i tv i nto mi nor i n divid u a liti e s lik e th os e j u s t

d escr ib ed i s l es s m a ni fes t Fig 1 4 0 rep res enti ng S p hwr ozou m
,
. .
p u nc ta tu m ,
grou p ill u s t ra t es this T h e s ceptic
one of th e ,
.
a lly mi nd ed m a y p erh a ps d ou bt wh eth er we ca n reg a rd t h e

-
”
c el l aeform b odi es c ont a i ned i n i t as the m orphol ogi ca l
“
,
u nit s Of th e a nim a l T he j elly lik e m as s i n whi ch th ey a re

.
-
i m b edd ed is but i nd efi nit ely divisibl e i nt o p orti ons h avi ng

,
*
e a ch a c ell or n u cl eu s for its c ent re .A m ong the
Thi s sta tem ent s eem s at v ari ance wi th the figure ; but the figure i s v ery
i na ccura t e I ts i nacc uracy c urious l y i ll ustra tes the v i ti a tion of ev idence
. .
When I s aw the dra wi ng on the bl ock I pointe d out to th e draughts m an

, ,
THE MORP H OL OGICA L C OMP OSIT I ON OF A NIMA L S .
89
F ora m i ni fer a , we fi nd ly i n d e fi nite evid ence of the c oa l

on
es cen ce of a gg re g a t es Of th e fi r s t or d er i nt o a gg r eg at es of th e ,
s econ d or d er T h ere a re s oli ta ry F ora mi ni fe r s a lli ed to th e

.
,
c reatu re rep res ent e d i n Fig 1 3 4 Ce rt a i n id ea l t yp es of

. .
c om bi na ti on a m on g th e m are s h own i n Fig 1 41 ,

A nd . .
s etti ng ou t f rom th e se w e m ay a s cen d i n v ar i ou s di recti on s

,
to kin ds c omp ou n d e d to a n i m m ense v a r i ety o f d eg rees i n a n

imm ense v ar i ety of w a y s I n a l l of th e m h ow ev er th e
.
, ,
se p a r a bility of the m a j or i n divid u a li ty i nto m i nor i n di vi
d u a li ti es i s v ery i n com pl et e
,
T h e p orti on of s a r cod e con .
t a i ned i n one Of th ese c a lc areous c h a m b e rs giv es or igi n to ,
a n e xt ern a l bu d ; a n d thi s p r es e n tly b e c om e s c ov e re d lik e ,
i ts p a re nt w i th ca l careou s m a tter : the p os i ti on i n whi c h

,
ea ch s u cce s s iv e ch a m b e r i s s o p rod u c e d d etermi ni n g th e ,
f orm of th e c om p ou n d s h ell Bu t the p or ti on s of s a r c od e

.
thu s bu dd ed ou t one from a noth er do not b e com e di s ti nctly ,
i n divid u a li zed Fig 1 4 2 r ep resenti n g th e livi n g net wor k

. .
,
-
whi ch r em a i ns w h en th e sh ell Of a n Or bi toli te h a s b een dis

s olv e d sh ow s th e c onti nu ity th a t e xi sts a m on g th e occu p a nts
,
of i ts a gg r e g a te d c h a m b ers
*
I n the c omp ou n d .
I nfus ori a th e c om p onen t u nits rem a i n q u it e di s ti n ct

, Be i n g .
,
a s a gg r eg a t es of th e fi rs t or d er mu ch m ore d e fi nit ely or ,
g a ni z e d th e i r un i on i nt o a ggreg a tes of th e s econd or d er d oe s

,
th a t h e h a d m a de th e s ur ro unding c ur v es m u ch m ore ob vi o u sl y rel a te d to

th e co n ta i ned bo di e s , th a n th ey were i n t he Ori gi na l
’
F or
(in D r Ca rp ente r
. s
am i ni rcr a ) ; a nd ha v ing l oo ked on wh il e he i n g re a t m e a s u re rem edi e d thi s

de fec t, t ho ugh t no fu rt h e r c a re wa s ne e de d . Now howe v er,
, on s ee i ng th e
fi gure i n th e p ri nte r’s proof, I fi nd th a t the en ra v e r s wa ye d by t h e sa m e
g ,
s up pos i ti on a s t h e d ra u ht sm an th a t s uch a rel a ti on w a s m ea n t t o be s h o w n,

g
ha s m ade h i s l i ne s rep res ent it s ti l l m ore dec i dedl y t ha n t hos e Of the d ra ugh t
m an be fore th e y were co rrec t ed . Thus , v a gue l i nea r re p res en ta t i ons , l ik e
v a gue v erba l ones , a re a pt t o gr ow m o re defi ni t e wh en rep eat ed . Hypoth e s i s
wa rp s p erc e pt i ons as i t w a rps th oughts .
Th ough the s ubd i v i s i on i nto cha m be rs of the s hel l does not corr e s p ond
to th e v is ion i nto c el l uni ts i t m ay stil l be hel d tha t s i nce i n the sol i ta ry
s ubdi -
types the s ubdi v i s i on O f th e nucl eus i s fol l owed b form a ti on of ne w i ndi

y
yp e s t he s ubdi v i s i on of
vi dua l s wh i ch se a ra te and si nce i n th e com o und t
p , p
the nucl e u s i s fol l owe d by g ro wt h and form a ti on of ne w ch a m b ers th e com
,
pound type m us t be rega rde d a s a n a gg rega te of the s econd orde r .

not d estroy th ei r origi na l i ndivid u a liti es A m ong the Vor ti .
cel l az of w hi c h two ki n d s a r e d e li nea t e d i n Fig s

, 1 4 4 a nd .
1 4 5 t h er e a r e v a r i ou s ill u s tr a ti on s of thi s : the m emb ers of

,
th e com mu nity b e i ng som etim e s a pp en d e d to a s i ngl e s tem ;

s om eti m es a ttac h e d by l on g se p ara t e s t e m s t o a c omm on
b as e ; a nd som etim es m a sse d t og eth er

T hu s fa r th ese a gg reg a t es of th e s ec ond or d er exhibit bu t
,
i n d efinit e i n divid u a liti es T he i nteg r a ti on i s phy s i ca l ; bu t

.
n ot phy s i ol ogic a l T h ou gh i n th e P ol ycytha ri a th ere is a

.
, ,
s h a p e t h a t h as s om e s y m m et ry ; a n d th ou gh i n th e F ora ,
m i ni fer a the f orm a ti on o f su cc es s iv e c h a m b er s p roc eed s i n

, ,
s u ch m eth odi c wa y s a s to p r od u ce qu it e regu l ar a nd t ol er -
a bly s p eci fi c s h ell s ; yet no m or e i n th es e t h a n i n th e S p ong e s

-
or th e c om p ou n d Vor tt cel l w do we fi nd s u c h c o or di na ti on as
’
-
,
give s the wh ol e a li fe p re d om i na ti ng ov er the liv es of its

p a rt s We h a v e n ot yet rea ch ed a n a ggreg at e of the s ec ond
.
or d er s o i n divid u a te d a s to be c a p a bl e of ser vi n g a s a u n it i n
,
'
s till high er c ombi n a ti on s Bu t i n t h e cl a ss C ael en ter a ta this
.
,
a dv a n c e i s di s pl a y ed The comm on H ydra h a bit u a lly t a k en

.
,
a s th e typ e of th e l owes t divi s i on of thi s c l as s h a s sp eci a li ze d ,
p arts p er for m i ng mu tu a lly su bs ervi ent f u ncti on s and thu s

-
,
e xhibi t i n g a t ot a l li fe di s ti nc t f rom th e liv e s of the u n it s .
Fig 1 4 6 rep resent s one of th ese crea t u res i n i ts cont ra ct ed

.
s tat e a n d i n i ts exp an d ed sta t e ; whil e Fig 1 4 7 i s a di a gra m .
s h ow i n g th e w a ll of thi s c re a
’
t u re s s a c lik e b ody a s seen i n
-
s ec ti on u n d er the mic ros cop e
a a n d b b e i n g the ou t er a nd
i nner cellu l a r l ayers ; whil e be

t ween th em is the m esogl oea
”
or s t r u c t u r el es s l a m ell a th e ,
s u pp orti n g or sk el et a l l ayer Bu t thi s l owly org a n i ze d

.
-
ti ss u e of the H yd ra illu st ra t es a ph as e of i nt eg ra ti on
,
i n whi ch th e liv es of th e mi nor a gg re g a te s a re on ly p a r

t i a l l y su b ordi n at e d to th e lif e of th e m a j or a gg reg a t e
-
For a H ydr a s s u b st anc e i s s ep a r abl e

’
f orm ed by th em .
THE MOR P H OL OGI CA L COMP OS IT I ON OF A NIMA L S .
91
i nto A m oeba lik e p orti on s c a p a bl e of m ovi ng a b ou t i n d e

-
,
p end ent ly I f w e b ea r i n m i n d h ow a na l og ou s a re t he
.
e x tr em e e xt e n s ibili ty a n d c ont ra c t ili t y of a H ydr a s b ody

’
a n d t en tac l es to the p rop erti e s di s pl a y e d by th e sa r cod e

,
a m on g Rhi z op od s ; w e m a y i n fer t h a t p r ob a bly th e m ov e
m en t s a n d oth e r a c ti on s of a H y dr a a re du e to th e h a l f ,
i nd ep en d en t co Op er a ti on Of th e A m oeba lik e i n dividu al s

- -
c om p os i n g i t .
§ 2 02 . A
t ru th w hi ch we be for e s a w a m on g pl a nts we ,
h ere s ee rep ea ted a m on g a n i m a l s— th e t ru th th a t as s oon as

th e i n teg r a ti on of a gg re g a t es of th e fi r st or d er i n t o a gg reg a t es
o f th e secon d o r d er p ro d u ces com p ou n d wh ol es s o s p ec ifi c i n
,
th ei r s h ap es a n d s i z es a nd s o m u tu a lly d ep en d en t i n th ei r
,
p a rt s a s to h a ve di sti n ct i n divid u a li ti es th er e s i m u lt an eou sly

,
a r i se s th e t en d e n cy i n th e m t o p r od u c e by g e m m a ti on oth er , ,
su c h a gg r e g a te s o f th e secon d or d er T h e a pp r oac h t ow a r ds
.
d efi n it e li m it a ti on i n a n or g a ni sm i s by i m pli ca ti on a n a p
, , ,
p r oa ch t ow a r d s a s t a te i n w hi ch g rowth p a s s i n g a cert a i n
p oi nt r esu l ts not i n th e i n crea s e of the Old i n divid u a l bu t
, , ,
i n the f or m a ti on of a n ew i ndividu a l T hu s it h a pp en s th a t
.
th e c om m on p olyp e bu d s ou t oth er p olyp es s om e Of w hi c h ,
v er y sh ortly do th e lik e a s s h own i n Fig 1 4 8 : a p roces s

,
.
p ar a ll el e d by th e frond s of
s u n d ry A l g a e a n d by th os e
,
of th e l ow er J u ng er m a n n i
'
a c ea . A n d j u st a s a m on g ,
th ese l a s t pl a nt s th e p ro ,
l i ferou sl y p r od u ce d f ron d s
-
,
a fter gr owi ng t o cert a i n

s i z e s a n d d e v e l opi ng root
l ets d eta ch th em s e lv es fr om t h e i r p a ren t fron d s ; s o a m on g

,
th e se a ni m a l s s e p a r a ti on of t h e y ou n g on es fr om th e b odi e s
,
of th e i r p a r en t s e ns u es w h en th e y h a v e a cqu i r e d t ol er a bly
com pl et e or g a n i z a ti on s .
T h er e i s re a s on t o thi n k th a t th e p a ra ll e l h olds still fu r

t h er . Wi thi n th e
li m i ts of the J u ng er m a nni a c ece w e f ou nd ,
t h a t w hil e s om e g en era e x h ibit th i s di s con t i nu ou s d ev e l op

m ent oth er g en er a e xhibi t a d e v e l op m ent th a t i s s imil ar to
,
i t i n a l l e sse nti a l r e s p ects s a v e th a t i t i s c onti nu ou s

,
A nd .
h ere withi n th e li m i ts of the H ydrozoa we fi n d a l ong wi th , ,
thi s genu s i n w hi ch the gem mip a rou s i n divid u a l s a re p re

s entl y c a s t off oth er g en er a i n whi c h th e y a r e not c a s t of
,
f ,
bu t f orm a p erm a n ent a gg reg a t e o f t h e th i rd or d er Fig s . .
1 4 9 a nd 1 5 0 ex e mpli fy th ese c om p ou n d H ydr ozoa — one of

,
t h em s h owi n g t hi s m od e of g rowth s o ca r ri ed ou t a s t o p ro
d u ce a si ngl e a xi s ; an d the oth er s h owi ng how by r ep eti ,
ti ons of the p roc es s l a t era l a x es a re p ro d u ced I nt eg ra ti on s

,
.
ch a ra ct eri zi ng c erta i n high er genera

of th e H ydr ozoa w hi c h s wi m or fl oa t
i ns tea d of b ei ng fix ed a re i ndi cat ed ,
by Fig s 1 5 1 a nd 1 5 2 : the fi rs t of
.
th em rep res enti ng th e typ e of a

g rou p i n whi ch the p olyp e s g rowi ng
from an a xi s or coenosa r c a re d ra wn
, ,
th rou gh th e wa ter by the rhythm i

c a l c on t ra cti ons of th e org a n s f rom
whi ch th ey h a ng ; a nd th e secon d of
th em rep resenti ng a P hysa l i a the
c omp onen t p olyp e s of whi ch are
u ni te d i nt o a c l u s ter a tt ac h ed to a n
,
a i r v es se l -
.
A p a r all e l s eri es of ill u s t r ati ons might be d r a wn f rom th a t

s e cond divi s i o n of th e C wl c n tcr a ta k n own a s the A c ti nozoa
, .
H e re , t oo we h a v e a g rou p Of s p eci es — the S ea a ne m on es

,
-
th e i ndivid u a ls of w hi ch a r e s oli ta ry H ere too we h a v e

.
, ,
a g a m og enetic m u ltiplic a ti on : occ a si ona lly by g emm a ti on bu t

,
m ore f req u en tly by th a t m odifi e d p roces s c a ll e d s p ont a neou s
fi ss i on A nd h ere too we h ave c omp ou nd forms resu lti ng

.
, ,
fr om the a rrest of this sp ont aneou s fissi on b efore it i s com
pl ete To giv e ex a mpl es is n eedl ess ; sinc e th ey w ou ld but

.
s h ow i n m ore v a ri e d wa ys th e t ru th a l ready m a d e sufii

, ,
'
THE MORP H OL OGICA L COM P OS ITI ON OF A NIMA L S 9 3 .
'
c i entl y cl ea r th a t the comp ou nd Cael en tcra ta a r e a ggr ega t es
,
of th e t hi r d or d e r p rod u ce d by i nt e g ra ti on of a gg r e g a tes of
,
th e secon d or d er su c h a s w e h a v e i n th e H ydra A s b e fore .

,
it i s m a n i fest t h at on th e hyp oth e s i s Of ev olu ti on th es e ,
h igh er i nt eg r a ti ons w ill i ns en s ibly ar i se i f th e sep a r a ti on of ,
th e gem m ip a rou s p olyp es i s l on ger a n d l on ger p ost pon e d ;

a n d th a t a n i n cr e a s i n g p o s tp on em en t w ill res u lt by s u rviv a l
o f th e fit tes t i f it p r o fi ts t h e g r ou p of i n divid u a l s to r em a i n
,
u n it e d i ns t e a d of disp er s i n g
*
.
§ 2 03 . T he
lik e rel a ti on s exi st an d i m ply th a t the lik e ,
p rocess es h av e b een gone th rou gh a m ong th ose m ore highly ,
or g a n i z e d a n i m a ls c a ll e d P ol yzoa a n d T u ni ca ta We h a v e .
s olit a r y i n di vid u a l s a n d w e h a v e v a r i ou s ly i n t eg r a t e d g r ou ps
,
-
of i ndivid u a l s : th e c hi e f di ffe r en ce b et wee n th e e vid e n c e
h ere fu rni s h e d a nd th a t f u rni sh ed i n the l as t case b ei n g th e

, ,
a bs en c e of a typ e obvi ou s ly li n ki n g th e s olit a r y s t a t e w ith
the a gg r eg a t e d s t a t e .
T hi s i nt eg r ati on of a ggr eg a t es of th e se con d or d er i s c ar ,
r i e d on a m on g th e P ol yzoa i n div er s w a ys a n d w ith di fi e r ent ,
d egree s of com pl eten ess The littl e p a t ch es of m i nu t e ce ll s

.
,
sh own as m a gn ifi ed i n Fig 1 5 3 s o com m on on th e fron d s of .

,
s ea w ee d s a n d t h e su r fa c es of r ock s a t l ow w a t er m a r k di s pl a y
- -
,
littl e b eyon d m ech ani ca l combi na ti on T he a dj a cen t i n di .
A c ri ti c sa vs th e q ue s ti on is wh a t a re th e for c e s i nt e r n al or e x terna l
w hi ch p r odu ce u ni on or s e pa ra t i on .
”
x
A p ro i m a t e repl y i s —d egre e of nu t ri .
t i on . As in a p l ant ne w i n di i d ua l s v o r ru di m en ts of t h em a re c a s t O ff wh e re
nu t r i ti on is fa il i ng, s o i n a com pou nd a n i m al . Th e connect i ng p a rt dw i ndl e s
if it cea s es to c a rry nu t ri m ent .
vi du a l s , th ou gh s ev era lly or igi nat ed by gem m a ti on f rom the

s a m e g erm h a v e bu t littl e phys i ol ogi c a l d ep en d ence
,
In .
ki nd re d ki n d s h owev er as s h own i n Fig s 1 5 4 and 1 5 5 on e

, , .
,
of w hi c h i s a m a gn ifi e d p orti on of t h e oth er the i nt eg r a ti on ,
i s s om ewh a t g rea t er : the co op e r a ti on of th e u n i te d i ndi

-
vi du a l s b ei ng s h own i n th e p r od u c ti on of th os e t u b u l a r
b ran ch es w hi ch f or m th ei r com m on s u pp or t and es ta bli s h ,
a m on g t h e m a m or e d ec id e d c omm un ity Of nu t r iti on .
A m ong th e Ascidi a ns t hi s genera l l a w of m orph ol ogic a l

c om p os iti on i s on ce m or e di spl a y e d E ac h of th ese c r ea tu r es
.
s u b s i st s on the nu tr i tiv e p a rti c l es cont a i ne d i n th e w ater

whi c h i t d ra w s i n th rou gh on e or ifi ce a nd s end s ou t th rou gh
a n oth e r ; a n d it m a y th u s s u b s i s t e ith er a l one or i n con ,
n exi on w i t h oth er s th a t a r e i n s om e c a s e s l oosely a gg r eg a t e d
a nd i n oth er c a s e s cl os ely a gg reg a t ed Fig 1 5 6 P ha l l us i a

. .
,
m en tu l a , i s one of th e s olit ary f orms A typ e i n whi ch the .
i ndividu a l s a re u n i ted by a s tol e n th a t giv es origi n to th e m

by s u c cess iv e b u d s i s sh ow n i n P er op hora Fig 1 5 7 A m ong
, , . .
th e B otryl li dae Of which on e ki n d i s d r a wn on a s m a ll sca l e

,
i n Fig 1 5 9 a n d a p orti on of th e s a m e on a l a rger s ca l e i n

.
,
Fig 1 5 8 th ere i s a c om bi nati on of th e i ndividu a l s i nt o a n

.
,
nu l a r c l u s t ers w hi ch a r e th em s elv es i m b edd e d i n a c om m on

,
gel a ti n ou s m a t rix A n d i n thi s g rou p th ere a r e i nteg r ati ons

.
e v en a s t a ge high e r i n w hi c h se v e r a l s u ch c l u s te r s Of c l u s te r s
,
g row from a s i ngl e b ase H e r e the c om p ou n di ng a nd re

.
comp ou n di n g a pp e a r s to be c a rr i e d fu rth er th a n a ny wh er e
el s e i n th e a ni m a l ki n gd om .
T hu s fa r h owev e r a m on g th es e a gg reg at es of the thi r d

, ,
or d er w e s ee w h a t we b e for e sa w a m on g th e s i m pl e r a gg re
,
g a t es of the sec on d ord er— we see th a t th e c om p onen t i n di

vi du a l i ti es a r e bu t t o a v ery s m a ll e xt en t s u b or di n at ed to
T HE MORP H OL OGI CA L COMP OSI T I ON OF A NIMA L S .
95
the i n divid u a lity m a d e u p of th em I n nea r ly a l l th e for m s .
i n di cat ed th e mu t u a l d ep en d en ce O f the u n i ted a n i m a l s i s s o

,
s light th a t th ey a re m or e fi tl y c omp a ra bl e to s oc i eti es of

, ,
whi c h the m em b e r s c o Op e r a t e i n se cu r i n g certa i n c omm on

-
b ene fits T h ere i s s ca rce ly a ny s p eci a li za ti on of fu ncti ons

.
a m on g th e m On ly i n th e l ast typ e d escr ib e d do we see a

.
nu mb er of i n di vid u a l s s o c om pl et e ly c om bi ne d a s to s i m u l a t e
a s i ngl e i n divi d u a l A n d e v en h er e t h ou gh th ere a pp ea rs to

.
,
be a n i nti m a te c om m u n ity of nu tr i ti on th ere i s no phy s i o ,
l ogi ca l i ntegr ati on b ey ond th a t i m pli ed i h s ev era l m ou th s and

*
st om a ch s h a vi ng a com m on v ent .
2 04 . We
m e n ow to an e xt rem ely i nt er esti n g qu es ti on
co .
D oe s th er e e xi s t i n oth er su b ki n gd om s com p os iti on of the-
thi r d d e gree a n a l og ou s to th at whi c h w e h a v e fou n d so

,
p re v a l en t a m on g th e C ozl cn ter a ta an d th e P ol yzoa a n d T u ni

ca ta ? T he qu esti on i s n ot w h eth er e l sew h ere th er e a r e
t erti a r y a ggr eg a t es p r od u ced by th e b r an chi n g or c l u st eri n g
o f s e c on d a r y a ggr e g a te s i n w a y s lik e th os e a b ov e t ra c e d ;
,
bu t wh eth er e l sew h er e t h ere a r e a gg reg a t es w hich th ou gh ,
oth erw i s e u n lik e i n th e a r ra n g em ent of th e i r p a rt s nev er ,
th el es s c on s i s t of p a r t s s o s imil a r to on e a noth er th a t w e
m ay su s pe c t th em to be u n i te d secon d a r y a gg reg at es The .
v a r i ou s comp ou nd typ es a b ov e d es crib ed i n w hi ch th e u n it ed ,
a n i m a l s m a i nt a i n th e i r i n divid u a liti es s o di s ti n ctly th a t th e
i ndividu a lity of the a ggr eg at e r em a i ns v a gu e a r e con s t ru ct ed ,
i n su c h w a y s th a t th e u n it e d a ni m a l s c ar ry on th ei r s ev er a l
a cti vi ti e s w ith s c a r ce ly a ny m u t u a l hi n d r a n c e T h e m em b e r s .
of a b r a n ch e d H dr ozoon su c h a s i s s h ow n i n Fig 1 4 9 or
y , .
Fig 1 5 0 a re s o pl a ced th a t th ey c a n a l l s p rea d t h ei r t ent a cl es

.
,
a n d c a t c h th e i r p rey a s w e ll a s th ou gh s ep a r a te ly a t ta c h e d t o
s ton es or w ee d s P a ck e d s id e by s id e on a fl a t s u r fa ce or
.
*
I t h as b een poi nte d o t t h a t I h a v e h ere unde s ta t ed th e e v i denc e of
u r
phys i ol o gi ca l i nt eg ra ti on A n i ns ta nc e of i t a m ong Hydr ozo a i s sh ow n i n

.
v v
Fi g 1 5 1 , b u t by a s t ra n ge o e rs i gh t I h a e fo rgot t e n t o n a m e t h e a ri ou s
. v
ca s es furni sh ed b th e S i ho no ho r a i n wh i c h th e i ndi idu a l
y p p p ol yp e s of a c om v
pound aggrega te a re grea tl y sp eci a l i ze d in a da pta t i on to differe nt funct i ons .
for mi ng a t ree lik e

-
mbl age th e a ssoci a t ed i n divid u a ls
a sse ,
a m on g th e P ol yz oa a r e not u n equ a lly c on di ti one d : or i f on e
h a s som e a dv a nt a ge ov er a n oth er i n a p a r ti cu l ar c a s e th e ,
m od e of g r owth a n d the rel a ti ons to su rrou n di n g obj ects a re

s o i rr eg u l a r a s to p r ev en t thi s a dv an ta g e re a pp ea r i n g w i th-
c on s t a nc y i n s u cc ess iv e g e n er a t i ons S i m il a r ly w i th th e
.
A s cidi a ns g rowi ng f rom a stol e n or th ose for mi ng a n a nnu l a r

c l u s ter : e ach of th e m i s a s w e ll pl a ce d a s ev er y oth er for
d ra wi ng i n the cu rr ent s of s ea wat er f rom whi ch it s el ect s i ts

-
food . I n th ese cases th e m od e of a ggr eg at i on d oes not e xp os e

th e u ni te d i n divid u a l s t o m u lti for m c i r cu m s t a n ces ; a n d
th erefore i s not ca l cu l at e d to p r odu ce a m ong th e m any
st ru ct u ra l m u l ti for mity For th e s a m e rea s on no m ark e d
.
phys i ol ogi ca l divi s i on of l ab ou r a r i s es a m on g th em ; a nd

c ons equ ently no c om bi na ti on cl ose enou gh to di s gu i s e th ei r
s ev era l i ndividu a liti es Bu t u n d er c onv er s e con diti ons w e
.
m ay e xp ec t c on v er s e r e su lts I f th ere i s a m od e of i nt egr ati on

.
w hich n ece s s a r ily su bj ects th e u nit e d i ndivid u a l s to u nlik e

s ets of i ncid en t forces and d oes this w ith c ompl et e u ni for mity
,
f rom gen erati on to g enera ti on it is to be i n ferre d th at th e

,
u n it e d i n divid u a ls will b ec om e u nlik e T h ey w ill sev er ally

.
a ssu m e s u ch di ff er ent fu ncti on s a s th e i r di fferent p os iti on s
en a bl e th em resp ectiv ely to c ar ry on with th e great est

a dv a n t a g e to th e a ss em bl a g e T hi s h et erogenei ty of fu ncti on
.
a r i s i n g a m on g t h e m will be foll owe d by h et erogenei ty of

,
s t ru ctu re ; a s a l s o by th a t cl os er com bi nati on whi ch th e

’
b ett er en a bl es th em t o u tili ze one a noth er s functi ons A nd .
h en c e whil e the origi na lly lik e i ndividu a ls a re rend ere d

,
-
u n lik e th ey w ill h a v e th ei r h om ol ogi es f u rth er ob s cu red by

,
th ei r p rogr es si ng fu s i on i nt o a n a ggr eg a t e i ndivid u a l of a

high er or d er .
T h ese c onv erse c on diti on s a re i n n early al l c ases fu lfill e d

wh ere the su cces s iv e i ndivid u al s a r i s i n g by conti nu ou s d ev el
O m ent a r e s o b u dd ed Of
p f as to f or m a li n ear s er i es
-
I sa y .
i n n early a l l ca ses b e c au s e th ere a r e s om e typ es i n w hi ch

,
the a s s oci a t ed i ndivid u a ls th ou gh j oi ned i n s i ngl e fil e a re

, ,
TH E MOR P H OL OGICA L COMP OSI TI ON OF A NI MA L S .
97
n ot th ereby rend ere d v ery un lik e i n th ei r rel a ti ons to the

e n vi ron m ent ; a nd th ere fore do n ot b ec om e di f fer enti a t ed a n d
i nt eg ra t ed to a ny cons id era bl e ext ent I refer to su ch A sci
.
di a ns a s the S a l p i dce T h es e c rea tu res fl oa t p a ss iv ely i n the

.
s ea, a t ta ch e d t oge th er i n s t r i ng s . Be i n g pl a c ed s id e by s id e
a n d h a vi ng m ou th s a n d v e n t s th a t op en l a t e r a lly ea c h o f
,
th em i s a s we ll ci rcu m s t a n ce d a s i ts n e ighb ou rs for a b s or b

i ng a n d emit ti n g th e s u rrou n di ng wa t er ; nor h a v e th e i n
di vi du a l s a t th e tw o e xt r em iti es a ny m ark e d a dv a nt a ges
o v er th e res t i n th e s e res p ec t s . H enc e i n thi s typ e a n d i n,
the a lli e d typ e P yr os om a whi c h h a s its com p on en t i n di vi

,
d u a l s b ui lt i nt o a h oll ow cyli n d er li near a ggr eg ati on m a y

,
e xi s t w ith ou t th e m i nor i n divid u a liti es b e c om i n g ob s c u re d
a n d th e m a j or i n divid u a lity m a r k e d : th e c on diti ons u n d e r
w hi c h a di fferenti a ti on a n d i nt eg ra ti on of th e c om p onen t
i ndivid u al s m ay be e xp ected a re not fu lfill ed Bu t w h er e
,
.
the ch a i n of i ndivid u a l s p rod u ce d by g e mm a ti on i s eith e r ,
h a bitu a lly fix e d to som e s olid b ody by one of i ts ext rem iti es

or m ov es a ctiv e ly th rou gh th e w a t er or ov er s u b m er ge d
s t one s a nd w ee d s th e se v e r a l m e m b ers of th e c h a i n b ec om e
,
differently c onditi one d i n the way ab ov e d escr ib ed ; and m a v

th erefore be e xp ect e d to be com e u n lik e whil e th ey b ecom e
u nit e d . A cl ea r id e a of th e c ont r a s t b et w een th es e two
li near a rra nge m ent s a nd th ei r two div ers e r esu lt s w ill be ,
Obt ai ned by con s id er i ng wh at h a pp ens to a row of soldi ers ,
wh e n ch a ng e d from th e or di n a ry p os iti on of a s i n gl e rank
to t h e p os iti on of I n di a n fi l e . S o l on g a s th e m en st a nd
s h ou ld er to s h ou ld er th ey a re s ev era lly abl e to u s e th ei r
,
wea p ons i n lik e w a y s w i th lik e effi c i en cy ; a nd cou ld i f ,
c a ll e d on s imil ar ly p e r f or m v a r i ou s m a n u a l p rocess es di rec tly

,
or i n di r ec t ly con d u c iv e t o th e i r w e lf a r e B u t wh e n on the
.
,
“ ”
w or d o f c om m a n d right fa c e ,
th ey s o pl a c e th em s elv es
th a t ea ch h a s one of h i s neighb ou r s b e fore him and a noth er
b ehi n d h i m nea rly a l l of th em b ecom e i n c a p a cit at e d for
,
fighti ng and for m an y oth er a cti ons T h ey ca n wa lk or ru n

.
on e a fter a noth er s o a s to p r od u c e m ov em en t of th e fil e i n
,
53
the di recti on of i ts l ength ; bu t i f the fi l e h a s to Opp ose a n

enem y or re m ov e a n ob st a c l e lyi ng i n th e li ne o f i ts m a rch ,
th e fron t m a n i s th e only one a bl e to u se hi s w ea p ons or

h and s to mu ch pu rp os e A n d m anifes tly su ch a n a rrange
.
m ent cou ld b ecom e a dv ant ageou s only i f the front m a n p os

s es se d p ower s p ecu li a r ly a d a pt e d t o hi s p os iti o n w hil e th os e
,
b ehi nd him f acilit a t ed his a c ti ons by c a rryi ng s u ppli es &c , .
T h i s s i m i l e g rotes qu e a s it seem s serv es to conv ey b ett er

, ,
p erh a p s th a n a ny oth er cou ld do a c l ea r id ea of th e rel ati ons

,
th a t m u st a r i se i n a ch a i n of i n divid u a l s a r i s i ng by g em m a
ti on a nd c onti nui ng p erm a n ently un i te d en d to end S u ch
,
.
a c h a i n c a n a r i s e on ly on con diti on t h a t c om bi na ti on is m ore
a dv a n t a g eou s th a n se p ara ti on ; a n d for it to be m ore a dva n
ta geou s the a nt eri or m e mbe r s of th e s er i es m u s t b ec om e

,
a d a pt e d to fu n c ti ons fa c ilit a t ed by th e i r p os iti ons whil e th e ,
p os teri or m em b ers b ec om e a d a pt ed to fu nct i ons whi ch th ei r

p os iti ons p ermit H en ce di rect or i ndi rect equ ilib r ati on or
.
,
b oth m u st t end conti nu a lly to est abli s h typ es i n whi ch the

,
c onnec t e d i n divid u a ls a r e m ore a n d m or e u nlik e one a noth er ,
a t th e s a m e tim e th a t th e i r se v er a l i n divid u a liti es a r e m ore
a n d m ore di sg u i s e d by th e i n t eg r a ti on con s equ ent on th ei r
m u t u a l d ep end enc e .
S u ch b ei n g the a nticip a ti on s w a rr ant e d by the gener a l l aw s

of e v ol u ti on we h a v e now to i n qu i r e wh eth er th er e a r e a n
, y
a n im a l s whi c h f u lfil th em Ve ry littl e s ear ch su ffi ces ; for
.
s t ru ct u re s of th e ki n d to be exp ec te d a re a b u n d an t I n th a t .
g rea t divi s i on of the a ni m a l ki ngd om a t one ti m e c a ll ed A n

n u l os a bu t now gr ou p e d i n t o A nn eli da a n d A r th r op oda w e
, ,
fin d a v a ri ety Of typ e s h a vi n g th e l ook ed for ch ar a ct ers-

L a .
u s c ontempl a t e s om e of th e m .
2 05 .d u lt Ch aet op od is c om p osed of s egm ent s w hi ch

An a
rep ea t one a n oth er i n th ei r d et a ils a s w ell as i n th e i r g en era l
sh a p e s . D i ssecti n g one of th e l ower ord ers su ch a s is , .
sh own i n Fig 1 6 0 p rov es th a t the su c cess iv e s egm ents be

.
, ,
sid es h a vi ng lik e l oc om otiv e a pp end ages lik e b ranchi ae and

, ,
T HE MORP H OL OGICA L COMP OS IT ION OF A NIMA L S .
99
s om e t im es ve en lik e p a i rs of e ye s a l s o h a v e lik e i nt e rn a l
,
org a n s . E ach h a s i ts e n l a rge m en t of th e a li m en t a ry c a n a l ;

e a c h i ts c o n t ra c til e dil a t a ti on o f th e g rea t bl oo d v e s s e l
-
ea c h
i ts p orti on of the d ou bl e ne r vou s c or d w i th g a n gli a w h en

,
thes e exi s t ; ea ch i ts b ra nch es from th e n erv ou s a nd v a s cu l a r

t ru nk s a ns wer i n g to th ose of i ts ne ighb ou rs ; each i ts s i m i
l a rl y a nswer i n g set of m u s cl es ; ea ch i ts p a i r of Op en i ng s
th rou gh the b ody w a ll ; a n d s o on th rou gh ou t ev en to the
-
,
or g an s of rep rod u c ti on . T h a t i s to s a y ev er y s egm en t i s i n

,
re a t m e a su r e a phy s i ol o gic a l w h ol e — e v e r y s e gm ent c on

g
t a i ns m os t of th e org a ns ess enti a l to i n dividu a l li fe a n d m u l
ti pl i ca ti on : su c h es senti a l org an s a s it d oes not con t a i n ,
bein g th ose which i ts p os iti on a s one i n th e mid s t of a c h a i n ,
p rev ent s it f rom h a vi ng or needi ng . I f we

a s k wh a t is th e m ea n i n g o f th e s e h om ol ogi e s ,
n o a d e qu at e
a ns w er i s s u ppli e d by a n y c u rr e n t hyp oth e s i s .T h a t t h is

v egetative rep eti ti on i s carri e d ou t to f u lfil a p re d e
term i n e d pl an w a s s h own to be qu i te an u nten a bl e noti on
,
On the on e h a n d w e fou n d nothi n g s a ti s

,
fa ctor y i n th e concepti on of a Cr eator wh o p r escrib e d to h i m

s el f a cert a i n u n it of c om p os iti on for a l l c r ea tu r es of a p ar
ti cu l ar c l a s s an d th en di s pl a ye d h i s i n g enu ity i nb u ildi n g u p

,
a gre at v a r i e ty Of for m s with ou t d e p arti n g f rom th e a rch e

”
typ a l id ea . On th e oth er h a nd e x a mi n a ti on m a d e it m a n i
,
fes t th a t e v en w er e su ch a con ce pti on worthy of b ei n g ent er

ta i ned it wou ld h a v e t o be r eli n qu i sh e d ; s i n ce i n e a ch cl a ss
,
th ere a re nu m erou s d evi a ti ons f rom the su pp ose d a r ch etyp a l

”
id ea. S till l es s can th es e t ra its of s t ru ct u re be a cc ou nt e d
1 00 MORP H OL OGICA L D E VE L OP ME NT .
for t el eol ogic a lly T h a t cert a in orga ns of nu t riti on an d re

.
s pi r a ti on a n d l ocom ot i on a re r e p ea t e d i n ea ch se g m e nt of a
d orsib r a nchi a t e a nnelid m ay be re ga rd e d a s fun cti on all y a d

,
va ntag e ou s for a c r ea t u re foll ow i n g i ts m od e of li fe Bu t .
why s h ou ld th e re be a hu nd re d or ev en two h u n d r e d p a i r s of
o v ar i es ? T hi s i s a n a rr a ng em ent a t v a r i a nce w ith th a t
phy s i ol ogi ca l divi s i on of l a b ou r whi ch e v ery org ani s m pro
fi ts by— i s a l e s s a dv a nt a geou s a rr a ngem en t th a n m ight h a v e
b een a d opt ed T h a t i s t o s a y the hyp oth es i s o f a d es igned
.
,
a d a pt at i on fa il s to e xpl a i n th e f a c t s Contra r iwis e .

,
th es e s t ru ct u r a l t r a it s a r e j u s t su ch a s m ight n a t u ra lly be
l ook ed for i f th es e annu l ose f orm s h a v e a ri s en by th e i n
,
teg ra ti on of simpl er for m s A m on g the v a r i ou s c om p ou nd

.
a ni m a ls a l rea dy gl a n ce d a t it i s v e ry g e ner a l for the u n it e d

,
i n divid u a l s to rep eat one anoth er i n a l l th ei r p a rt s— rep ro

du cti ve org a n s i ncl u d e d H en c e i f i nst ea d of a cl u s tere d or
.
,
b ran ch e d i nt eg ra ti on su ch as the C ael en tera ta P ol yzoa and

, ,
T u ni ca ta e xhibit th er e occu r s a l on gitu di n a l i nt eg rati on ; we

,
m a y exp ect th a t th e u n it e d i n divid u a l s w ill h a bit u a lly i ndi
ca t e th e i r or igi na l i n d ep e n d en ce by s e v era lly b e a ri ng g er m
p rod u ci n g or s p er m p rod u c i n g org ans -

.
T he reasons for b eli evi ng one of th ese creatu res to be a n

a gg re g a t e of th e thi r d or d er a re gr ea tly s t r ength en ed wh en,
we t u rn f rom th e a d u lt s tru ct u r e t o the m od e of d e v el op

m ent A m on g th e D ors i br a nchi a ta a n d T u bi col ae the em
.
,
~
bryo l ea v e s th e eg g i n th e sh a p e of a c ili at e d ge mm u l e not ,
m u c h m ore di f fer enti a t e d th a n th a t of a p olyp e A s sh own .
i n Fig 1 6 2 it i s a n e a r ly gl ob u l ar m a ss ; a n d its i nt e r i or
.
,
con s ist s Of u nt r a n s for m e d c ell s Th e fi rs t a pp re ci a bl e ch an ge

.
i s a n el on g ati on a n d a sim u lt an eou s c om m encem ent of s eg

m ent a ti on T h e s egm ents m u ltiply by a m odifi ed gem ma
.
ti on whi ch t ak es pl a c e f rom th e hi nd er e nd of the p enu lti m at e

,
s egm ent A nd c ons id e r a bl e p rog res s i n m arki ng ou t th ese

.
divi s i ons i s m a d e b ef ore th e i ntern al org ani z ati on b egi ns .
Figs 1 6 3 1 6 4 1 6 5 r ep res ent s om e of th ese ea rly s t a ges I n

.
, , ,
.
a nn el ids of oth er ord ers th e emb ry o as s u m e s th e s e gm ent e d

,
THE MORP H OL OGICA L COMP OSITI ON OF A NI MAL S 1 01 .
fo r m whi l e s till i n the egg Bu t it d oes thi s i n j u s t the

.
s a m e m anner as b efore I nd eed th e es senti a l id en tity of the

.
,
two m od e s of d ev el opm en t i s s h own by th e f a ct th a t th e s eg

m ent ati on withi n the eg g is only p a rti a lly c ar r i e d ou t : i n
m s
al l typ es the s egm ents continu e to i nc rea se i n nu mb er

t h es e
for som e t i m e a fter h a tchi ng . N ow this p roces s i s a s
like th at by which com p ou n d a ni m als i n genera l a re f or m ed ,
a s th e di ffer en t c on di t i on s of th e c a s e p e r mit Wh en new

.
in divid u a l s a re b u dd ed ou t l at era lly thei r u n foldi ng is not

-
,
hi nd ered— th ere i s nothi ng to di sgu i s e eith er the p rocess or

the p rod u ct B u t ge mm ae p rod u c e d one f rom a noth er i n th e
.
s a m e st r a ight li ne ,
a nd r em a i n i ng c onn ect e d re s t r ict one
,
a n d th a t th e r esu lti n g s egm en ts a r e

’
a noth er s d ev el op m en t s
so m a n y g e m m ip a rou sly p r od u c e d i n divid u a l s

-
i s n ec es s ar ily
,
l ess Obvi ou s .
§ 2 06 E vid ence r em a i ns which a dd s v ery gr ea tly to the

.
weight of th a t a l r ea dy a ss ign ed T hu s fa r w e h a v e s tu di ed
.
only t h e i n divid u a l s egm en t e d a n im a l ; c on s id eri n g w h a t m a y
be i n fer re d from i ts m od e of ev ol u ti on a n d fi na l or g ani zati on .
We h a v e now to s tu dy s egm ent ed a n im a l s i n genera l C om .
p ari s on of di fferen t grou ps of th em a nd of ki nd s w ithi n each

gr ou p will di s cl os e v ar i ou s ph a ses of p rogressiv e i nt e gr ati on
,
of th e n a t u r e t o be a n ti c ip a t e d .
A m ong th e s i m pl er P l a tyh el m i n thes a s i n s om e ki n d s of

,
P l a na ri a t r ans v er se fi s s i on occu rs
, A p or ti on of a P l a na ri a
.
sep a r a t e d by s p ont aneou s con st r icti on b ecom es a n i nd e

,
1 02 MORP H OL OGI CA L D E VE L OP ME NT .
p end ent i n dividu a l S i r J G D al yel l fou n d th a t i n s om e

. . .
c as e s nu m e r ou s fr a g m ent s a rtifi c i a lly se p a r ated g re w i nto ,
*
p erfec t a n i m a ls I n th ese c rea t u res whi ch thu s r emi nd u s
.
of th e l ow es t H ydr ozoa i n th e i r p ow e r s of a g a m og en e ti c
mu l tipli cati on the i ndivid u a ls p rod u ce d one from a noth er

,
do not c onti nu e c onn ect e d A s th e y ou n g ones l at er a lly

.
bu dd ed off by the H ydra s ep a ra t e wh en c om pl ete so do th e

-
,
you ng on es l ongitu di na lly b u dd ed Off by th e P l a na ri a -

.
Fig 1 6 6 i ndi ca t es t hi s
. B u t th er e a r e a lli ed typ es whi ch
.
sh ow u s a m ore or l e ss p er si s t ent u n i on of h om ol og ou s p a r ts ,
or i n divid u a l s s i m il a r ly a r i s i ng by l ongitu di na l g e m m a ti on ) ;
,
T he c es t oi d E n tozoa f u rn i s h ill u st ra ti on s With ou t d welli ng .
on th e fa c t th a t ea ch se g m ent of a T aeni a lik e ea ch sep a r at e ,
P l a na ri a i s a n i n d ep en d en t h er m a ph r odit e ; a n d w ithou t
,
s p ecifyi ng th e s u n d ry c omm on s t ru c tu r a l t ra it s whi c h a dd
p rob ability to th e s u s pi c i on th at th ere i s s om e ki nship be

t w een th e i n dividu a ls of the one ord er and th e seg m ent s of
th e oth er ; it will s uffi c e to p oi nt ou t th a t t he two typ es a re
so far a lli e d a s to d em an d th ei r u n i on u n d er th e sa m e s ub
cl a s s titl e A nd rec ogn i zi ng this ki nship we see s ig nific anc e

.
,
i n th e f a ct th a t i n the on e c a se th e l on git u di n ally p rod u c ed -
gem m ae s ep ar at e as c ompl et e i n divid u al s a n d i n the oth er ,
c onti nu e u n i ted a s s egm en t s i n s m a ll er or l arger nu mb er s

a n d for sh ort er or l ong er p er i od s I n T a mia echi nococcus .
,
Rec entl y Mr T . H . Morgan h a s m a de el a bora te e xperim ent s wh i ch

sh o w tha t P l a na r i a M a m l a ta m ay be cu t i nto m any p i e ces from v a ri ous
pa rts a nd of v a ri ou s sh a p es— e v en a s l i ce out o f th e s i de — a nd e a ch, if not
too s m wi l l p rod uce a p erfect ani m al

al l , .
{ Si nce thi s wa s wr it ten i n 1 8 6 5 th ere h a s c om e t o l i gh t evi dence m o re

s ubd i v i s i on Of
c om pl et el y t o th e p oi nt th a n a ny a t t h a t ti m e k no wn In t h e .
P l a l yhel m i nthes k nown a s fl r bel l a ri a , th e re a re s o m e , the M erca tom i da

whi ch , by a p roce ss of s egm ent a ti on form ch a i ns of 4 , th en 8 , th en 1 6 , a nd
s om et i m es e v en 3 2 i ndi i du a l s v .
"
E a ch form s a m outh [l a tera l ] a nd for
s om e t i m e th e c h a i n pers i s ts , b u t th e i ndi v idu a l s ul t i m a t e l y b eco m e se x ua l l y
fce
”
m a t ur ed a nd th en s e p a r a te (Sh i pl ey, Z ool ogy
. o I n vertebra ta , p .
He re i t sh oul d b e rem a rk ed th a t th e l a t e ra l m out h s e n a b l e t h e m e m b er s of
a s t ri ng to fee d s ep ara tel y, a n d th a t nu t ri t i on not b ei ng inte rfe red wi th they

doub tl e s s g a in s om e v
a d antage by te m po rary m ai nte na nce o f th e i r uni on
p roba bl y i n cree pi ng .
T HE MORP H OL OGI CA L COMP OS I TION OF A NI MA L S 1 03 .
re p res ente d i n Fig 1 6 7 we h a v e a s p eci e s i n w hich th e

.
,
nu m b er of s egm en t s t hu s u nit ed d oes n ot e xc eed fou r In .
E c hi n obothri u m typ us th er e a r e eight or ten ; a n d i n c es t oid s

g ener a lly th ey a re nu m erou s *
A con s id er a bl e
.
hi a tu s oc cu r s b etween thi s ph a se of i nt egr a ti on a nd the n ex t

high er ph a se whi ch we m eet w i th ; bu t it i s not g re ater
th a n th e hi a t u s b et ween the typ e s Of th e P l a ty hel m i n th ea a n d
the C hwtop oda which p r esent the two ph a ses
, T h ou gh i t i s .
d ou btfu l wh eth er s ep arati on s i n gl e s e g m ent s occu rs a m on g
th e A nn el i da i yet v er y often w e fi n d s t r i ng s of s eg m en ts
, ,
I find tha t th e rea s ons for re a rdi ng th e s egm ent of a Taem a a s a ns weri ng
'
to a n i ndi v i dual Of the s econd ord er of a gg ega ti on a r e m uch s tronge r th a n r ,
I s up pos e d wh en w iti ng th e a b ov er Va n B en eden s a ys . L e P rogl otti s

(s egm ent) a yant a c q ui s tout s on d é v el op pe m ent se d é ta eh e ord ina i rem en t de ,
la col oni c et conti n u e en core a cr oi tr e

’
d a ns l inte sti n du m em e a ni m a l ; 11
ch an e
g m em e sou v ent de form e e t s em bl e d oué d
’
u n e nou v el l e v i e ; ses an
gl e s
’
s ef
fa cent, P l a na i re i l i eu
’
t out l e corp s s a rro ndi t , et il na ge com m e u ne au m
”
de s m u s cos i t é s i n t e s t i na l e s .
i Th ou gh th i s wa s doub tful i n 1 8 6 5 i t i s no l onger d oubtful . In an i ndi ‘
v i du a l ’
Ctenodr i l u s m onos l yl us , whi ch m ul ti p l i e s by di v i di ng a nd s u b di v idi ng
i ts el f, “
p a rt s a ri se whi ch a re de s t i tute of both h ea d a nd anu s a nd a t ti m es
”
C p a r da l es , t he re i s
'
con s i st of onl In a n o th e r s p e ci e s ,
y a s ingl e s e gm ent . .
'
se p ar at i on i n to m a ny s egm ent s ; a nd e a ch s e gm ent b efor e s ep ara ti ng form s
a budding zone o ut o f w hi c h oth e r s e m en ts a re a ft e r wa rd s p roduced , com

g
pl et ing th e a ni m al (Kors chel t an d He i de r, E m bryol ogy, i , 3 01
1 04 MORP H OL OGI CA L D E VE L O P ME NT .
i i g by r ep ea ted l ongi tu di n a l bu ddi ng whi ch a fter rea ch

ar s n ,
i ng c ert a i n l eng th s u n d e rgo s p onta n eou s fi ss i on : i n s om e

c a s es d oi n g this s o a s to for m two or m ore s i m il a r s t ri ngs
o f se gm en ts con s tit u ti n g i n d ep en d en t i n divid u a l s ; a nd i n
oth er c ases d oi ng i t s o th a t th e s eg m ent s s p ont aneou sly

s ep a r a t e d a r e bu t a s m a ll p a rt of th e s t r i n g T h u s a S yl l i s
.
,
Fig 1 6 8 a ft er rea chi ng a cer ta i n l eng th b egi ns to t r a n s

.
, ,
for m it s elf i nto two i n div i d u a l s : on e of th e p os ter i or segm ents

d ev el op s i nto a n i m p erfect h ea d a n d s i m u l tan eou s ly n arrows
,
i ts c onnexi on w i th th e p r ece di ng s eg m en t s f r om whi c h it ,
e v e ntu a lly s e p a r a t es S till m or e re m ark a bl e i s the e xtent to

.
whi c h t hi s p roce s s i s c arri e d i n cert a i n ki nd re d typ es ; w hi ch

e xhibit to u s s ev e r a l i n divid u a ls th u s b ei n g s i m u lt a neou s ly
for m ed ou t of g rou p s of s eg m en t s Fig 1 6 9 copi ed ( om i t

. .
,
t i n g the a pp end a g es ) from one c on t a i ned i n a m em oi r

“
by M Mil ne E d wa r d s rep res ents s i x worm s of di fferent

.
-
,
a g es i n cou rs e of d e v el op m en t : th e t e rm i n a l on e b e i ng th e
e ld es t ,
th e on e h avi ng th e g reat est n u m b er of s egm ents ,
a nd th e on e t h a t w ill fi r s t d et a ch it s e l f ; a nd th e s u ccess
i vel y a nt er i or on es w ith th ei r s u ccess iv ely s m a ll er nu mb ers

,
o f s egm ent s b ei ng s u ccess iv ely l es s a dv a n ced t ow a r d s fit ness

,
for s ep a r a ti on a n d i nd ep end en c e H er e a m on g g rou p s of

.
s eg m ents w e see r ep ea ted wh a t i n th e p r evi ou s c a ses occ u r s
w i th s i ngl e segm ent s A n d th en i n oth er a nnelid s w e fi nd th at

.
the st ri n g of segm ent s a ri s i ng by gem m a ti on from a s i ngl e

germ b ecom es a p er m a nen tly u ni ted wh ol e : the ten d ency to
a ny m or e c om pl ete fi s s i on th a n th a t whi c h m a rk s ou t th e seg
m ents b e i ng l os t ; or i n oth er w or d s the i nt eg r a t i on h a vi ng

, , ,
b ec om e r el ativ ely c om pl ete . L ea vi n g ou t of s igh t t he

qu esti on of alli an ce a m ong th e typ es a b ov e g rou p ed t ogeth er ,
th a t whi c h it h ere con cer ns u s to noti ce i s th a t l ongit u di n a l

,
gem m a ti on d oes g o on ; th a t i t i s di s pl a y e d i n th a t p ri m i tiv e

f orm i n whi ch the gemm ae sep a rat e a s s oon a s p rod u ce d ; th a t
we h a v e typ e s i n w hi c h s u c h g em m m h a ng t og eth e r i n
gr ou p s of fou r or i n g rou p s of eight and ten f rom whi ch
, ,
h owev er the g emm ae s u c c es s iv ely s ep arat e a s i ndivid ua l s ;

MORP H OL OGI CA L COMP OS IT ION OF A NI MA L S
.
TH E .
1 05
th a t a m ong higher typ es we h a v e l on g s t r in g s of s i m il a r ly

f or m ed g em m ee whi c h do n ot b ecom e i n divid u a lly i n depen d
ent , bu t s ep a r a t e i nt o or g a n i zed gr ou p s ; a n d th at from
th ese we a dv a nce to for m s i n whi c h a l l th e g e mm ae rem a i n
p a rts of a s i ngl e i n divid u a l One oth e r s ign ifi c a n t
.
f act mu st be a dd ed T h ere a re ca ses i n whi c h a n nelid s

.
m u ltiply by l a tera l g em m a ti on *
T h a t the l on git u di na lly .
p rodu ced g em m ze whi c h com p ose a n a nn elid sh ou ld th u s ,
h a v e one of them or sev er a l of th em the p ow er of l a t er a lly

, ,
bu ddi ng Off gem m ae f rom whi ch oth er a nnelid s a ri se giv es

-
, ,
fu rth er su pp ort to the hyp ot h es i s th a t p r im or di a lly th e seg , ,
m ent s we r e i n d ep en d en t i n divid u a l s A n d it su gges t s t h i s .
be li e f the m ore s t rongly b ecau s e i n c ert a i n typ es of Ga l en ,
tera ta we s ee th a t l ongit u di n a l a n d l a t era l g em m a ti on d o

,
. Oc cu r t ogeth er w h ere the l on git u di na lly u nit ed gem m m a re

,
-
d em on st r a bl y i nd ep end ent i ndivid u a l s .
§ 2 07 T h ou gh
it s eem s next to i m p oss ibl e th a t w e s h a ll
.
ev er be a bl e to fi n d a t yp e su ch a s th a t w hi c h i s h ere su p
p osed to be th e u n it of c omp os iti on of th e a n nu l ose typ e ,
si n c e we mu st a ssu m e s u c h a typ e to h a v e b ee n l ong s i n ce
e xti n c t
yet th e for egoi ng e vid enc e goes far t ow a r d s s h owi ng
,
th at an a nnu l ose a n i m a l i s a n a ggreg a t e of th e thi r d or d er .
T hi s rep et iti on of se gm ent s som eti m es nu m b e r i n g s ev er a l ,
hu n d red s lik e one a noth er i n a l l th ei r orga ns e v en d own to

,
th os e of r ep r od u cti on whil e i t i s oth erwi se u n a cc ou n ta bl e i s

, ,
fu lly a c cou n t e d for i f th ese se g m ent s a r e h om ol ogou s with

th e sep a r a t e i n divid u a l s of s om e l ower typ e The g em m a .
ti on by whi c h th es e s eg m en t s a r e p rodu ce d i s as s i m il a r a s ,
the conditi on s a ll ow to th e gem m a ti on by w hi c h com p ou n d

,
In pl a ce of th os e ori gi na l l y h e re i ns ta nce d a b ou t wh i ch th er e a re di s
put e s , I m a y gi v e a n un dou bt e d one desc ri b e d by McI ntos h, th e S y l l zs
’
r a m os a a s ec i e s of ch aeto od l i v i n
, p p g i n h e x a cti nel l i d s p onge s from th e A ra
fura Sea , wh i ch bra nch es l at e ra l l y re p ea t ed l y so a s to e xtend in al l di re ct i ons
th rou gh th e cana l s Of th e s p ong e . In m os t c a s e s th e b uds t erm i na t e i n o alv
segm e nt s wi th t wo l ong c i rr i e a ch . B ut m a l e a nd fem a l e bu ds we re found,
v
p ro i de d e ach wi th a h e a d, a nd c onta i ni ng o a ri e s a nd tes tes v . Som e ti m es
x
th e se se ual bu ds h ad b ec om e se p a ra te from th e b ra nch e d s to ck .
1 06 MOR P H OL OGICA L D E VE L OP ME NT .
i l i n g enera l a re p rodu ced A s a m ong pl ant s and a s

an m a s .
,
a m ong d e m on s t ra bly c om p ou n d a nim a l s we see th a t th e only

-
,
t hi ng req u i re d for th e for m a ti on of a p er m a nent c h a i n of

ge mmip a rou sly p rod u c ed i ndivid u a l s i s th at by rem a i ni ng
-
,
a ss oci a t e d su ch i n divid u a l s w ill h a v e a dv a n t a g es g rea ter th a n
ar e to be g a i ne d by s ep a r a ti on .F u r th er c omp aris on s of ,
the annu l oid a n d l ower a nn u l os e f or ms di scl ose a nu m b e r ,
o f th os e t ra n si ti ona l ph a s e s of i n t eg ra ti on which the hypo
th es i s l ea d s u s to e xp ec t A nd l a s tly th e differences a m ong

.
, ,
th es e u nite d i n divid u a l s or s u c c ess iv e s egm ents a re not ,
g rea t er th a n the differenc es i n th ei r p os iti on s and fu ncti ons

e xpl a i n— not g r ea t er th a n su ch diffe re nc es a r e k nown t o p ro
du c e a mong oth er u nit ed in divid u a l s : witness su n d ry com

p ou n d H ydrozoa .
I n di re ct evid enc e Of m u ch weight h a s still to be giv en .
T hu s far we h a v e c on s id ere d on ly th e l ess d ev el op ed A nnu -
l osa . T h e m ore i nt eg r at ed a n d m or e di fferenti a t e d typ es of

the cl as s rem a i n . I f i n th e m we fin d a c a rryi ng f u rth er of
th e p rocesses by which th e l ower typ es a re h ere su pp osed to
h a v e b een evolv ed we sh all h a ve a dditi ona l r eason for be
,
li evi n g th em to h a v e b een s o ev olved I f we fi nd th a t i n .
th es e su p eri or ord ers th e i ndivid u aliti es of th e u nit ed seg

,
m ent s a re mu ch l es s p ronou nce d th a n i n the i nferi or we ,
s h a ll h a v e g rou n d s for su s p ec ti n g th a t i n th e i n f er i or th e
i ndivid u a liti es of the s egm ent s a re l ess p ronou nce d th an i n

th ose l ost f or ms whi ch i niti at e d th e a nnu l ose s u b ki ngd om -
.
N O — P a rtly f rom the wi sh to i ncorp ora t e f u rther evi

[ T E
d ence and p artly f rom the w i s h to p res ent the evid ence ol d
, ,
a n d n ew i n a m ore effe ctiv e or d er I d ecid e h er e to rec a st th e

, ,
foregoi ng exp os iti on .
S ign ifi c a nt t ra it s of d e v el opm e nt are exhibit ed i n c omm on

by two g rou p s oth erwi se una lli ed— c ert a i n of th e P l a tyhel o
m i n th es a nd c ert a i n o f t h e l ow er A nn u l os a Of th e P l a tyhel .
m i n th es th e ordi n a ry typ e i s a n u ns egm ent ed c rea t u re a

TH E MORP H OL OGI CA L C OMP OSIT I ON OF A NIMA L S 1 07 .
P l a nar i an or a plifyi n g it A m ong the f ree

T rem a t od e e em x .
for m s as i n som e P l a n a ri ans th ere oc c u r s t r ans v ers e fi ss i on

, , ,
a nd p rompt s ep a ra ti on of th e s egm en ts ; whi l e a m on g s om e
oth er fr ee for m s a s the M i cr os tom i da the tw o se gm ent s fi rs t

, ,
p rod u ced th em selv es bec om e s egm ent e d w hil e still a dh erent

, ,
a n d thi s p r oc es s i s repe a t ed u ntil a st r i ng i s f or m e d A noth er .
gr ou p of the P l a tyh el m i n th es th e Ces toid E n tozoa exhibi t

, ,
a na l ogou s p r oc e s se s T h ere a re un s egm ent e d f or m s a s the

.
,
Ca ryOp hyl l a eus a nd th ere a re form s i n whi ch th e seg m ents

, ,
now few now m a ny a dh ere t og eth er i n ch a i ns ; th e te rmi na l

,
m e mb er s of which h owe v er ev entu a lly s e p ara t e and h a vi ng

, , ,
be for e s ep ara ti on app roa che d the t rem atod e s t ruc tu re b ecom e ,
i nd epen d en t i ndivid u a l s which gr ow c ree p a b ou t and con , ,
ti nne the race I n both of th ese typ e s th e c onditi on u nd er

.
whi ch the g em m ip a rou s ly p rodu c e d m em b er s rem a i n c on

-
nected i s th a t th ey sh a ll be a bl e to fee d in divid u a lly : i n

,
the one ca se by l a t er a l m ou ths i n th e oth er c a se by absorp ,
ti on th rou gh the i nt egu m ent I t i s fu rth er ob s erv abl e th a t

.
i n b oth c a s es s ep ar a ti on of th e c omp onent i n divid u a ls occ u rs

a t sex u a l m a t ur ity wh en a dv an t a g e i n n u t r iti on h as c ea s ed
"
to be th e d omina nt n ee d a nd di s p e rs i on of th e s p eci es h a s
ta k en i ts pl a ce i n d egr ee of import anc e A m on g .
A nnelid s high er th ou gh th ey are i n type we find p ar a ll el

, ,
i s m s Usu a lly i n i ts fi rst s t a ge an ann elid i s u ns egment ed

.
,
bu t a s fas t a s it el ong a t es l i n e s of se gm ent a ti on i n d en t i ts

s u r f a ce T hi s s egm ent a ti on p roc ee ds i n v a ri ou s w ays an d
.
,
the segm ent s exhibit v a r i ou s d e gr ees of d e p end ence In .
s om e l ow type s s pont aneou s fi s si on goes on to th e e xt ent Of

,
p rodu ci n g si n gl e s egm ents each of whi ch h as su ch vit a lity

,
th a t it b u ds ou t ant er i or a nd pos te r i or p arts a t i ts two e nd s .
T h u s a lik e i n the simpl e for m whi ch exi st s be for e se gm en t a

tion and i n the f orm e xhibit ed by a d et ach ed s egm ent we ,
h a v e a u nit a na l ogou s to ea ch of th e u nit s whi ch a re j oi ned

togeth er i n c ert a i n f ree T a r bel l a r ia a n d i n th e Ces t oid s : the
f
di fferen ce b ei ng th a t i n the A nnelid s th e s ex u ally m a t u re -
un it s do not i ndivi d u a lly di su nit e B ut th ou gh th ere d oe s

.
“
1 08 MORP H OL OGI CAL D E VE L OP ME NT .
not ta k e p ara ti on of si ngl e c ompl et ed s e gm ent s th ere

pl a ce se ,
t a k es pl a ce s e p ara ti on of gr ou p s of s egm ents whi c h a re ,
e ith e r s e x u a lly m a tu r e a t th e ti m e th ey d rop of f or p res ently

be c om e so A n d th e grou p s of s eg m ent s w hi ch h a v e b ecom e
.
s e x u a lly m a tu r e b efor e t h ey d rop of f h a v e s i m u lt an eou sly,
a cqu i re d s w i m m i n g or g a ns a n d d ev e l op e d ey es en a bli n g t h e m ,
to s p r ea d a n d di ffu s e th e s p ec i es S u n d ry bi ol o gi s t s r ec ogni ze
.
a p a r a ll eli s m b etwe en th a t d et ac h m ent of d e v el op e d seg m ents
w hi ch g oe s on i n th e c est oid E n tozoa a n d t h a t whi ch g oes on ,
i n th e S cyp hom edus ce T h e su cce ss iv ely d et a ch ed m em be rs

.
of th e s t r obil a a r e s e x u a lly m a t u r ed or m a tu r i n g i n divid u a l s

-
'
whi ch a s m e d u sae a r e fitt ed for s w im m i ng a b ou t m u l tiply

, , ,
i ng a nd rea chi n g oth er h a bit at s ; w hil e e a ch d et a ch ed p ro

,
gl otti s of the cestoid i s by the na tu re of i ts m ediu m li m i ted

, ,
t o c ree pi n g a b ou t C l ea rl y thi s fi ss i p a rou s p roces s i n s u ch

.
A nnelids a s th e S yl l i daz whi ch h a s s i m il arly b een c omp a red

,
to th e s t robi l iz a ti on of th e S cyp hom edus ce diff er s s imply i n ,
th e r es p ect th a t si ngl e s egm ents are not a d a pt ed for l ocom o

ti on and it th erefore p rofit s the sp eci es to s ep ara t e i n grou p s
,
.
Al l th ese f a ct s a n d a na l o gi es p oint t o th e c on cl us i on th a t th e
r em ot e a n c es t or of th e A nn elids wa s a n u ns e gm ent ed crea
t ure h om ol og ou s with ea ch of the se gm ent s of a n exi s ti ng

A nn elid .
T hi s c oncl u s i on i s su pp ort e d by oth er ki nds of evid enc e

h ere to be a dd ed T he l arv ae Of A nn elid s a re v ery v ari ou s ;
.
bu t a m id th e i r differen c es th ere i s a rec ogni z abl e typ e T he .
T roch oph or e is th e typic a l l a r v a l f or m of th e A nnelid s t e m

a t roch oph or e b ei n g a c u ri ou s s ph eroid a l cili at ed s t ru c t u r e
su gg estiv e of c oel ent er a t e a f fi niti es A nd this u nsegm ent e d

.
l a rv a rep r es enti n g the re m ot e a nc estor from whi ch the m any

,
A nn elid typ e s div erged i s simil a r to th e l a r v ae of th e Roti fera

,
a n d th e M ol l us ca : a t roch oph or e i s c om m on to al l th ese gr ea t
c l a ss es Moreov er si n ce a m on g the Rhi zota ( a su b cl a s s o f

.
,
-
th e Roti ferce ) th ere is a sp eci es T r ochosp hcera s olit a ry and

, , ,
free s wi mm i ng resembli n g i n for m a nd s t ru ct u re a t roch O

-
,
ph ore th ou gh it i s not a l arv a bu t an a d ult we get f urth er

, ,
T HE MORP H OL OGICA L COMP OSI T ION OF A NIMA L S 1 09 .
e vid ence th a t th ere wa s a p rimitiv e cr ea tu re of thi s g ener a l

ch a r a ct er,
of w hi c h th e t roc h oph or es O f M ol l us ca R oti fera , ,
a nd A nn el i da a re div erg ent m odifi c a ti on s a nd w hi c h w a s ,
u n se g m ented : the i m pli ca ti on b e i ng th a t th e s egm ent a t i on
or th e A n n el i da w as s u p er i n d u c ed T h a t t h i s s egm ent a ti on
.
resu lt e d fr om g e mm a ti on is impli e d by w h a t a r e c a ll e d p oly
t roch a l l a rvze T h ese s om eti m es a pp ear a s a st ag e s u cc ee d

.
i ng oth e r l a rv a l typ es T h u s th os e of A r eni c ol a m a ri na a ri s e

.
f ro m l a rv as w hi c h a t fi rs t w er e m onot roc h a l l a te r b eca m e ,
te l ot roc h a l a n d fi n a lly by th e a pp eara nce of n ew cili a te d

, ,
r i n g s b et ween th ose a l rea dy p r es en t a s s u m e d th e s t a g e of

,
p olyt roch a l l ar vae T hi s con diti on w a r ra nt s th e

.
a ss u mpti on th a t th e s eg m ent e d f or ms a r e to be l ook e d u p on
a s the y ou n g er th e u n se gm en t ed
,
on th e oth er h a n d , as th e ,
”
p hyl ogeneti ca lly old er ( Korschel t and H eid er i
. A nd , ,
th a t the a b ov e d e sc r ib ed r i ng s of c ili a m a r k Off s eg m en ts i s

-
s h ow n by t h e c a s e o f Op hryotr oc ha p u er i l i s w hich re m a i n s , ,
a s it w e r e ,
i n a l a rv a l c on diti on s i nc e the s egm ent s r et a i n
,
”
th ei r c ili a ti on th rou gh ou t li fe (I b . Y et one m ore
.
,
s ig ni fi c a n t fa ct m u s t be n a m ed I n ea r ly st a g es of d ev e l op
.
m en t ea ch segm ent of a n a r ch i an nel i da n h a s coel om i c s p a c e s
s epa r a t e f r o m th ose of n e ighbou r i n g s e g m ent s bu t i n th e ,

“
a d u lt th e s e pt a g en er a lly b r ea k d own e ith er p a rti a lly or
compl et ely ,
s o th a t th e p e r ivi s cer a l c a vity b e c om e s a c on
”
ti nu ous sp a c e f rom end to en d of th e a nim a l ( S edgwi c k .
,
T ex t B ook ,
Whil e thi s f a ct i s c ongru ou s w ith the
hyp oth es is h er e m a i nt a i n ed it i s i nc ong ru ou s with th e hyp o
,
th esi s th a t th e annelid wa s or igi na lly a n el ong a t e d c reat u r e

whi ch a ft erwa r d s b ec a m e se gm ent e d ; s i nc e i n th a t c a s e th e
i m pli c ati on w ou ld be th a t th e coel om i c s ept a not a r i s i n g from ,
r ec a pit u l a ti on of a n a n c est ra l s t ru c t u r e bu t or igi n a t e d by the ,
p roces s of s egm ent a ti on w ere fi rs t su perfl u ou sl y for m e d and

,
th en d est roye d .
Va r i ou s li nes Of evid enc e th u s c onv erge to th e c on cl u s i on

th a t an annu l os e a n im a l i s an a ggreg a t e of th e thi r d or d er .
I n Ju n e 1 8 6 5 wh en N O 1 4 of my s eri a l c ont a i ni ng th e
, ,
.
110 MO RP H OL OGICA L D E V E L OP ME NT .
f oreg oi ng ch a pt e r w a s i ssu ed I s u pp osed my self to be a l one

,
i n h oldi n g t hi s b e li e f res p ec ti ng th e a nnu l os e typ e a nd l o ng

,
conti nu e d to su pp os e so Ov er thi rty y ea rs l a ter h owev er i n

.
, ,
M E d m on d P err i er s work L a P hi Z OS Op hi e Z ool ogi qu e a va n t

.
’
,
D a r wi n I fou nd m enti on of a l ectu re d e liv ere d by M L a c a ze

, .
D u thi ers at th e E col e N or m a l e S u p é ri eu re i n P a r i s an d r e ,
p ort e d i n th e R evu e des Cou rs S ci enti fiqu es for J a nu a ry 2 8 ,
1 8 6 5 i n w hi c h he enu nc i a t ed a lik e b eli e f

,
. Ju dgi ng how ,
e v er by th e a ccoun t of thi s l e c tu re w hich M

,
P err i e r giv es
.
h p re sen t ) it pp r s th a t M L a c aze D u th i er s s imply

( e w a s ,
a ea .
-
c ont en d ed th a t thi s vi ew of th e a nn u l os e s tr u ct u re a s a r i s i n g
by u ni on of once i n d ep end ent u n i ts i s su ggeste d by cert a i n

-
,
a p r i or i c ons id er ati on s
. T h ere i s no i ndi c ati on th a t h e
a ssign e d a ny of th e cl a sse s of f a ct s a b ov e giv en which go to
,
s h ow th a t it ha s th u s a r is en .
F or fu rth er f a ct s a n d a r guments c onc ern ing the g enesis

o f the a nnu l os e type s ee A pp endi x D
,
CHA P T E R V .
TH E M O RP H O L O GI C A L C O M P O S I T IO N OF A N M A L S , I
C ON TI N UE D .
§ 208 . I N S E C T S A ra ch nid s C ru st ac eans

, , , a nd My ri a p od s ,
'
al l of
i‘
a re th a t high er divi s i on of the A n nu l osa
m em b e r s
c a ll e d A r ti cu l a ta or n ow m ore g ener a lly A r thr o oda T h ou gh
p .
i n thes e c rea t u r es the f orm a ti on of s egm ents m a y be i nt er
preted a s a di sgu i sed gem m ati on ; a n d th ou gh i n s om e of ,
th em the nu m be r Of segm ents i nc rea ses by thi s m odifi ed bu d

,
di ng a ft er l eavi ng the egg as it d oes a m ong the A n nelid s ;

,
yet the p roc es s i s not nea r ly s o d omi na nt : th e s egm en t s a r e

u sua lly mu ch l es s nu m erou s th an we fi nd th em i n the typ es
la s t con sid ered I n m os t c a s es too the s egm ents a re i n a
.
, ,
g rea t er d eg ree di ff erenti at ed one f rom anoth er at th e s a m e ,
tim e th a t th ey a re s ev e ra lly m ore differenti a te d withi n th em

selv e s N or is th ere a ny i nst an ce of s p ont a neou s fi ss i on
.
t aki ng pl a c e i n the s er i e s of s egm ents comp os i ng a n a rti cu

'
l at e an i m al On the cont r a r y the i nt e gr a ti on a l way s g reat

.
, ,
enou gh p e rm a nen tly to un it e th e s egm ent s i s f requ ently ,
c a r ri ed s o fa r a s to hid e v e ry c ompl et ely th e i n divid u a liti es
of s om e or m an y of th e m ; a n d o cca si on a lly as a m ong th e ,
A ca ri the con solid a ti on or th e a r rest of s egm ent a ti on i s s o

, , ,
Th e nam e A n n ul os a , once u sed to em br ace th e A nnel i da a nd A rthr o
p 0da , ha s of l a te ce a s ed to b e u s e d It s e em s to m . e b ett er th a n A pp e n
dwul a ta , both as be i ng m ore obvi ou sl y de sc ri pt i v e a nd as be ing m ore
x v
e cl u s i e .
1 12 MOR P H OL OGI CA L D E VE L OP ME NT .
d ecid e d as tol ea v e s ca rcely a tra ce o f the a rti cu l at e s t ru e

tu re : the typ e b ei ng i n th es e c a se s i n dic a t e d chi e fly by the
p resence of th ose ch a r a cteri sti ca lly for m ed li m b s whi ch giv e
-
,
t h e a lt ern a tiv e n a m e A r th r op oda t o a l l th e high er A n nu l os a .
Om it ti ng th e p a ra s i tic ord ers whi ch as i n oth er c a ses a re

, , ,
a b e rr a nt m em b er s of th e i r s u b ki ngd om
-
c om p a r i son s b e t ween
,
the di ffer en t or d e r s p rov e th a t the high er a re s t ron gly di s

ti ngu i sh ed f rom the l ow er by th e m u ch g rea ter d eg r ee i n
,
whi ch th e i n divid u ali ty of th e ter ti a r y a gg reg ate d om i na t es

o v er the i n divid u a li ti es of th os e sec on d a ry a gg r e g a t e s c a ll ed
“ ”
segm ent s or s o mit es ,
o f w hi c h it i s c om p os ed Th e s u c .
c es s i ve Fig s 1 7 0— 1 7 6 r ep re s ent i n g ( w ith ou t th ei r li m b s ) a

.
,
Ju l u s a S c ol op end ra an i sop od ou s C ru st a c ean and f ou r

, , ,
ki nd s of d eca p od ou s C ru s t a ce a ns endi ng wi th a Crab will

, ,
con v ey a t a gl a n c e a n id ea of th e w a y i n w hich th a t g r e a t er
s i z e a nd h et er o gen eity r ea c h ed by th e high er typ es i s a ec o m ,
n i d by a n i nt e g r a ti on w hi ch i n th e e xt r em e c a s es n e a rl y
p a e , ,
oblit er at es a l l t r a ce s of c omp os i te s t ru ct u re I n th e C ra b
.
the p ost eri or s egm en t s u s u a lly f old e d u nd ernea th th e sh ell

, ,
a l one p rese r v e th ei r p rimitiv e disti n c t n ess S o compl etely

.
c onfl uen t are th e rest th at it s eem s a b su r d to say th at a

,
C ra b s c a r a p ac e i s c omp os ed of a s m any segm ent s a s th ere a re

’
p a i rs of limb s f oot j a w s a nd a ntennas a tt ach ed to it ; and

,
-
,
w ere it not th at d u ri ng e ar ly st a g es of the Crab s d ev el op ’

T HE MORP H OL O GICA L COMP OSIT I ON OF A NIMA L S .
113
ment the g m en t a ti on i s fa i ntly m a rk ed th e a s serti on might

se ,
be con s id e re d ill egiti m a t e .
T h a t a l l a rt i cu l a t e a n i m a l s a re thu s c om po s ed f rom end to

end of h om ol og ou s s eg m ent s i s h owe v er a n a ccept e d doc
, , ,
tri n e a m on g n a tu ra li s t s . I t i s a d oc t r i ne th a t res t s on ca re fu l
ob s er v a t i on of th ree cl a ss e s of f a ct s —th e c orr es p on d e nce s
of p a r t s i n th e s u cc es s iv e s om it e s o f a n a d u lt a r ti cu l a t e
a ni m a l ; the s till m ore m a r k e d c orres p on d e nc es of s u c h p a r t s
a s th ey e xi s t i n th e e mb ry on ic or l a r v a l a r ti c u l a t e a n i m a l ;
a n d th e m a i nt en a n c e Of s u c h c or re s p on d enc es i n s om e typ es
whi ch a re a b sent i n typ es oth erwi se n ea r a ki n to th em .
The n atu re of the c on c l u s i on w hi c h th e s e e vid enc es u ni te i n

s u pp orti n g w ill b e st be sh ow n by th e a nne x e d c opi es from
,
the l ectu re di a g r a m s o f P rof H u xl ey ; e xhibi ti n g th e typi c a l

-
.
st ru ct u res o f a My ri a p od a n I ns ect a S pid er a nd a Cru s t

, , ,
~
a cean w i th th e i r r el a ti on s to a c omm on pl a n a s i nt er p ret e d

, ,
by h i m .
T rea ti n g of
th ese h om ol ogi es P rof H u xl ey s a ys th at a
, .
st r iki n g u n i for m ity of com p os iti on i s to be f ou n d i n th e h ea ds
of a t a n
, y rat e th e m ore highly or g a n i ze d m em b er s of th ese
,
fou r c l a sses ; an d th at typ ic a lly th e h ea d of a C ru st a cean

, , ,
a n A r a ch n id a My r i a p od
, or a n I nse c t i s c om p ose d of s i x
, ,
54
so mi tes ( or s e gm ent s corr es p ondi n g wi th those of the body )

and th ei r a pp en d a g es th e l a tt er b e i n g m odifi e d s o as t o s e rv e
,
the p u rp os e of s ensory an d m a nd u c a t ory org a n s ”

.
T h u s ev en i n the high er A r thr op oda th e m u ch gr ea t er c on ,
s olid a ti on a nd m u c h g rea t er h eterogene i ty do not oblit era te

a l l e vid enc e of th e fa ct th a t th e or g a n i s m i s a n a ggr eg a t e of
,
t h e thi r d or d er Com p a ris on s sh ow th a t it i s divi s ibl e i nt o a

.
nu mb er of p roxi m a t e u nit s e a ch of w hi c h i s a ki n i n c e rt a i n
,
f un d a m ent a l t ra it s to i ts n eighbou rs and each of whi ch i s a n ,
a gg reg a t e of th e s econ d or d er i n s o fa r a s it i s a n org a ni z ed ,
c om bi n ati on of th os e a ggreg at es of the fi rs t or d er w hi ch we

c a ll m orph ol ogi c a l u n it s or c ell s A n d th a t th ese s eg m ents .
or s om it e s w hi ch m a k e u p a n a nn u l os e a n im a l wer e origi n
, ,
a lly a gg re g a t e s o f t he s ec on d or d er h a vi n g i n d ep end en t i n
di vidu a l i ti es i s a n hypoth esis whi ch ga th er s fu rt h er s u pp ort

,
f rom the c ont ra st be tween th e high er a n d the l ower A rth ro

p od s a s w ell a s from the cont r a s t b et ween the A rth rop od s
,
“
Th e fu si on of th e se gm ent s form ing th e A rt h ropo d h ea d a nd th e
x
e trem e ch a nges , or pe rh a ps i n som e c as es di s a pp ea ra nce s , of th e ir a p pen
dag es , p ut g rea t di ffi cul ti e s i n the w a y of i de nti fica ti on ; s o th at th ere a re
differenc es of opi ni on res pe ct ing th e nu m b er of in cl uded s egm ent s P rof . .
Ma c Bride wri t es I t i s h i ghl y p rob a bl e t h a t a p ri m a ry h ea d ( p recors i

l obe orp rae s tom i um ) h as be en deri v ed from a nnel i d a nces tors , but t he
s econda ry fus i on of bo dy-s eg m e nt s w i th th i s h ea d, i n o t her w ords the form a
ti on of a s ec onda ry he a d, h a s g one on i ndep endentl y i n th e di ffe re nt cl as s e s

of th e phyl um A r throp oda , vi z , A r a chnida , Crus ta cea , a nd Tr a ckeata
.
( i ncl u di ng I ns ects a nd Myri a pods) . J dg d

u e by the num be r of a pp enda ge s
(whi ch gi v es a n i nfe rior l i m i t ) t he h ea d of a m a l a co s t ra co us Crus ta cea n

con s i s ts of p rass t om i um a nd 8 s e gm en ts ; th e h e a d of a n i ns ect of p raes to
m i um a nd 4 se gm ent s ; th e h ea d of a Myri apod of p raes t om i um a nd 3

se gm ents ; a nd the hea d of a n A ra chni d of
praes tom i um a nd 3 s egm ent s .
A ga in , th e com m ent of Mr . J . T . Cunni ngh a m i s A cco rdi ng t o Cl a us

a nd m os t m odern a u tho ri t i e s the re a re onl y 5 s egm ent s i n th e h e a d of a n
A rth ropod, th e eye s not count i ng as a pp endag e s ; a nd furth er i t sh oul d be

"
not ed th at th e s econd pa i r o f a nt ennae a re w a nti ng i n In s ect s .
Of cou rs e d ifference of o p i ni on re s p ecti ng th e num be r of s om i tes i n the

h ea d in v ol v es di fference of op i ni on r es p e ct i ng th e n u m be r con s t i tut i n g th e
enti re body, whi ch , i n th e h i gh er A rt hropod s , i s s a id by s om e to be 1 9 a nd

by oth ers 2 0 But those wh o thus differ i n det ai l , agree i n regarding a l l
.
th e se gm ents of hea d and b ody a s hom ol ogou s , a nd th i s i s th e e s s ent i a l p oi nt

w i th whi ch we a re he re conce rne d .
T HE MOR P H OL OGICA L COMP OSIT I ON OF A NI MA L S 1 1 5 .
in g enera l a nd the A nnelid s For if th a t m as ki ng of th e

.
i ndivid u a liti es of the segm ents which we fi nd di sti ng ui s h es

the hi gh er for m s f rom th e l ower h a s b een goi n g on f rom the
,
begi nni ng as we m ay f a i rly a ssu m e ; it i s to be i nfer re d th a t

,
the i ndivid u a liti es of th e s egm en t s i n th e l ower f orm s w ere ,
or igi n a lly m ore m ar k e d th an th e y n ow a re Rev ers i ng th os e

.
p r oc esses of ch ange by which th e m ost d ev e lop e d A nnu l osa

h a v e a ri sen f rom the l eas t d ev el op ed ; and a pplyi n g i n
th ou ght thi s re v er se d p roc ess to the l ea st d ev el op e d a s th ey ,
were d es cr ib e d i n the l a st C h a pt er ; we a re b rou ght t o the

conc e pti on of a tt a c h e d s e gm ent s th a t a r e a l l c ompl e t ely a lik e ,
an d h a v e th e i r i n divid u a liti e s i n no a pp reci a bl e d e g r ee s u b
or di n a t e d to th a t of the ch a i n th ey c omp os e F rom whi c h .
there i s bu t on e st e p to the c oncepti on of g e mmip arou s ly

‘
p rodu ced i ndivid u a l s which se v era lly p ar t one f rom a nother

a s soon a s th ey a r e for m e d .
§ 2 09 . We mu st now t
j u ncti on whence we
r e u rn to a
div erged som e ti m e a go A s b e fore e xpl a i ned u n d er the

.
h ea d of Cl ass ifi ca ti on or g an i s m s do n ot a dmit of u ni s er i a l
,
-
a rr a ng em en t e ith e r i n g ene r a l or i n d et a il ; bu t e v e r ywh ere

,
for m g rou p s w i thi n g rou p s H ence h a vi ng t race d the ph a s es

.
,
of m or ph ol ogic a l c omp os iti on u p to th e high e s t form s i n a ny
su b ki ngd om
-
w e fi nd ou rse lv es a t th e ext re mity of a g rea t
,
b ranch fr om whi ch there is no a cc ess to anoth er g rea t b ran ch

, ,
ex ce pt by g oi ng b a ck to s om e pl a c e of bif u r c a ti on l ow d own
i n the t ree .
’
T h er e exi s t s u c h simil a r iti es of sh a p e a n d s t ru ctu r e be
t ween th e l arv a l for m s of l ow Mollu s c s and th ose of A nnelid s
a nd Roti fers a s to sh ow th a t th e r e w a s a n ea r ly typ e c omm on
,
to th e m a l l ; and its p rob a bl e ch ara ct e rs s u gge st e d by com,
p ari son s eem to imply th a t it h a d ar i sen f rom s om e coel en

,
ter at e typ e i nt er m edi a t e be tween the C ni da ri a and th e C teno

,
phom Bu t th ere i s thi s n ot ew orthy di fferen c e b et ween th e

.
m ollu sc an l arv a and the a lli e d l a rv ae th a t it gives origi n to

,
only one a n i m a l a nd not to a g rou p of an im a l s u nit e d or ,

di su nit ed N o t ru e Moll u s c mu ltip l i es by gemma ti on ei th er

.
,
c onti nu ou s or di s conti nu ou s ; bu t th e p rod u c t of e v e ry ferti
l i z e d ger m i s a s i ngl e i ndivid u a l .
I t is a s i g nifi c ant f a ct th at h ere w h er e for the fi rst ti m e

,
we h a v e h om og enes i s h oldi ng th r ou gh ou t a n enti re su b ki n g -
dom we h a v e al so th rou gh ou t an enti re su b kin gd om n o c a se

,
-
i n w hich th e or g a nism i s divisibl e i nto two th r ee or m ore , , ,
lik e p a rt s T h ere is neithe r a ny su ch cl u steri ng or b r an c h

.
i ng a s a c oel ent era t e or m oll u scoid a n im a l u s u a lly di s pl a y s ;

n or i s t h er e a ny t r a c e of th a t seg m ent a ti on whi c h ch a ra c
ter i zes th e A n nu l os a . A m ong th ese a n i m a l s i n w hi ch no

s i n gl e eg g p rod u ce s s ev er a l i ndivid u als no i ndivid u a l i s
,
s ep a r a bl e i nt o s ev e r a l h om ol og ou s divi s i ons T hi s c onn e xi on

.
w ill be s een to h a v e a p rob a bl e m ea ni ng on rem e mb er i ng ,
th a t it is the con v er se of th e conn exi on which obt a i n s a m ong

the A n nu l osa con s id ere d a s a g rou p
,
.
A Moll u s c th en i s a n a gg r eg a t e of th e se c on d ord er
, ,
Not .
only i n th e a d u lt a n im a l i s t h ere n o s ign of a m u l tipli c ity o f
lik e p a rt s th at h a v e b ecom e ob scu red by i n t eg r a ti on ; bu t

th ere i s no s ign of su ch mu l tipli city i n the emb r yo A n d thi s .
u n i ty i s j u s t a s c on s pic u ou s i n th e l ow es t L a m ellib ra nch a s i n
th e high est C eph a l op od .
I t m ay be well to not e h owev er m ore es p eci a lly b ec a u se

, ,
it ill u st r at es a d ang er of mi si nt er p ret a ti on p r es ently to be

gu a rd ed a g a i nst th a t th ere a re cert a i n Moll u scs w hi c h s i m u
,
l at e the segm ent ed st ru ctu re E xt erna lly a C hi ton Fig 1 8 8

.
,
.
,
a pp ea rs to be m a d e u p of
divi s i ons su b s t anti a lly lik e

th os e of the c rea tu re Fig .
1 8 9 ; a n d one wh o j u dg e d
only by e xt e rn a l s wou ld s a y ,
th a t th e c rea t u re Fig 1 9 0 .
di ffe rs as m u ch f rom th e
cre a t u r e Fig 1 8 9 a s thi s .
,
d oes f rom the p rec e di ng one Bu t th e t ru th i s th a t w hil e

.
,
1 9 0 a n d 1 8 9 a re cl osely a lli ed typ es 1 8 9 di ff ers f ro m 1 8 8

-
,
THE MOR P H OL OGI CA L C OMP OSI T I ON OF A NIMA L S 1 1 7 .
m uc h m ore th an a m a n d oes f rom a fi sh A nd the

w id e ly .
ra di c a l di s ti n c ti on b et ween th e m i s thi s — Wh erea s i n t h e
C ru s ta cea n th e s e gm ent a ti on i s c a r ri e d t r an s v ers ely th rou gh

the wh ol e m ass of th e b ody so a s to rend er th e b ody m ore ,
or l ess c l e a r ly divi s ibl e i nt o a s er i es o f p a r t s w hich a r e s imi
l a rl y c om p os e d ; i n th e Moll u s c th e s egm ent a ti on i s limit e d

to th e sh ell c arri e d on i ts u pp er s u r fa ce a n d l e a v es i ts ,
b ody a s compl etely u n divid e d a s i s th a t of a c omm on s l u g *

.
Wer e th e b ody cu t th rou gh a t each of the divi s i ons the ,
s e ct i on of i t a tt a ch e d to e a c h p orti on of th e s h ell wou ld be
u n lik e a l l th e oth er s ec ti on s H er e the s eg m e nt a ti on h a s a

.
p u r ely fu ncti on a l d er iv ati on— i s a d a ptiv e i n st ea d of g eneti c .
The si m il a rly f or m e d a n d s i m il a r ly pl a c ed p a rt s a re not

- -
,
h om ol ogou s i n the s a m e s en s e a s a re the a pp end a ges of a

ph aenoga mic a xis or th e limb s of a n i n s ect .
§ 21 0 In. t dyi n g t
s u h e r em a i n i n g a n d high e s t s u b ki n g -
dom of a ni m a l s it i s imp or t a nt to r ecogn i z e thi s r a dic a l dif

,
f erenc e i n m ea ni ng b et ween th at lik en ess of p a rts whi ch i s

p rodu ced by lik eness of m odi fyi n g forces and th a t lik en ess ,
of p a rt s whi c h i s du e to p r i m or di a l id en ti ty of or igi n On .
our r e c ogn iti on of thi s di ffer en c e d ep en d s th e vi ew we t a k e
of c e rt a i n d oc t r i ne s th a t h a v e l ong b een d om i n a nt a n d h a v e ,
s till a w id e cu rrency .
A m on g th e Ver tebra ta a s a m on g the M ol l us ca h om og en e

, ,
s i s i s u niv ersa l T h e two su b ki ngd om s a re lik e one a noth er

.
-
a n d u nlik e th e r em a i ni n g su b ki n gd om s i n thi s th a t i n a l l
-
,
the typ es th ey s ev er a lly i ncl u d e a si ngl e fe rtili zed ov u m p ro ,
d u ces only a s i ngl e i ndivid u a l I t i s t ru e th at a s th e e ggs of

.
ce r t a i n g a st er op od s oc ca s i on a lly exhibit s p on t a neou s fi ss i on
P rof MacB ri de corr ects th i s s ta t em ent b y sa yi ng th a t The cteni di a or

.
“
gi l l s (whi ch i n M ol l us ca gen era l l y a re represented onl y by a s ingl e p a i r) a re

here rep res ent ed by a l a rge num ber of v
p a i rs ; th ey do not , h owe e r, c orr es pond
”
in e i th er num b er or p os i ti on t o t he s h el l pl a t e s. I t m a y, I thi nk , b e con
tended th a t if the s e h a d a ny m orp h ol ogi c al si gnifi ca nce , th ey woul d not diffe r
in arr angem ent from th e sh el l pl a te s , and woul d not b e l i m i ted t o thi s s p eci al
typ e of Mol l u s c .
of th e vit elli ne m ass whi ch m a y or m ay not resu l t i n the

,
form a ti on of tw o i n divid u a l s ; so a m on g v ert eb r at e a n i m a l s

we now a n d th en m eet w i th d ou bl e m ons t ers whi ch a pp e a r ,
to i m ply su c h a s p onta neou s fi s si on i m p er fectly c a rr i e d ou t .
Bu t th es e a n om a li es s er v e to r end er c on s pi c uou s the f a ct ,
th a t i n b oth th ese su b—ki n gd om s the n orm a l p roces s i s th e

i nteg r a ti on of the whol e ger m m a ss i nt o a s i ngl e org an i s m
-
,
whi c h a t n o ph ase of its d eve l opm ent di spl a y s any t en d ency

to s ep ara t e i n to two or m or e p art s .
E qu ally a s th rou gh ou t th e M ol l us ca th ere h old s th r ou gh ,
ou t th e Vcr te bm ta th e c orr e l a tiv e f a c t th a t n ot ev en i n its l ow

,
e st a n y m ore th a n i n its high es t typ es i s th e b ody divi sibl e ,
i nto h om ol ogou s seg m ents T he v ert eb r a t e a ni m a l u nd er its

.
,
si m pl es t a s u n d er i ts m os t c om pl ex f or m i s lik e the m ollu sc ,
ou s a n im a l i n t hi s th a t you c a nn ot cu t it i nto t ra ns v er se
,
sli ces ea ch of w hi ch c on t a i n s a digest iv e or g an a res pi r a tor y

, ,
or g a n a re p r od u c tiv e or g a n &c
,
T he orga n s of th e l ea s t
, .
d ev el op ed fi sh a s w e ll as thos e of the m os t d ev el op ed -
m am m a l for m bu t a si ngl e phy s i ol ogi c a l wh ol e ; a n d th ey

,
s h ow not th e r em ot es t t r a c e of h a vi n g ev e r b een divi s ibl e
i nt o two or m or e phy s i ol ogi ca l whol es T h a t s egm ent ati on

.
which th e v ert eb r a t e a nim a l u su a lly exhibits th rou gh ou t

p a rt of i ts org an i z a ti on i s the s a m e i n origi n and m eani ng
,
a s the s egm en t a ti on of a Chi ton s sh ell ; an d n o m or e impli es

’
i n th e v ert eb rat e a n i m a l a c omp os it e s t ru ct u re th a n do th e ,
su cce ss iv e p a i rs of bra nch i m of th e D a ta or th e t ra nsv ers e ,
r ows of b r an chi ae i n th e E ol is imply c omp osit e s t ru ctu re i n

,
th e m oll u sc ou s a nim a l T o s om e this will s eem a v ery qu es

.
ti on a bl e p rop os iti on ; a nd h a d we no evid en c e b ey on d th a t

which a d u lt v e rt eb r a t e a ni m a l s of d ev el op ed typ es su pply i t ,
w ou ld be a p rop os i ti on not ea sy to sub st anti a t e But a bu nd a nt .
s u pp ort for it i s to be fou nd i n the s t ru ct u re of the v ert eb ra t e

e m b ry o a nd i n the c omp a ra tiv e m or ph ol ogy of th e Ver tebr a te:
,
i n g en era l .
E mb ry ol ogi st s t ea ch u s th a t th e p ri m ordi a l rel a ti ons of

p art s a re m ost cl early di s pl a yed i n the early st a ges of evo
T HE MORP H OL OGICA L COMP OS I TI ON OF AN I MA L S 1 1 9 .
l u ti on ; a nd t h ey g enera lly b ecom e p a rti a lly or com

th a t
p l e t e l y di s gu i s e d i n i ts l a ter st a ges H enc e were th e v ert e

.
,
b ra t e a n i m a l on the s a m e l ev el a s the a nn u l ose a ni m a l i n

d eg ree of c ompos iti on— d i d it s imil arly c ons i s t of s eg m ent s
whi c h a re h om ol og ou s i n the s en s e th a t th ey a r e th e p rox
i m a te u n i ts of c om p os iti on ; we ou ght to fi nd thi s f u n d a
m ent a l fa c t m os t s t ron gly m a r k e d a t th e ou ts et A s in .
the a nnelid em b r yo the fi rs t con s pic u ou s ch a ng e i s th e

-
e l on g a ti on a n d divi s i on i n t o s e g m en t s by const ricti ons th a t

,
e n c i r c l e th e wh ol e bod y ; a n d a s i n th e a r th ropod e mb ry o
th e bl a st od erm be com es m a r k ed ou t t ra ns v er se ly in to pi e c e s
whi c h e xt en d th em s elv es r ou n d th e y elk be f ore th e i nte rn a l
or g a n i z a ti on h a s m a d e a n
y a pp rec i a bl e p rog r ess ; s o i n th e
e m b ry o of e v e r y v er t eb ra te a n im a l h a d i t an a n a l og ou s com
,
p os iti on t he fi rs t d ecid ed ch a n ge s h ou ld be a seg m ent a ti on

,
implica tin g the enti re m a ss B u t it i s not s o S und ry i m

. .
p orta n t di ffer enti ati on s occu r b efore a ny di vi si on s be g in to

s h ow t h e m se lv es T h ere i s th e d e fi ni n g of th a t e l on g a te d
.
,
el ev a t ed a rea with i t s l on gi tu di n a l g r oo v e w hi ch b ec o m e s t h e,
s ea t of su b s e qu e nt c h a n g es ; th e re i s th e for m ati on of th e
not och or d lyi n g b en e a th thi s g roov e ; th ere is th e g rowt h
u pw a r d s o f th e b ou n d a r i es o f th e g r oov e i n t o t he d or s a l
l a mi nae whi ch r a pidly d ev el op a nd fold over i n th e regi on o f

,
th e h ea d Ra thk e a s qu ot ed a nd i n d or se d by P rof H u xl ey
.
,
.
,
d escr ibe s the su bs eq u ent ch a nges a s follows T h e gel a ti n

ou s i n v es ti n g m a s s whi ch a t fi rs t s ee m s on ly to con stit u t e
, , ,
a b a nd to th e r ight a n d t o th e l eft of th e not oc h or d for ms
a r ou n d it i n th e fu rth er c ou r s e of d ev el opm ent a s h ea t h

, , ,
whi ch en d s i n a p oi nt p os t eri or ly A nt er i orly it send s ou t

.
,
two p rocesses whi c h u n d erli e th e l a t era l p art s of th e s k u ll ,
bu t v er y s oon c oa l esc e for a l on ger or s h ort er di s t a n c e .
P os t er i or ly th e s h ea th p roj ec t s bu t littl e b ey on d th e not o

,
c h or d ; bu t a n t e r i or ly for a c on s i d er a bl e di s t a n ce a s fa r a s th e
, , ,
i nfu n dibu lu m I t s en ds u p wa rds two pl at es whi c h em b r ace

.
,
the fu t u r e cen t r a l p a rt s of th e ner v ou s s y s te m l a tera lly p rob ,
”
a bly th r ou gh ou t th e i r enti r e l e n th T h t i s t o s a y i n th e
g a .
,
1 20 MORP HOL OGICAL D E VE L OP M E NT .
Vertebra ta the fi rst ste p i s th e ki n g ou t on th e bl a s t oderm

m ar
of a n i nt eg r at e d s t ru c t u r e w i thi n whi ch s eg m en ts su bse

,
qu ently a pp ea r Wh en th es e do a pp ea r th ey a re for s om e
.
,
tim e limit e d to the m iddl e regi on of the spin al a xi s ; an d n o

m or e th en th an ev er a ft er do th ey i m plic a t e the g ener a l
,
m a s s of th e b ody i n th ei r t r a n sv erse divi s i ons On the .
c on t r a ry b e fore v er t eb r a l segm en t a t i on h a s m a d e m u c h p r o
,
gre s s th e ru dim ents of the v a s cu l ar s ys tem a re l a id d own i n

,
a m a nn er sh ow i ng n o t r a c e of any p r i m ordi a l cor res p on d ence
of i ts p a rt s with th e divi s i on s of th e a xi s E qu a lly .
at v ar i a nce wi th the be li e f th a t th e v ert eb r a t e a n i m a l i s

es s enti a lly a s er i e s o f h om ol og ou s p a rt s i s t h e h et erog enei ty ,
which e xi st s a m on g th es e p a r ts on th e i r fi rs t a ppe a r a nce .
T h ou gh i n th e h ea d of a n a d u lt a rti cu l a t e a nim a l th ere i s

littl e s ign of divisibility i nt o s egm ent s lik e th ose of the
b ody ; yet su ch s egm ent s with th ei r a pp r op r i at e g a ngli a a nd
,
app en d a g es a r e e as ily id en tifi a bl e i n the a rti cu l a t e e mb r y o

, .
Bu t i n th e Ver tebra ta thi s a n tith es is i s rev er se d

. A t the .
tim e wh en s egm en t a ti on h a s b ec om e d ecid ed i n the d or s a l

r e gi on of th e spi ne th ere i s n o t r a c e of s e gm ent s i n th e p a rt s
,
which a r e to for m th e s ku ll— n othi ng wh at ev er to su gg es t th at

th e s ku ll i s b ei n g f or m e d ou t of divi s i ons h om ol ogou s with
v ert eb rae *
. A n d mi nu t e ob s er v ati on n o m ore di s cl os es a ny
”
s u ch h om ol ogy th a n d oes g ener a l a pp ea ra n c e Rem a k
“
.
,
s ays P r of H u xl ey
. h a s m ore f u lly p rov e d th a n a ny oth er
,
“
Ob s erv er th e s egm ent ati on i n t o u rw i rbel or p rot o v e rt eb rae

’
-
, , ,
whi ch i s ch aract er i s ti c of the v ert eb ra l c ol u m n st op s a t ,
the occipit a l m a r gi n of th e s kull— th e b a s e of w hi ch b ef ore ,
o ss ifi c a ti on p r esents no t ra ce of th a t s egm entati on whi ch

,
occu r s th rou gh ou t th e v ert eb ra l col u m n .
C on s id er n ext th e evid ence su ppli ed by comp a rativ e m or

p h o l ogy . I n p rec e di ng s ecti on s 2 06 it ha s b een ,
Th ough i t i s a l l eg e d th a t at a l a ter s tag e t he p osteri or p a rt of the s ku l l

is form e d by fus i on of di vi s i ons wh i ch a re a s s um e d t o repres ent v ert eb rae, ye t
i t i s a dm i t ted th a t th e a nteri or p a rt of th e s kul l nev er sh ow s a ny s i
gns of
s u ch di vi s i on More ov er i n bo th p a rts the bone s s h ow no tra ce

. of p ri m i ti ve
s e gm ent a ti on .
T HE MOR P H OL OGICA L C OMP OS I T I ON OF A NI MAL S .
1 21
s hown th a t a m ong a nn u l os e a n im a l s the divi s ibility i nto ,
h om ol ogou s p a rt s i s m ost cl ea r ly d e m ons t r abl e i n th e l owes t

typ es . T h ou gh i n d ec a p od ou s C ru st a cea ns i n I n s ect s i n , ,
A r a c hn id s th er e is diffi cu lty i n id en tifyi ng s om e or m an y of

,
the c omp onent som it es ; and th ou gh wh en id en tifi ed th ey , ,
displ ay on ly p a rti a l corr es p ond enc es ; yet on d escen di ng to

A nnelids the c omp os iti on of the en ti re b ody ou t of su ch
,
s omit e s b e c om e s c on s pi cu ou s a n d t h e h om ol ogy b et ween e a ch

,
s omit e a n d its n eighb ou rs i s sh own by th e re p eti ti on of one

’
a noth er s st ru ct u r a l d et a il s a s w ell a s by th e i r c om m on
,
g emmip arou s origin : i nd eed i n som e c ases w e h a v e the

,
homol ogy di rectly d em onst ra t e d by seei ng a somi te of th e

b ody t ra nsform ed i nt o a h ea d I f th en a v ert eb r at e ani m a l
.
, ,
h a d a s e gm ent a l co mp ositi on of ki n d re d na t u r e we ou ght t o ,
fi n d it m os t c l ear ly m a r k ed i n th e l owes t Ver tebr a ta a nd

m os t di s gu i se d i n th e high es t Ver tebra ta Bu t h ere a s b e fore
.
, ,
‘
the fa ct i s j u st the re v erse A m on g the Ver te bra ta of

.
d ev e lop ed typ e s u c h s egm en t a ti on a s really exist s rem a i ns

,
cons picu ou s— i s bu t littl e ob s cu re d e v en i n p a rt s of the s pi na l

c ol u m n f orm e d ou t of i nt e gr a te d v ert eb r ae Wh erea s i n the .
u n d e v e l op e d v ert eb r a t e t yp e s e gm ent ati on i s sc a rce ly a t a l l

,
t racea bl e !
T he A m p hi oxus Fig 1 9 1 i s not only with ou t
,
.
,
os s i fi e dv ert eb ra—3 ; not on ly i s it wi th ou t c a rtil a gi nou s r epre

s ent at i ve s o f th e m ; bu t it i s e v en wi th ou t a n yt h i n g lik e
di sti nct m em b ranou s divi s i on s T h e s pi n a l c ol u m n e xi sts

.
a s a cont i nu ou s notoch or d : th e on ly s igns of i n cipi ent s eg
m ent a t i on b ei n g giv en by its m emb r anou s s h ea th i n th e ,
u pp er p a rt o f whi c h qu a d r a t e m a ss es o f s om ewh a t d ens er

“
See not e a t th e end of th e ch apte r .

ti ssu e s eem f a i ntly to rep resent neu ra l s pi nes Moreov er .

,
th rou gh ou t s u nd ry grou p s of fi s h es a nd a mphibi a n s the ,
segm ent a ti on re m a i ns v ery imp er fect : on ly c ert a i n p eri

h e r al a pp en d ag es of th e v e rt e b ra b ecom i n g d efi n e d a nd
p
s olidifi ed w hil e i n pl a ce o f th e b odi e s of th e v e r t e b ra th e r e
,
s till c onti nu es th e u n divid ed not och or d T hu s i n stea d of

.
,
b ei ng m or ph ol ogi c ally comp ose d of v ert eb ra l segm ent s the ,
v ert eb ra te a nim a l i n i ts p r i m i tiv e for m is enti rely with ou t

v ert eb ra l segm ents ; a nd v ert eb r a l seg m ents b egi n to a pp ear
only as we a dv a nc e t ow a r d s d ev el op e d for m s Onc e .
m ore evid ence equ a lly a dv erse to the cu rrent hyp oth es i s
,
m eets u s on obs ervi n g th a t the di fferences b et wee n th e p art s

s u pp os ed to be h om ol og ou s a re a s g rea t a t fi rst a s at l a st
,
.
D i d th e v ert eb ra t e a n im a l p r im ordi a lly c ons i s t of h om o

l ogou s s egm en t s f rom s nou t to t a il ; th en th e s egm ents s a id
to c omp ose th e s ku ll ou ght i n th e l owes t Ver tebra ta to sh ow
, ,
th ems elv es m u ch m or e lik e the rem a i ni n g s egm ent s th an

th ey do i n the high es t Ver tebr a ta Bu t th ey do not
. Fi sh es .
h a v e cra ni a m a d e u p of b ones th a t are no m ore cl ea rly

a rr a ng ea bl e i nto s e gm ents lik e v e rt eb ra th an a re the c ra ni a l
“
b one s of the high est m am m al N ay i n d eed th e c a se i s

.
, ,
mu ch s t ronger . T he simpl es t fi sh p ossess i ng a s k el et on ,
h a s a c ra niu m c omp os ed of c a rtil a ge th at i s not s egm ent ed

a t al l
Bes id e s b ei ng i ncons i st ent wi th the l ea din g t ru ths of

E mb ry ol ogy a nd Comp a ra tiv e Morph ol ogy th e hyp oth e si s of ,
Goeth e a n d Ok en i s i n c on s i st ent wi th it self Th e f a ct s .
b rou gh t forw ard to sh ow th a t th ere exi s ts an a rch etyp a l

v ert eb ra and th a t the v ert eb rat e an i m a l i s c omp osed of
,
a r ch etyp a l v ert e b r a ar r an ged i n a s er i es a nd se v er a lly m odi

,
fi ed to fi t t h ei r p os iti on s — th es e f act s I say s o fa r f rom

, ,
p rovi ng a s m u ch s uffi ce wh en i m p a rti a lly con s id ered to di s

, , ,
p rov e i t N o a s s igned nor any con ceiv a bl e at trib u t e of the

.
, ,
s u pp ose d a r ch etyp a l v e rt eb r a i s u n ifo rmly m a i nt a i ne d T he .
p a rt s comp os i ng it a re c onst a nt neith er i n th ei r nu mbe r nor ,
i n th ei r rel a tiv e p ositi on s nor i n th ei r m od e s of os s ific ati on

, ,
T HE MOR P H OL OGICA L COMP OSIT I ON OF A NI MA L S 1 23 .
nor i n the s ep a ra te ness of the i r s ev era l i ndivid u a li ti es w hen

p res ent T h ere i s no fi xi ty of a ny one e l em ent or con
.
,
nexi on or m od e of d ev e l op m e nt
,
which j u stifi es ev en a ,
s u s pi c i on th a t v ert eb ra are m od ell ed a ft er an id ea l p a tt e r n .
T o s u b s t a nti a t e th es e a s s erti on s h ere w ou ld req u i re t oo m u c h

s p a ce ,
a nd a n a m ou n t of t ec h n ic a l d et a il w ea r i s om e to t h e
g enera l read er T he w a rran t for th e m will be fou n d i n a

.
c r iti ci s m on the os teol ogi ca l work s of P rof Owen or igi na lly .

,
p ubli s h e d i n the B ri ti s h a nd F or ei gn M edi co C hi ru rg i ca l R e -
vi ew for Oct 1 8 5 8 T hi s c r iticism I a dd i n th e A pp endi c es

. .
,
for the c on v eni en ce of th ose who m a y w i s h to s tu dy the

q u es ti on m ore fu lly ( S ee A pp endix B ) . .
E v er ythi ng th en goes to sh ow th a t the s e gm ent a l c omp o

, ,
s i ti on which ch a r a c t er is es th e a pp a ra t u s o f e xt er na l rel a ti on
i n m ost Ver tebra ta i s n ot p r im or di a l or genetic bu t fun cti on

, ,
a lly d et er mi ned or a d a ptiv e Ou r i n f erenc e m u st be th a t the

.
v ert eb ra t e anim a l i s a n a gg reg at e of the s ec on d or d er i n ,
which a r el a tiv ely su p erfi ci a l s egm ent a ti on h as b een p r o

du ced by m ech a ni c a l i nt ercou rs e wi th th e en vi ron m en t We .
s h a ll h erea ft er see th a t thi s c on ce pti on l ea d s u s t o a c on
s i s tent i nt er p re t a ti on o f th e f a cts —s h ow s u s why th er e h a s
a r i sen su ch u n i ty i n v a r i ety a s exi s ts i n e v ery v ert eb ra l c ol u m n ,
a n d why thi s u n ity i n v a r i ety is di s pl ay e d u n d er c ou ntl e ss

*
modifi ca ti ons i n different sk el et on s .
21 1 . On
gl a nci ng b ack at the f a ct s b rou ght t ogeth er i n
th ese two ch a p ters w e s ee it to be p r ob abl e th a t th ere h as gone
,
on a m on g an im a l s a p roc e s s lik e th a t whi c h we sa w rea s on
to think ha s gone on a m on g pl a nt s Mi nu t e a gg reg ates o f .
th ose phy s i ol ogic a l u n it s whi ch com p ose livi n g p rot opl a s m ,
A q u al ifyi ng fa ct shoul d be na m ed . Wh en th e p rod uct i on of v ertebral

segm ent s h a s be com e cons ti tuti ona l l y e s t ab l i shed, s o t h a t th ere i s an i nn a te
t endenc y t o form th em , th ere a ri se s a lia bi li ty to form su pe rnum e rary on es
a nd th i s , from ti m e to ti m e r ecu rri ng, m ay l ength en th e s eri e s , as i n th e

body of a s n ak e o r t h e ne ck of a _
s w an .
Thi s q u al ifi cati on ,
howe v er, af
fect s
q
e u al l y th e h ypoth es i s of an i dea l t yp e a nd th e hyp othe s i s o f m ech a ni ca l
ge nes i s .
I 24 MORP H OL OGICA L D E VE L OP ME NT .
e xis t a s P r otozoa : som e of th em i ncoh eren t i nd efi nit e and , ,
a lm ost h om og en eou s a n d oth ers of th em

, m or e c oh er ent de ,
fi ni t e and h et erogeneou s By u ni on of th es e nu cl eat ed p a rti

,
.
c l e s of s a r c od e a r e p r od u c ed v a ri ou s i n d e fi ni te a ggr e g a tes of
,
the s econ d ord er— S p ong es P ol ycytha ri a F or a m i n i fers &c

, , ,
.
i n whi ch th e comp ou nd i ndivid u a lity i s s c a r cely en ou gh

m a rk ed to su b ordi na t e the p r imitiv e i ndivid u a li ti es Bu t i n .
oth er typ e s ,
a s i n H ydr a th e liv es of the m or ph ol ogic a l
,
u n i ts a r e i n a c on s id e r a bl e d e gr e e th ou gh not wh olly m er g e d
, ,
i n the li fe of th e i nt eg ra t ed b ody th ey f or m A s th e p r i m ary

.
a gg reg a t e wh en it p a sses a c ert a i n s i ze u nd erg oes fi s s i on o r

, ,
gem m a ti on ; s o d oes the s ec on d ary a ggreg at e A n d a s on .
th e l ower st a ge s o on the high er we see c a s es i n w hi ch th e ,
g em mip arou sly p rod u ced i n dividu a l s p a rt as s oon as for m ed

-
,
a nd oth er c a ses i n w hi c h th ey c on ti nu e u n it e d th ou gh i n ,
g reat m easu re i n d ep end ent T hi s m a ss i ng of s econd ary aggr e

.
g at e s i nto t erti ary a ggreg at es is v a ri ou sly ca rri ed on a m ong

,
th e H ydr ozoa , th e A c ti nozoa , th e P ol yzoa , an d th e T u ni ca ta .
I n m os t of th e typ es so p rod u c ed , the c om p onen t i ndivid u

v ery littl e su b or di na t ed to the i ndivid u a li ty of th e
a l i ti es a r e
c omp os it e m as s— th er e i s on ly phy s i ca l u nity a n d n ot phy s i o
l ogic al u n ity ; bu t i n cert a i n of th e oceanic H ydrozoa the ,
i ndivid u al s ar e s o fa r diff erenti at ed and c ombi ned as v ery

mu ch to m as k th em F or m s sh owi ng u s cl early th e t ra ns i
.
ti on to w ell d ev el op ed i ndivid u al s of th e thi rd or d er are not

-
,
to be f ou nd . N ev er th el es s i n th e g rea t su b ki ngd om A nnu

,
-
l osa th e re a re t r a it s of s t ru ct u re d ev el opm ent an d m od e of

, , ,
mu l tipli ca ti on whi ch g o fa r to sh ow th a t i ts m e mb ers ar e

,
s u c h i ndivid u a l s of th e thi rd or d er ; a n d i n th e re l a ti ons t o
e xt er n a l con diti on s i n v olv ed by th e m o d e of u ni on w e fi n d ,
a n a d e qu a t e c a u s e for th a t ob s c u r a ti on of th e s ec ond a r y i ndi
vi du al i ti es whi ch we m u s t su pp os e h as t a k en pl a ce T he .
two oth er grea t su b divi si on s Mol l us ca a nd Ver tebra ta

-
, ,
b etween the l ower m emb ers of whi ch th ere a re s u gg es tiv e

p oi n t s of comm u nity p resent u s only with agg reg at es of th e
,
s econ d ord er th at h av e i n m any c ases b ec om e v ery l a rg e and

,
T HE MORP H OL OGICA L COMP OSITION OF A NI MA L S .
1 25
v ery com pl ex We fi nd i n the m no t ra ce of the u ni on of

.
g em mip a rou sly p rod u ced i n divid u a l s N ei th er th e m ollu s cou s

-
.
nor the v er t e b ra te a n im a l s h ow s th e f a i nt es t t rac e of a seg
m ent a ti on a fi ecti ng the t ot a lity of i ts s t ru ct u re ; a nd we s ee

good g roun ds for c oncl u di ng th at su ch s eg m ent a ti on a s ex
c epti on a l l y occu rs i n th e one a nd u s u a lly oc c u r s i n th e oth er
i s s u p er i n d u c e d .
[ N O TE — A q uesti on the st a t em ent on p 1 2 1

c ritic c a lls in .
r esp ecti n g the A m p hi oxus A t th e ou t set h owev er he

.
, ,
“
a dmi ts th a t i n th e A m p hi oz rus th e c ent ra l n er vou s s y st em
”
a nd th e not oc h or d a re not s e gm ent e d I n the A nnelid .
,
h owev er the c ent ra l ner v ou s s ys t em is s eg m ent ed a nd th ere

, ,
is s egm ent a ti on of the p a rt which as a su pp orti ng st ru ct u re , ,
is ana l ogou s to the n otoch or d i n r es p ect of f uncti on— th e

ou t er p art w hi ch r ep res ent s th e exo s k e l e t on i n c on t ra st to -
i
the end o s k el et on H e goes on to s a y th a t
.
“
th e gu t i s not
inv olv ed [ i n the s e gm ent a ti on ] and exhibit s i n A m p hi ox us
j u st a s it d oes i n w orm s differenti ati ons enti rely i nd ep en d ent
”
of th e s e gm en t a ti on of th e m es obl a s t P a rt of thi s s t a t e .
m ent is I thi nk not c ong ru ou s with al l th e fa cts I n P roto

, , .
dri tas one of th e l owest of the A r chia nn el ida “

, the i nt es ti ne ,
i s m on ili form th ere b ei ng a c ons t r icti on b et ween e a ch s eg

,
m ent a n d the ne xt ( S hipl ey ) Compl et e segm ent a ti on of

. .
th e i nt es ti n e i s Obvi ou s ly i m p os s ibl e s i nce were th e ca na l , ,
divid ed i nt o po rti ons by s ept a no food cou ld p ass B ut th e

,
.
f a ct th at th e gu t h a s thes e su ccess iv e exp ans i ons a nd con

st ri cti on s c orres p on di ng
, to th e s u c c e s siv e s eg m e nt s a n d ,
givi n g to eac h s egm ent a p a rti a lly s ep a ra t e s t om a ch s h ow s -

,
th a t seg m en ta ti on h a s g on e a s far as c on s i sts w ith th e

car ryi n g on o f th e liv e s of th e s egm en t s N o s u ch p arti a l .
s e m enta ti on e xi st s i n the A m h i T h th n th
g p ox us u s e r ee .
, ,
fu n d a m ent a l st ru ct u res — th e di re ctiv e s t ru c t u re th e su p ,
p ort i n g stru ctu re and the a li m ent ary st ru ctu re— a r e respec
,
ti vel y si m pl e i n th e l owes t v ert eb r at e and s egm ent e d or ,

1 26 MOR P H OL OGI CA L D E VE L OP ME NT .
p a rti ally s egm ent ed i n the l owes t A nn elid A g a i n whil e i t

, .
,
i s s a id t h a t th e g ill cl e fts e xhibit s egm ent ati on it i s a dmit ted

-
,
th at thi s h as no r el ev ance to a ny c ons titu ti ona l s eg m ent a

”
ti on : th ey a re s egment e d on a pl an of th ei r own i r re
s p ecti ve of oth er org an s A n oth er a ll eg a ti on i s th a t the
.
ov ar i es of A m p hi oxus a re s egm e n te d T h e i r s egm ent a ti on

.
,
h ow ev er lik e th a t of th e gill s i s i sol at ed and m ay be con

, , ,
s i dered a s ill u s t ra ti ng th os e rep e titi on s o f lik e p a rt s seen i n
s u p ernu m e r ary v e rt eb ra i n v a r i ou s c r ea t u r es— a rep etiti on

whi c h b ec om es h a bit u a l if th e resu lti n g s t ru ct u r e is a dva n
tageou s to th e s p eci es On th e st a t em ent th a t w hil e th e
.
A m p hi oxus h a s no ru di m ents of a r en a l s y st em th e E l as m o
b ra nch emb ry o ha s s u ch ru dim ents w hich a re a s di s ti nctly
,
s egm ente d a s th e ne ph r idi a of a wor m two c om m ents m a y

,
be m a d e . Th e fi rst i s th a t if i n th ese Vert eb r at es the

n e ph ridi a b ea r a r el a ti on to th e g enera l st ru ctu re lik e th a t
w hi ch th ey do i n A nnelid s th en one wou ld e xp ect to fi nd

,
th e s eg m ent a l a rra n gem ent sh own in the l owes t typ e a s i n ,
A nnelid s r a th er th a n i n a typ e c onsid era bly a dv a nc e d i n

,
d ev el opm ent S h ou ld it he r epli ed th a t i n the A m p hi ox us

.
a n exc r et ory sy st em h a d not yet a ri s en th ou gh one is re

,
qu i red for th e high er org ani zati on of a n E l asm ob ra nch then ,
the an sw er m ay be th a t s i n c e th e s eg m ent a l a r ra n gem ent i n

th e E l asm ob r an ch c orres p onds w ith th a t of th e my ot om es it ,
h a s no re fer enc e to a ny p r im ordi a l s egm en t a ti on s i nce the ,
my otom es h av e b een fu ncti onally gener a t ed The s ec ond .
c om m ent i s th a t w h erea s the neph r idi a of the A nnelid h a ve

i nd ep en d ent ext erna l op eni ng s the neph ridi a i n the E l a s m o
,
b r a nch h a v e not T h es e di sch a rge th ei r s ec reti ons i nto c er

.
t a i n g ener a l tu b es of exit comm on to th em a l l ; sh owi ng th at

ea ch of th em i ns tea d of b e i ng a m e mbe r of a p arti a lly i nd e
,
p end ent st ru ctu re is u nit ed with oth ers i n su b ordi nati on to

,
a g en er a l s t ru ct u r e T h a t is to s ay the s egm en t at i ons a re

.
,
far f rom b ei ng p ara ll el i n th ei r ess enti a l na t u res T h e a s ser .
ti on a ccomp anyi ng th es e c ritici s m s th a t th ere i s no di ffer

,
enc e i n pr i nci p l e b etw een th e s egm ent a ti on of A m p hi oxus

TH E MORP H OL OGICA L COMP OS IT ION OF A NIMA L S 1 2 7 .
a nd A nne lid is di ffi cu l t to il e w i th the vi s ibl e con

r ec onc
t ras t be tween th e two Wh at ev er l oc a l s egm enta ti ons th ere

.
a re i n a n A m p hi oxus a pp ear to m e q u it e u nlik e

“
i n p ri n
c i pl e to th os e w hi ch a n A nn e lid e xhibit s Cou ld i ts
.
p orti on of gu t be d u ly s u ppli e d with nu t r i m ent the segm ent ,
of a l ow A nne lid c ou ld c ar ry on i ts vit a l f u n c ti ons i nde pe nd
ently . I n th e p a rt s of th e A m p hi ox us w e s ee nothi n g
a pp roa chi n g to t hi s . Cu t it i nt o t ra ns v er s e se c t i on s a n d no
on e of th e m c ont a i ns a nythi n g lik e th e a sse m bl a g e of s t ru e
t u res requ i red for livi n g The A m p hi oxus i s a phy s i ol ogica l

.
w h ol e a n d i n th a t res p ect di ffer s ra di ca lly from th e A nnelid

, ,
e a c h s e gm en t of which i s i n chi e f m e a s u r e a phy s i ol ogi c a l
wh ol e . N o occ u rren ce of l oc a l se gm ent a ti on i n the A m
p h i ox us c a n oblit er a t e thi s fu n d a m ent a l c ont r a st .
A n a c com p a nyi ng c on t ra s t t ell s th e s a m e st ory On a s .
c endi n g f r om th e l ow e s t to the high es t a nnu l os e typ es w e

se e a p rog res si ng i nt e g ra ti on m or ph ol ogi c a l an d ph ys i ol og i
,
c a l ; s o th a t w h ere a s i n a l ow a nn e lid the s u ccess iv e p a r t s
a re i n l a r g e m e a su re i n d ep en d en t i n th e i r s t ru c t u r es a nd i n
th ei r liv es i n a high a rth rop od a s a c ra b m os t of th e p arts

, , ,
h av e l ost th ei r individ u a liti es and h a v e b ecom e m erg e d i n a

c onsolid a t e d org an i s m w ith a s i ngl e li fe ! u i te oth erwi se i s
.
i t i n th e v ert eb r ate s er i e s I ts l owest m emb er is a t the v ery

.
ou ts e t a co mpl et e m or ph ol ogic a l a n d phy s i ol o gi ca l w h ol e a n d ,
the f orm a ti on of th ose s er i a l p a rt s w hi c h s om e thi nk a na l o

gou s to th e s eri a l p a rt s of a n A nnelid b egi n s at a l a t er s t a ge
,
a n d b ec om e s gr a d u a lly p r onou nc e d T h a t i s to say the c ou rs e

.
,
of t ra nsf or m a ti on is rev erse d ]

C H A P T E R VI .
M O RP H O L O GI CA L I
D FFE RE N T A T I IO N IN P L A NTS .
21 2 . WH I L E ,
i n th e th ei r ev olu ti on pl a nt s a nd
c ou rs e o f ,
a n i m a l s h a v e di s pl a y e d p rog r e s s iv e i n t e g r a ti on s there h a v e ,
a t th e s a m e t i m e g on e on p ro g r ess iv e d i fi e r enti a ti on s of th e
r esu lti n g a gg r e g a t e s b oth a s wh ol es a nd i n th ei r p arts

,
.
T h es e di fferen ti a ti on s an d the i n te rp ret a ti ons of th em f orm ,
the sec on d c l a ss of m orph ol ogi ca l p r obl em s .
We com m ence a s b e f ore w i th pl a nt s We h a v e to con .
s id e r,
fi rst the s ev era l ki n d s of m odifi ca ti on i n sh a p e th ey
,
h a v e u n d erg one ; an d secon d the rel ati on s b etw een th ese

, ,
ki nd s of m odifi ca ti on and th ei r fact ors L et u s gl an ce at the .
l ea di n g qu esti on s t h a t h a v e to be a nsw ere d .
21 3 . I rresp ectiv e of th ei r d eg r ees of comp os iti on pl a nts ,
m ay, a nd do,
b ecom e ch an ged i n th e i r gen era l form s A re .
th ei r c h a ng es ca p a bl e of b ei ng for m u l at ed ? The i nqu i ry

whi ch m eet s u s a t th e ou t set is — d oes a pl a nt s s h a p e a d m it
’
o f b e i n g exp r ess e d i n any u niv ers a l t erm s —t er m s t h a t

r em a i n th e sa m e for a l l g en era or d ers a n d cl a ss es
, ,
.
A f ter pl ant s c on sid ere d a s wh ol es h a v e to be cons id er ed

,
th ei r p roxi m at e com p on ents whi ch v a ry with th ei r d eg rees

,
of c om p os i t i on a nd i n th e high es t pl a n t s a re wh a t we c a ll
,
b ranch es I s th ere a ny l a w t ra c ea bl e a m on g th e contrast ed

.
sh a p es of different b ran ch es i n the s a m e pl a nt ? D o the rel a

tiv e d ev el op m ent s of p a rt s i n th e s am e b ra nch confor m to
a ny l a w ? A n d a re th ese l aw s i f th ey exi st a lli e d wi th one
, ,
1 28
MORP H OL OGICA L D I FF E R E NTI A T I ON I N P L A NT S .
1 29
a not h er a nd w i th th a t to whi ch the s h a p e of the whol e pl ant

con for m s ?
D es c endi ng to th e c omp onents of th e s e c om p on ent s w hich ,
i n d ev e l op e d pl a nts w e di sti n gu i s h a s l ea v es th ere m eet u s

,
ki nd red q u e sti ons res p ect i ng the i r re l a tiv e s i zes th e i r r el a

,
tiv e s h a p es a n d th e i r sh a p es a s c omp a r e d with th ose of

,
foli ar orga ns i n g en er a l Of th e i r m or ph o l og ic a l di fi e r enti a

.
t i on s a l so i t h a s to be a s k ed w h eth er t h ey e x e mplify a ny
, ,
t ru th th at i s ex em plifie d by th e enti re pl a nt an d by its l arger

p a rt s.
T h en a s t e p l ower w
,
e c om e d own to th os e m orph ol ogi ca l
,
u nit s of w h i c h l ea v es a n d fr on d s co ns i s t ; a n d conce r n i n g
thes e a r is e p a r a ll e l i n qu i ri es t ou chi n g t h ei r div er g ences from

one a noth er a nd fr om c e ll s i n gen e r a l .
T h e p r obl em s th u s pu t t og eth er i n s ev er a l g rou p s c annot

of c ou r s e be r ig or ou s ly s ep a r a te d E v ol u ti on p res u pp os e s
.
t rans iti ons whi ch m a k e al l su c h cl a ss i ngs m ore or l es s con

venti on a l ; a n d a dh e re nc e to th e m m u st be s u bo rdi n a t e to
the n ee d s of th e occas i on .
21 4 . st u dyi ng the c a u ses of th e m orp h ol ogi ca l differ

In
enti a ti on s th u s divid e d ou t a n d p ros p ec t iv e ly g enera li ze d ,
we sh a ll h a v e t o b ear i n mi n d s e v er a l or d ers of f orc es whi c h

it will be well b ri efly t o s p ec i fy .
Growt h ten d s i n evi t a bly t o i niti at e ch a ng e s i n the s h a p e

of a n
y a gg reg a t e by a lt eri n g b oth the a m ou nt s of the i nc i
,
d en t f orces a n d th e for ces whi ch th e p a rt s ex ert on one

a n oth er . With the m ech ani ca l a cti ons thi s i s obvi ou s .
Matter th at i s s en s ibly pl a stic c a nn ot be i n creased i n m a s s

wi th ou t u n d erg oi n g a ch a ng e i n i t s p r op or ti on s cons eq u en t
,
on th e di m i n i sh e d r a ti o of i t s c oh e s iv e f or c e t o th e for c e of
gra vi ta ti on With the phys i ol ogi ca l acti on s i t i s eq u a lly

.
obvi ou s . I nc rea se of s i ze oth er t h i n g s equ a l a lt ers th e rel a

, ,
ti ons of th e p art s to the m a teri a l a n d dyn a m ic a l fa ct or s of

nu t ri t i on ; a n d by s o a ffecti n g di ffere nt ly th e n u t r iti on of
di fferent p a rts i niti at es fu rth er ch anges of p rop orti ons

,
.
55
In pl ants o f th e thi r d ord er it is thu s w ith th e p roxi m a t e

c omp onen t s : t h ey a re su bj ect to m u t u a l i n fl u ences th a t
a r e u n lik e on e a noth e r a n d a r e c on ti nu a lly c h a ngi n g T he .
e a r li e r for m e d u nit s b ecom e m ech a n i ca l su pp ort er s of th e

-
l ater for m e d u n i ts a nd s o e xp er i en c e m odi fyi n g f or ces from

-
,
whi c h th e l a te r form e d u n i ts a re e x em pt
-
F u rth er th es e
.
,
e ld er u n i ts s i m u l ta n e ou s ly b egi n to serv e a s c h a nne l s th r ou gh
whi ch m at er i a l s a re ca rr i e d to an d from th e y oun ger u n i ts

a n oth er c au s e of differen t i a t i on t h a t g oes on i n c rea s i n g i n i n
t ens ity On ce m ore t h ere a ri se ev er s tren gth en i n g cont ra s t s

.
,
-
b etween the a m ou nts of light whi ch f a ll u p on the you ngest or

ou ter m os t u ni ts a n d t h e e ld es t or i nner m os t u ni ts ; w h en c e
res u lt s t ru c t u ra l c ont r a s t s O f E vid en tly

yet a n oth er ki n d .
,
th en a l ong wi th th e p rogr ess iv e i n tegr a ti on of cell s i n to

,
f ron d s of fron d s i nto a x es an d of a x es i nt o pl a nt s s till m ore

, ,
c om p os i te th ere com e i n to pl a y su n d ry cau ses o f di fferen

,
t i a ti on whi ch a ct on th e wh ol e a n d on ea c h of i ts p arts ,
w h a tev er th e i r g r a d e T he forc es t o be ov er com e th e f orc es

.
,
t o be u tili ze d a n d th e m atte rs to be a pp rop r i a t ed do not

, ,
rem a i n th e s a m e i n t h e i r p r op or ti ons a n d m od es of a cti on for
a ny t wo m em b er s of th e a gg r eg a te : be th ey m e mb er s of th e
fi r s t s ec ond thi rd or a ny oth er ord er

, , , .
21 5 . th ese th e only ki nd s and cau ses of h et ero

N or a re
g e n e i ty w h i c h w e h av e t o c ons id er B ey on d
. t h e m or e
g ener a l ch an ges p r od u ced i n the rel ativ e s i zes a nd sh a p es of

pl ant s an d th ei r p a rts by p rog res s ive a ggreg a ti on th ere a re ,
the m ore p art ic u l a r c h a nges d eterm i ne d by th e m or e p a r

ti cu l ar c on diti on s .
P l a n ts as wh ol es a s su m e u nlik e a ttit u d es t owa r ds th ei r

envi ron m en t s ; th ey h a v e m an y w a y s of a r ti cu l ati n g th e i r
p art s w i th on e a noth er ; th ey h a v e m any w a ys of a dj u s ti n g
th ei r p a rt s t ow a rd s s u rrou ndi ng a gen ci es T h es e a re c a u ses
.
of s p e c i a l d i tter enti a ti ons a dditi on a l to th ose g ene r a l di ffer en
ti a ti on s th a t resu l t from i n c rease o f m a s s a nd i nc re ase of com

p ositi on I n e a ch p art con sid ered i ndivid u ally th ere a ri s es
. ,
MORP H OL OGI CA L D I F FE RE NTIA TI ON I N P L A NTS 1 3 1 .
a ch a r a ct eri sti c s h ap e c on s equ en t on th a t rel ativ e p os iti on

tow a r d s ex te rna l a nd i ntern a l f orc es w hich th e m od e of
,
g row th enta il s E v er y m e m b er of th e a ggr eg a t e p resents

.
i tself i n a m ore or l ess p ecu li ar w ay t owa r d s th e light t owa rd s ,
th e a i r a n d t ow a r d s its p oi n t o f s u pp ort ; a n d a cc ordi n g to

,
the re l a tiv e h om ogen e ity or h et eroge ne ity i n the i nc id en ce of

the a genc i es th u s b rou ght to b ear on i t w ill be the rel a tiv e
,
h om ogeneity or h eterogene i ty of i ts sh ap e .
Be f ore
p i g f r o m thi s a p r i ori Vi e w o f the
§21 6 . a ss n ,
m orph ol ogic a l di fferenti ati ons w hi ch necessa rily a cc omp an y

m orph ol ogi ca l i nteg r ati ons to a n a p os teri or i vi ew of th em it
, ,
s ee m s nee d fu l to s p ec i fy th e m ea n i ng s of c ert a i n d e sc r iptiv e
t erm s we sh a ll h av e to e mpl oy .
T a kin g for ou r b roa d es t divi s i on a m ong f or m s th e r egu l ar

,
an d th e i rr e gu l a r w e m a y divid e th e l atter i n t o th os e which

,
a r e w h olly i r re gu l ar a nd th os e w hi c h ,
b ei ng bu t p ar ti a lly
i rregu l ar su gge s t som e r egu l a r for m to whi ch th ey a pp roa ch
,
.
By s lightly s t ra i ni ng the differen c e b et ween th em tw o cu r ,
ren t wor d s m a y be con v eni e ntly u s e d to d esc r ib e t h es e su b
divisi on s T he enti rely i rr egu l ar forms we m ay cl a ss as

.
a s ym m etr i ca l — lit er a lly a s form s with ou t a ny e qu a liti es of

'
'
dim ens i on s T h e form s whi ch a pp roxi m a te t ow ar d s r egu

.
l a ri ty W i th ou t r e ac hi ng it w e m a y di s ti n gu i s h a s u ns ym m etri
,
ca l : a wor d which th ou gh it a ssert s i n eq u a lity of di m en s i on s

, ,
h as b een a ssoci a t e d by u s e r ath e r w ith su ch s light i nequ a lity

a s c on stit u t e s a n ob se r v a bl e d ep a r t u re f r om e qu a lity .
Of th e regu l a r for m s th er e a r e s ev e r a l c l a ss es di ffer i ng i n

,
the nu m b er of di rect i on s i n w hi c h equ a li ty of di m en s i on s i s

rep ea te d . H enc e resu lt s th e n ee d for n a m es by w hi c h sym
m et ry of s ev era l ki n d s m ay be e xp res sed .
T he m ost r egu l ar of figu re s i s the s ph er e : its dim en s i ons

a r e the sa m e fr om ce n t r e to s u r fa c e i n a l l di r e cti on s ; a n d if
cu t by a n
y pl a n e th r ou gh th e cent re th e se p a r at ed p a r t s a re
,
equ a l a n d s i m il a r T hi s i s a ki n d of s y m m e t ry wh i c h s t a nd s
.
a l on e an d w ill be h er ea ft er s p ok en o f a s s heri ca l s m m etr

, p y y,
Wh en asph ere p a s ses i nt o a s ph eroid either p rol a t e or

,
obl a t e th e r e re m a i n s bu t on e s et of pl a nes th a t w ill divid e it

,
i nto h a lv es whi ch a re i n a l l res p ect s a like ; na m ely the

, ,
pl a nes i n whi ch i ts a xi s li e s or whi ch h a v e i ts a xi s for th ei r

,
li ne of i n tersecti on P rol a t e a nd obl a te s ph eroid s m a y

.
s ev era lly p a ss i nto v a ri ou s f orm s wi thou t l os i ng thi s pro

p erty The p r ol a t e s ph eroid m a y b ecom e egg s h a p ed or py ri
.
-
for m and i t w ill s till con ti nu e c a p a bl e of be i n g divid e d i nto

,
two equ a l a n d s i m il ar p a rt s by a ny pl a ne c u tti n g i t d own

it s a xi s ; nor will the m a ki ng Of con str i cti on s d ep r iv e it of
this p rope rt y .S imi l a rly wi th the obl ate s ph eroid T he .
t ra ns iti on from a slight obl a t eness like th a t of a n or a n ge

, ,
t o a n o bl a t en es s r ed u ci n g it nea r ly to a fl a t di s c d oes not ,
a lt er its divi s ibility i n t o lik e h a lv e s by e v ery pl a n e p a ss i n g
th rou gh its a xis .A nd cl ea r ly th e m ou ldi ng of a ny su ch

fl att ened obl ate s ph eroid i nto the sh ap e of a pl at e l ea ves it ,
a s b e for e ,
sym m etric a lly divi s ibl e by a l l pl anes at right
a n gl es to its su r fa c e a n d p a s s i ng th rou gh its c ent r e
.
T his .
s pe c i es of s ym m et ry i s c a ll e d r a di a l s ym m etry I t is f a mili ar
.
l y ex em plifi ed i n s u ch fl owers a s th e d a isy th e t u lip and th e

‘
, ,
d a hli a .
F rom sph eri ca l sym m et ry i n which we h a v e an i nfi nit e

,
nu m be r of a x es th rou gh eac h o f w hi c h m ay p a ss a n i nfi nit e
nu m b er of pl a nes s ev er a lly dividi ng th e a ggr ega t e i nto equ a l

a n d si m il a r p a rt s ; a n d f rom r a di a l s ym m et ry i n whi c h w e
,
h a v e a si ngl e a xis th rou gh which m ay p ass a n i nfi nit e nu m ber

o f pl a n es s ev er a lly dividi n g th e a ggr eg a te i nt o eq u a l a nd
s i m il ar p a rt s ; we now t u r n to bi l a ter a l sym m e try i n whi c h ,
the divi sibili ty i nt o eq u a l a n d si m il a r p art s b ec om es m u ch

r est r ic te d. N oti n g for th e s a k e of c om pl et enes s th a t th ere is
, ,
'
a se x tu pl e bil a te r a l nes s i n th e c u b e a n d i ts der i va ti ve f orms
whi ch a d m it of divi s i on i nto e q u a l a nd s i m il ar p a rt s by pl a nes

p a ss i ng th rou gh the th ree di a gona l a x es an d by pl anes p a s si ng
t h rou gh th e th ree a x es th a t j oi n th e cent res of th e su r f a c es ,
l e t u s limit ou r a t tenti on to the th ree ki n d s of bil a t e ra l nes s

which h ere c onc ern u s . T he fi rst Of th ese i s tri p l e
MORP H OL OGICA L D IFFE RE NTIA T I ON I N P L A NT S 1 3 3 .
bi l a tera l s ym m e try . T his is the

sy m m et ry of a figu re h a vi n g
t h ree a x es a t right a n gl es to one a noth er th rou gh e ach of ,
w hich the r e p a sses a s i ngl e pl a ne th a t divid e s the a gg re g ate

i nto corres p ondi ng h a lv es A com m on b r i c k w ill s erv e a s an
.
e x a m pl e ; a n d o f obj e ct s n ot qu it e s o s i m pl e th e m os t f a mili a r ,
i s th a t m od er n ki n d of sp ect a c l e c a se w hi ch i s o pe n a t b oth
-
en d s T hi s m a y be divid e d i nt o corr es p on di ng h a lv es a l ong

.
i ts l ongit u di n a l a xi s by c u tti n g it th rou gh i n th e di re c ti on

of its thi c k nes s or by c u tti n g it th r ou gh i n th e di re cti on o f
,
i ts b rea d th ; or it m a y be divid e d i nt o c orres p ondi n g h a lv e s

by c u tti n g i t a cros s the middl e
'
Of obj ect s which

.
illu st rat e dou bl e bi l a tera l sym m etry m ay be na m e d on e of ,
t hose boats bu ilt for m ovi ng with equ a l f a c ili ty i n e ith er

di recti on a nd th erefore m a d e a lik e at stem a n d s t ern Ob
, ‘
.
vi ou sl y s u c h a b oa t i s s ep a r a bl e i nt o equ a l a n d s im il ar p a rts
by a vertic a l pl ane p a s s m g th rou gh s t em and s t ern ; and it is

a l s o s ep a r a bl e i nt o e qu a l an d s i m il a r p a r t s by a vertic a l pl a ne
bu tti ng it a mid s hips .

_
T o ex e m pli fy si ng l e bi l a tera l
s ym m etry it nee d s bu t to t u r n to the or di na ry b oa t of w hi c h
the two end s a re u n lik e H ere th ere rem a i n s bu t the one

.
pl a ne p a ss i ng v erti c a lly th rou gh st em a n d st ern on the oppo! ,
sit e s id es of w hi ch the p a rt s a re sym m et ri ca lly di s p os e d

'
T h ese s ev er a l ki nd s of sym m etry a s pl a c e d i n the fore goi n g

ord er imply i nc rea s in g h et erogeneity
,
The g re at est u ni.
f ormity i n sh a p e i s s h own by the divi s ibility i nt o lik e p a rts

i n a n i nfi n it e nu m b er of i n fi nit e se r i es of w a ys ; and th e
grea t est d egr e e of m u lti for mity c onsist ent with a ny re gu l a r ity ,
i s sh own by the divi s ibility i nt o lik e p a rts i n only a s i n gl e ,
wa y . H ence i n t r a c i ng u p org a n ic e v olu ti on a s di s pl a ye d i n

,
m orph ol ogi ca l di ffe renti a ti ons we m ay exp ect to pa s s from,
the one ext re m e of s ph er ic a l sy m m et ry to the oth er ext rem e ,
of si ngl e bil a t era l s ymm et r y T his exp ect ati on we sh all fi nd

.
to be c ompl et ely f u lfill ed .

CH A P T E R VI I .
TH E GE N E RA L S H AP E S OF P L AN T S .
§ 21 7 . p rot ophytes th os e e x emplifi ed by P l euro

A M ON G
coccus vu l ga r i s a re by g enera l c on s ent c on s id er e d th e s i m pl es t .
A s s h own i n Fig 1 t h ey a re gl ob u l a r c ell s p r esenti ng no

.
,
obvi ou s di ffe r enti a ti on s a v e t h a t be t wee n i nn er a nd ou ter
p a rt s T h ei r un i for mi ty of figu r e coe xi st s w ith a m od e of

.
lif e i nv olvi ng the u ni for m exp osu re of a l l thei r s id es to

i ncid en t forces For th ou gh ea ch i n divid u a l m ay h av e its
.
e xt er n a l p arts di fferently re l a t e d to en vi r on i n g a ge n c i es yet ,
th e n ew i ndivid u a l s p rod u ce d by s p on ta n eou s fi ss i on wh eth er ,
th ey p art comp an y or w h eth er th ey f or m c l u s ters an d a re

m a d e p olyh ed ra l by m u t u a l p ress u re h a v e no m ea ns of m a i n,
t a i ni ng p a ra ll el rel a ti ons of p os iti on a m on g th ei r p art s .
On the c ont r a ry the i n defi n i ten ess of the a tti tu d es i nto

,
which su ccess iv e genera ti on s f a ll m u st p rev ent the r i se of

,
a ny u nlik ene s s b etween one p orti on of the s u rfa c e a nd

a noth e r S ph eri ca l s ymm etry c onti nu es b ec a u s e on the
.
,
a v era g e of c ase s i n cid en t f or c e s a r e e q u a l i n a l l di r ecti on s

, .
Oth er or d ers of P r otop hyta h a v e m u ch m ore s p eci a l

f orm s a l ong w ith mu c h m ore s p ec i a l a tti tu d es : th ei r ho
,
m ol og ou s p a rts m a i nt a i n i ng f rom g enera ti on t o g ener a ti on

, ,
u n lik e re l a ti on s to i nc id en t f or c es T he D es m i di a cece a nd
.
D ia tom a cece o f whi ch Figs 2 a nd 3 s h ow ex a m pl es s ev era lly

,
.
,
i nclu d e g enera ch aract eri zed by t r ipl e bil a t era l sym m et ry .
A N a vi cu l a i s divi s ibl e i nt o c orres p on di n g h a lv e s by a t ra ns

T HE GE NE RA L S H A P E S OF P L A N T S .
1 35
v erse pl ane a nd by two l ongi tu di na l pl a n es —on e cu tti ng i ts

v a lve s a t r ight a ngl es a nd the oth er p a ss i n g b etw een i ts
v a lv es
. T he lik e i s t ru e
1
of th ose n u m er ou s t ra ns
v ers el y— con s tr i c te d f or m s of
D es m i dia c eae e x em plifi e d by

,
t he s econ d of th e i n divid u a ls
r ep re se nte d i n Fig 2 I f now we a sk how a N a vi cu l a i s
. .
r el a t ed t o i t s en vi ron m ent w e s ee t h a t i t s m o d e o f li fe ex
,
p os es it to th ree di fferen t s e t s of f orces : ea ch s et b ei ng

res olv a bl e i nt o tw o eq u a l a n d Opp os it e s et s A N a vi cu l a .
m oves i n the di recti on of i ts l en gth w i th e i th er en d foremost ,

.
H enc e on th e a v era g e i ts end s a re su bj ect to lik e a cti ons

, ,
from the a genc i es to w hi c h i ts m ot i on s su bj ect it F u rth er .

,
e i th e r en d w hil e m ovi n g e xp os es i ts r ight a n d l eft s id es t o
a m ou n t s of i n fl u ence w hi c h i n th e l on g r u n m u st be equ a l .
I f th en the two end s a r e not only lik e one a n oth er bu t h a v e

, , ,
corresp on di n g r ight a n d l e ft s id es th e s y m m e t r i c a l di st r i bu
,
ti on of p a rt s a nsw er s to the sy m m et ri ca l di str ibu ti on of

f orces P ass i n g to the t wo e dges an d the two fl at su rfa ces
.
,
w e s imil a rly fi n d a c l u e t o th ei r lik enes ses a nd di fferences i n

th ei r resp ectiv e r el a ti on s to t h e thi n gs a rou n d th e m T h es e .
l ocomotiv e p rot ophytes m ov e th rou gh th e en tan gl ed m asses

o f fr a g m e nt s a n d fib re s p rod u c e d by d eca yi n g org a n i s m s a nd
c on fe r v o id g r ow th s T h e i nter s ti c es i n su ch m a tt e d a cc u
.
m u l a ti on s a r e n ea r ly a l l of th em m u c h l ong e r i n one di m en
s i on th a n i n th e re s t— for m c r evi ces r a th er th a n re gu l a r
m esh e s H enc e a s m a ll or g a ni s m w ill h a v e m u ch gr ea t er

.
,
fa cility of i ns i nu a ti n g it s elf th rou gh thi s debr i s i n w hich it ,
fi nd s nu tr i m en t if its t rans v er se sec ti on i s fl a tt ene d i nstea d

,
o f s qu a r e or c i r c u l ar A nd whil e w e s ee h ow by su r viv a l Of
.
,
the fitt est a fl a tt en ed f or m is lik ely to be a cqu i re d by

,
di a tom s h a vi n g this h a bit ; we a ls o see th a t lik eness will be

m ai nta i n e d b etween the two fl at s u r f a ces an d b e tween th e
two e dges For on th e a v er a ge th e r e l ati ons of th e two fl at
.
, ,
su r f ac es to th e s id es o f the Op eni ng s th rou gh whi c h th e

1 36 MORP H OL OGICA L D E V E L OP ME NT .
di a tom p a ss es will be a like ; a nd so too on the a vera ge

, , , ,
w ill be the rel a ti on s of the two e dg es I n d es mids

.
o f th e typ e e x e mplifi e d by the s e c on d i n divid u a l i n Fig 2 .

,
a ki n d re d e q u a li za t i on of di m e ns i on s i s oth erw i s e 1 nsu r e d .
T h ere i s nothi n g to k eep on e of th e t wo s u r fa ces u pp e rm ost

ra th e r t h an th e oth e r ; a n d h en c e i n th e l ong s u c ce ss i on of
,
i ndividu a l s the two s u rfaces a re s u re to be s imil arly exp os ed

,
t o light a n d a gen ci es i n gener a l Wh en to thi s i s a dd e d th e

.
f a ct th a t s p ontaneou s fi s s i on occu r s t r a n sv ersely i n a c on st a nt

w ay it b ecom es m a nif e st th a t th e two end s w hil e th ey a re
, ,
m a i nt a i ne d i n c on diti on s lik e on e a n oth er a re m a i nt a i n e d i n

,
c on diti on s u n lik e t h os e of th e two e dg es H ere th en as .

,
b efore t ripl e bil a t era l s ym m et ry i n f or m coexi st s wi th a t ripl e

, ,
bil a t era l sym m etry i n th e a v era ge di str ibu ti on of a cti ons .
S till c onfi n i n g ou r a tt enti on to a gg r eg at es of th e fi rs t

or d er l et u s next not e wh a t re su lt s wh en th e two e n d s a re
,
p erm an ently s u bj ect to di fferent c onditi ons T he fix e d .
u n i ce ll u l ar pl a nt s of whi c h ex a mpl es a re giv en i n Fig s 4

‘
,
.
,
5 a n d 6 s ev era lly illu st ra t e the c ont ra st i n s h a p e a ri s i ng

, ,
b etween th e
p a rt th at i s a ppli ed to the su pp orti ng su rfa ce
a n d the p a r t th a t e xt end s i n t o th e s u rrou n di n g m edi u m .
T h ese two p a rts w hi c h a r e the m os t u nlik e i n th ei r r el a ti on s

to i ncid ent for c es a re th e m ost u nlik e i n the f orm s
, Ob s erv e .
,
n ext th at the p art whi c h li fts it self i nt o the w at er or a i r is

, ,
m ore or l ess d ecid edly ra di al E a ch out wa rd g rowi ng t u b u l e

.
of C odi u m a dhcer ens Fig 4 h a s its p arts disp os ed w ith s om e

, .
,
regu l a rity a rou nd its a xis ; the u pp er st em a n d s p ore v ess e l -

TH E G E NE R A L S H A P E S OF P L A NT S .
1 37
of B otrydi u m di s pl a y a l a t era l g row th th a t i s approxi

, Fig . 5,
m at ely eq u a l i n ev ery di recti on ; a n d the stem s of the .
Mu cor Fig 6 s h oot u p with an a pp roa c h to ev enn es s on a l l

,
.
,
sid es P l a nts of thi s l ow typ e a re nat u ra lly v ery v ar i abl e

.
i n t h e i r m od e s of g row th : ea ch i ndivid u a l be i ng g re a tly m odi

fi ed i n for m by its sp eci a l ci rc u mst ances Bu t th ey nevert he .
l es s s h ow us a gener a l lik enes s b etween p a rt s exp os e d to lik e

f orc es a s well a s a genera l u nlik eness bet ween p a rt s exp os ed
,
to u nlik e for c es .
Resp ecti ng th e form s of th ese a ggr e g at e s of the fi rst or d er ,
it h a s only to be a dd ed th a t th ey a re a s y m m et r i ca l whe re
th ere i s tota l i rr egu l a rity i n the i n cid enc e of f orc es We .
h ave an ex am pl e i n the i nd efi n it ely c ontort e d and b ranch ed

sh a p e of a f ungu s—ce ll growi ng as a my celi u m a mon g the ,
p articl es of s oil or th rou gh the i nt er stice s of org anic tis su e .
ill s t ra ti on s o f the g enera l t ru ths w hich t he

§ 21 8 u . Re -
f or m s of these v eget a l a gg reg a t es of the fi rs t ord er displ ay ,
a re fu r ni s h e d by v eg et a l a gg reg a t e s o f th e s e c on d or d er The .
e qu a liti es a nd i nequ a li t i e s of g row th i n di f fe r en t di re cti on s ,
p rov e to be s imil arly rel a t ed to th e equ ali ti e s a nd i n equ aliti e s

of en vi r oni ng a c ti on s i n di f fer ent di rec ti ons .
Of sph eric a l sy m m et ry a n i n st a nc e oc cu r s i n E u dor i na

e l ega ns T h e c ili a t e d c ell s a re h er e s o i m i ted a s to p r od u c e a
.
sm a ll mu lb erry sh a p ed h oll ow b a ll which b ei ng simil a r ly

,
-
, ,
c ondi ti one d on a l l s id es sh ow s no u nlik enesses of s t ru ctu re

, .
A n a lli e d f or m h owe v e r Vol vox g l oba tor p resent s a highly

, , ,
i nstru c tive th ou gh v ery t rifli ng m odifi cati on

, I t i s n ot
, .
a b sol u t ely h om og eneou s i n i ts s t ru ctu r e a n d i s n ot a bs ol u tely
hom og eneou s in its m oti ons T h e wa vi ng c ili a of its c om p o .
nen t ce ll s h a v e f a ll en i n t o su ch slight h et e rog ene iti es o f
a cti on as to c a u s e rot a ti on i n a con s t a nt di rect i on ; a n d
a l ong with a fix ed a xis of rot a ti on th e re h a s a r i s en a fix e d
a xi s Of p rog re ssi on A c onc omit ant f a ct i s th a t th e c ells

.
of th e c ol on y e xhibit an a i bl diff ti ti in l ti n
'
pp r ec a e e r en a on r e a o
to th e fix ed a xis T h ere is a n i ncipi ent div ergenc e f rom

‘
.
s ph er i ca l u n i form ity a l ong wi th thi s s light dive rg enc e f rom

u n i for m ity of c on di t i on s .
Veget a l a gg rega t es of th e s econ d ord er a re u s u a lly fix ed

l ocom oti on i s exc ept i ona l Fixi ty i m pli es th a t the su r f a c e
.
o f a tt a c hm en t i s di fferently ci r cu m s t a n ce d f rom th e f ree s u r
f a c e H ence we m ay exp ect to fi nd a s w e do fi nd th a t

.
, ,
a m on g th es e roo t ed a gg re g a t es o f t he s econ d ord er a s a m on g ,
th os e of the fi rs t or d er the p r i m ary cont r a st of sh ap e i s

,
b et ween th e a dh erent p ar t an d the l oose p a rt S ea weeds .

-
v ari ou s ly e x empli fy th i s I n som e th e fron d s a re v ery

.
i rregu l ar and i n som e t ol er a bly regu l a r ; i n s om e the f or m is

p seu do foli ar a nd i n s om e p seu d a xi a l ; bu t difi eri ng th ough
- -
they do i n th es e res p ects th ey a g ree i n h a vi ng the end

,
which is a t tach ed to a solid b ody u n lik e th e oth er end The .
s a me t ru th i s s een i n su ch second a ry a gg reg at es a s the com

m on A g ar i cs or r a th er i n th ei r i m m en s ely d e v el op e d or g an s
,
-
of fru cti fi ca ti on A p u ff b a ll Fig 1 9 2 p re s ent s no oth er

.
-
, .
,
obvi ou s u nlik eness of p a rt s th a n th a t b et w ee n i ts u nd er a n d
u pp er s u r f a c es S o too wi th th e st a lk ed ki nd s th at f re qu ent
.
ou r w ood s a n d p a s tu r es I n th e typ es which Fig s 1 9 3

. .
,
1 94 1 95
,
d eli nea te the u n lik eness es b et we en the roote d
, ,
en d s a n d th e e xp a n d e d e nd s as w e ll a s b et w e en th e u n d e r
,
an d u pp e r su r fa c e s o f th e exp a n d e d en d s a r e obvi ou sly,
rel a t e d t o this f u nd a m en t a l c ont ra s t o f con diti on s N or i s .
this rel ati on l es s cl e arly di s pl a y e d i n the sessi l e f un gi w hi c h

gr ow ou t f rom the s id es Of t rees as sh own at a b Fig 1 9 6
, , , . .
T h at which i s c omm on to this and the p rec edi ng typ es is the ,
c ont ras t b etw e en th e a tt ach ed en d and th e f ree end .

THE GE NE RA L S H A P E S OF P L A NT S .
1 39
F rom wh a t th ese form s h a v e i n com m on l et u s t u rn to ,
th a t w hi ch they h av e not i n comm on a nd Ob s erv e th e ca u ses,
o f th e wa nt of co mm un ity . A p u ff b a ll s h ows u s i n th e
-
Simpl es t w ay the lik ene s s of p a rt s a ccom p a nyi n g lik enes s of

,
con di t i ons a l on g w i t h the u n lik en ess of p a rt s a c com p a nyi ng

,
u n lik en e ss of conditi ons For whil e i f w e cu t v erti c a lly

,
.
th rou gh i ts c ent re we fi nd a di ffer en ce b etwe en top a n d

,
b ott om i f we cu t h or i zonta lly th rou gh i ts c en tre w e fi nd no

, ,
di fferen ces a m ong i ts sev era l s id es B e i n g on th e a ver age of

.
,
c a s es
, s i m il a r ly r el a t e d to th e e n vi r on m e nt a l l rou n d it ,
r em a i ns the s a m e a l l rou n d T h e r a di a l s y m m et ry of the

.
m u s h room a nd oth er v er ti c a lly—gr owi ng fun gi ill u st r at es ,
thi s connexi on of c au se a nd effect s till be tt er B u t now .
m ark wh a t h a pp ens i n the g rou p of A ga ri cus n ol i ta ng er e -

,
s h ow n i n Fig 1 9 5 . Ra di a lly s y m m etr i ca l as i s the typ e a n d

.
,
r a di a lly s ymm et r ic a l a s a re th os e c ent r a lly pl a c ed i n divid u a ls-
w hi c h a re e q u a lly c row d e d a l l roun d we see th a t th e pe r i,
p h e r a l i ndivi d u a l s di ss i m il a r ly c i rcu m st a nced on th ei r ou t er

,
s id es a n d on th ei r s id e s next th e g r ou p h a v e p a rti a lly

,
c h a n g e d th ei r r a di a l s y m m et r y i n t o bil a t e r a l s ym m e t ry It .
i s no l onger p oss ibl e to m a k e t wo corre s p on di n g h a lv es by

‘
any v ert i ca l pl an e cu t ti ng d own th r ou gh th e pil eu s a nd the

s t e m ; bu t th ere i s on ly on e v e rt i ca l pl a n e th a t w ill th u s
pr o
d u ce cor res p ondi n g h a lv es— th e pl a ne on th e opp os it e s id es
of whi c h th e re l a t i on s t o t h e e nvi ron m en t a re a lik e A nd .
th e n m a r k th a t th e div erge nc e f rom a l l s id e d s y m m etry -
t owa r d s two s id ed s ymm et ry h er e c au se d i n th e i ndivid u a l

-
,
by s p eci a l ci rcu m st a nces i s ch ara ct eri s tic of th e ra c e wh ere

,
the h a bi ts of th e ra c e c ons t a n tly i n v olv e two s id e d n es s of -
c ondi ti on s . B es id e s b ei ng e x em plifi e d by s u ch comp ara tiv ely

u ndi fferent i a t e d typ es a s ce rt a i n P ol ri Fig 1 9 6 a b this
yp o ,
.
, , ,
t ru th is ex em plifi ed by m em b ers of th e genu s j u st n am ed .
I n A g a ri cus hori zon ta l i s Fig 1 9 6 c w e h a v e a d e p art u re

,
.
, ,
f r om ra di a l s y m m etr y th a t i s c on spi c uou s only i n th e f or m

of t h e s t e m . A m or e d e cid ed bil at er a l n es s e xi sts i n A s u b .
p a l m a tus sh own i n el e v a ti on a t d a n d i n s ecti on a t d

’
, A nd .
L en ti nus fla bel l i for m is of whi ch 6 a nd e a re di fferent vi ews

’
, ,
e xhibi ts c om pl e te bil a te ra l nes s— a bil a ter al nes s i n w hi c h

th e r e is th e g rea tes t lik eness of th e p a rt s th at a re m os t simi
l a rl y c on diti oned a n d th e g rea t es t u nlik enes s of the p a rt s
,
th a t a re m os t di ss imil arly c ondi ti on ed .
A m ong pl a nt s of th e secon d or d er of c om p os iti on it w ill ,
s u ffi c e to n ote one fu rth e r c l a ss of fa c ts w hich a r e th e c on
v erse of th e foregoin g a nd h a v e th e s a m e i m pli ca ti ons T h ese .
a re th e f a ct s s h ow i ng th a t a l on g w ith ha bit u a l i rregu l a rity i n
t he rel a ti ons t o ext erna l f orces t h ere i s h a bi tu a l i rregu l a r ity

,
i n the m od e of g rowth B es id es fi ndi ng su c h fa ct s a m ong

.
T h a ll ophyt es a s i n th e t u b ers Of u nd ergrou nd fu n gi a nd i n

,
t he c reepi ng film s of s es s il e li ch ens w hi c h s ev er a lly sh ow u s

,
v ar i ati ons of p roporti ons obvi ou s ly cau sed by v a ri a ti ons i n

the a m ou nt s of the i n fl u en c es on th e i r di fferent s id es we a l so , ,
a m ong A r c h eg on i a t es of i n fer i or typ e s fi n d i rr e,

gu l a riti es of
f or m a l ong wi th i rre gu l ar iti es i n envi roni ng a cti ons T he .
f ronds of the Ma r cha n tia c ece or s u ch J u ng er m a nni a cece as a re

sh own i n Fig s 4 1 4 2 4 3 ill u st ra t e th e w ay i n w hi ch each
.
, , ,
l owly O rg an i zed a gg rega t e of the sec ond ord er not i ndividu a t e d

-
,
by th e m u t u a l d e p end e nce of i ts p art s h a s i ts f or m d et er

,
mi ne d by the b a l anc e oi f a ciliti es and res i st ances whi ch each

s id e of th e f ron d m ee ts w ith a s it s p rea ds .
§ 21 9 . Am
g pl a nt s displ a yi ng i n t egra ti on of the thi rd
on
d eg ree a nd a m ong pl ant s s till f u rth er com p ou nd ed th ese

, ,
s a m e tru th s a re equ ally m anifest I n the form s of su ch pl a nts

.
we s ee p r im a ry c on t ra st s a n d s econ d a ry c ont ra s ts whi c h n o ,
l ess cl ea rly th an th e foreg oi ng a re rel a t ed to c ont ra s ts of

,
c onditi ons .
T h at fl oweri ng pl ant s f rom th e d a isy u p to th e oa k h a v e ,
i n c omm on the f u nd a m ent a l u nlik eness b etw een the u p wa r d

g rowi ng p art a nd the d ownwar d g rowi ng p a rt ; and th a t
this m ost m ark ed unlik eness c orresp onds with the m ost
m a rk ed unlik eness b etween the two p arts of th ei r envi ron
m ent s oil a nd a i r ; a re f a cts too c onspic u ou s to be n a m ed
,
TH E G E NE RA L S H A P E S OF P L A NTS .
1 41
were th ey not i m p ort a nt it em s i n the ar gu m ent Mor e .
i ns tru ctiv e p erh a p s b ecau s e l es s f a mili ar i s th e f a c t th at we

, ,
mi ss thi s e xt rem e c ontra st i n fl ower i n g pl ants which h a v e not

th ei r high er a n d l ower p orti on s e xp os e d to c on di ti on s thu s
e xt rem ely c ont r a s t e d A p a r a s i te lik e th e D odd er g rowi ng
.
,
i n ent a ngl ed m a sses u p on oth e r pl a nt s from whi c h it su c k s

,
the j u i ces i s not th u s divi s ibl e i nt o two st rongly di sti n gu ish ed

,
-
h a lv es .
L ea vi ng ou t of cons id er a ti on th e di ff erenc e b e twee n the

s u pp orti n g p a rt a nd t h e s u pp o rt e d p a rt i n ph a n og a m s a nd ,
l ooki ng at th e su pp ort ed p ar t only w e ob serv e b etween i ts

,
f orm an d th e h a bitu a l i ncidence of forces a r el a ti on lik e th a t

,
wh ich we ob serv e d i n the s i m pl er pl ant s P ha nog a m s th a t .
a re p ra ctic a lly i f n ot li t era lly u n i a xi a l a nd th o s e whi c h de

,
ve l op th e i r l a t era l a x e s on ly i n th e s h a p e of a xill a ry fl owers ,
wh en u n i nterfer ed w i th c om m only send u p v ert i ca l st em s

r ou n d w hi c h th e l ea v e s a n d fl ow er s a r e di s p ose d with a m or e
or l ess d ec id e d ra di a l sy m m e t r y .Ga r d en s a n d fi eld s s u pply

u s w ith s u c h i nst a nc es a s th e T u lip a n d th e Or c hi s a n d on ,
a l ar g er s ca l e th e P a l m s and th e A l oes a re fertil e i n ex

,
a m pl e s T he ex c e pti ons too a re i ns t ru ctiv e

.
, , Bes id es th e
.
i n divid u a l div erg enc es a r i s i ng f rom s p eci a l i nter feren c e s th ere ,
a r e to be t r a ce d g en era l div e r g en ces wh ere th e h a bit s of the
pl a nt s exp os e th em t o genera l i n t er f erences i n a nythi n g

a pp r oa chi n g to c on st a nt w a y s . P l ants whi ch lik e th e F ox
,
gl ov e h a v e s pik es of fl ower s th a t a re b orne on fl exibl e f oot

,
s ta lk s h a v e th ei r fl owers h a bit u a lly b e n t r ou n d to one f a c e of

,
the st em : an u n lik enes s of di strib u ti on p r ob a bly cau se d by

Th e wi ld H ya
’
u n lik eness i n th e r el a ti on to th e S u n s r a y s .
c i nth too with s t e m s o fl exibl e th a t i ts u pp er p a rt d r oop s

, , ,
s h ows u s h ow a c on s equ en t di f fe r enc e i n th e a c ti on of gr a vity

on th e fl ower s c au s es th e m to d evi a t e f r om
,
th ei r typic a lly
r a di a l a r r a ng e m en t t owa r d s a bil a t era l a r r a ng em en t .
Mu ch m ore c on s pi cu ou s a re t h es e g enera l a nd s p eci a l rel a

ti ons of f orm t o genera l a nd s p eci a l a c ti on s i n the envi r on
m ent a mong ph a nog a m s th at a r e m u lti a xi a l
, T h at whe n
.
st a ndi ng a l one a nd i n pl aces wh ere the wi nd s do not i nj u re

,
the m nor a dj a cen t th i n g s sh a d e th em sh ru b s and t rees d ev el op

,
w ith t ol er a bl e e v e nn es s on al l s id es i s a n obvi ou s t r u th
,
.
E qu a lly obvi ou s i s th e t ru th th a t wh en g row i ng t ogeth er i n a

,
wood a nd m u t ua lly i n t erfered w ith on a l l s id es t rees s till

, ,
s h ow ob s cu r e ly ra di a l di s tr ib u t i ons of p a r t s ; t h ou gh u n d er ,
su ch c on di t i on s th ey h a v e t a ll t a p er st em s w ith b ra nch es
,
di rect ed u pw a rd s— a di fference of s h a p e cl ea rly du e to th e

di fferent i ncid ence of for ce s A nd a lm ost equ a lly obvi ou s i s
.
the t ru th th a t a t ree of thi s s a m e ki n d g row i n g a t the e dge

,
o f th e wood h as i ts ou t er b ra nch e s w ell d e v e l op e d a nd i ts

,
i n ner b ranch es comp ara tiv ely ill —d ev el op ed Fig 1 9 7 which

. .
,
in accu ra t ely rep resents this diff erenc e will s erv e to m ak e it

,
m a n i fes t th a t whil e one of the p eriph era l t rees can be cut .
i nt o s om ethi ng lik e tw o s imil ar h alv es by a v erti ca l pl a ne

di rect e d t owa r ds th e cent re of the wood— a pl ane on each s id e .
of whi ch th e c on diti on s a r e a lik a —i t c a nnot be cu t i n t o s imi
l ar h a lv es by a ny oth er pl a ne . A lik e div ergen ce from a n .
i n defi n i tel y ra di al symm et ry t ow ar ds a n i ndefi ni tel y bil a t er a l

- -
s y m m et r y
,
occu r s i n t re es th a t h a v e t h ei r c onditi on s m a d e
bil a t era l by growi ng on i n c li ned s u rfaces T wo of th e c omm on

.
f orm s ob serv a bl e i n su ch c a se s a re giv en i n Fig 1 9 8 H ere

. .
th er e i s divi s ibili ty i nt o p arts th at ar e t ol er a bly s imil ar by ,
a v e r ti ca l pl ane ru nn i n g di re ctly d own th e hill ; bu t n ot by
a ny oth er pl an e T h en furth er th er e i s th e bil at era l ness

.
, , ,
si m il ar i n g enera l m eani ng th ou gh di ff eren tly ca u sed oft en ,
s een i n t rees exp os ed to s trong p rev a ili ng wi nds A lm ost

.
TH E G E NE RA L S H A P E S OF P L A NT S .
1 43
v y s ea coas t h as a bu n d ant ex am pl es of s tu nt ed t rees w hi ch

e er -
,
l ik e the one sh own i n Fig 1 9 9 h a v e b een m a d e t o d evi a te

.
,
from th ei r or di na r y equ a l g rowt h on a l l s id es of a v er ti ca l

a xi s to a g rowt h th a t i s e q u a l on ly on th e opp osit e sid es of a
,
’
v erti ca l pl an e di rect ed t owa r d s the wi nd s eye .
F rom a m ong v eget a l a ggreg at es of th e thi rd or d er we h av e ,
n ow on ly to a dd e x a mpl e s of the enti rely a sym m et r ic a l f or m
w hi ch a ccomp a n i e s a n en ti rely i rre gu l a r dist r ib u ti on of i nci

d ent forces C reepi ng pl ant s fu rni s h su ch e x a m pl es T h ey
. .
s h ow bot h wh en c li m bi n g u p v erti ca l o r i n cli ne d s u r f a ces an d

,
wh en t r a ili ng on th e g rou nd th a t th ei r b ra nch es g row hit h er

,
a n d thith e r a s th e b a l a n ce of f or c es a id s or Opp os es ; a nd th e
g ener a l ou tli ne i s w ith ou t s ymm et ry of any ki n d be c au se ,
the en vi ron i n g i n fl u enc es h a v e no ki n d of regu l a rity i n th ei r

a r r a n g e m en t .
2 20 A l on g with
. s om e u nfa mili a r f a ct s I h a v e h ere s et
,
d own f a ct s whi ch a re so fa mili ar a s to s eem sca rce ly worth

n oti n g .I t is b eca u se th es e fa ct s h a v e b ec om e m ea ni ngl e ss
to p er c e p ti on s d ea d ene d by i n fi nit e r ep etiti on s of th em th a t ,
it i s needf u l h er e to p oi nt ou t th ei r m ean i ngs N ot a l one for .
i ts i ntr i ns i c i m p orta nc e h a s the u n lik en ess b et w ee n th e

a tt ac h e d e n d s a n d th e f re e en d s b een t r a ce d a m on g pl ant s of
a l l d eg r ees of i nt eg r a ti on N or i s i t s imply be ca u se of th e
.
s ign ifi c an c e th e y h a v e i n th em s elv es th a t i n sta nces h a v e

,
b een giv en of th os e v a ri eti es of s ymm et r y a n d a s ymm et ry

whi ch the f ree en d s o f pl a nts eq u a lly di s pl a y : be th ey pl a n t s
'
o f the fi r s t s e con d t h i r d or a n
, , , y high e r or d e r N e i th er h as
.
th e on ly oth er p u r p ose b een th a t of s h owi n g h ow i n th e r a di a l ,
s ym m et ry of s om e v e g et a l a gg r e g a t es a n d th e s i ngl e bil a ter a l
s ym m et r y of oth e r s th er e a re t r a c ea bl e the sa m e u lti m a t e

,
p ri n cipl es a s i n the s ph eri ca l symm etry an d t ripl e bil a t era l

sy m m etr y of cert a i n mi nu t e pl a nt s fi rs t d e s cr ib e d Bu t th e .
m a i n Obj ect ha s b een to p r esent u nd er th e i r s i m pl es t a sp ect s

, ,
those gener a l l aw s of m or ph ol ogic a l differ enti ati on which are

fulfill ed by th e c omp onent p a rt s of ea ch pl ant .
If g a nic f or m i s d et er m i ned by th e di s t ribu ti on of f orc es ;

or
a n d th e a pp r oa c h i n e v ery c ase t ow a r d s a n e qu ilib r i u m of
i nner acti on s with ou t er a cti ons ; th en t hi s rel a ti on b etween

form s a n d for c es m u st h old a l ik e i n th e orga ni sm a s a w h ol e
i n i ts p r oxim a te u nit s an d i n i ts u n i ts of l ow er or d er s
, For .
m u l as w hi ch exp re s s th e sh a p es of enti re pl ant s i n ter m s of

s u rr ou n di ng c on di ti on s m u s t be for m u l as w hich a l so e xp r es s
,
t he s h a pe s of th ei r se vera l p ar ts i n t er m s of s u r rou n di n g
c on di ti ons . I f th erefore w e fi n d th a t a pl ant a s a wh ol e is
, ,
ra di a lly s y m m etr i c a l or bil ate r a lly s y m m e t ri ca l or a sym m e
t r i c al a ccor di n g a s the in c id ent f or ce s a ffec t i t equ ally on a l l

,
s id es of a n a xi s or a f ,
f ect it equ a lly on ly on the opp os it e s id es
o f one pl a n e o r a f
,
f ect it equ a lly i n no two di r ecti on s ; th en ,
w e m a y exp ect th a t i n lik e m a n ner each m e m b e r of a pl an t

, ,
w ill di s pl a y ra di a l sy m m et r y w h e re envi ron i n g i n fl u en ces a re

a lik e a l on g m a n y ra dii bil a tera l s y m m etry wh ere th er e is
,
bil a tera l ness of envi roni n g i nflu enc es and u nsym m et ry or

,
a s y m m et r y wh er e t h ere i s p a rti a l or en ti r e d ep a rt u re f rom a
b a l a n ce of su rrou n di ng acti on s .
T o s h ow th a t thi s exp ect a ti on i s b orne ou t by the f a ct s ,
will be th e obj ect of th e f oll owi ng fou r ch a pt ers L e t us .
begi n with the l arges t p art s i nt o whi ch pl ants a re divisible ;

a nd p roc ee d to th e s u cc essiv ely sm a ll er p a rts .
C H A P TE R VI I I .
TH E S H AP E S OF B RA N C H E S .
§ 2 2 1 A GGRE GA TE S of the fi rst or d er s u pply a few ex ampl es

.
o f f o rm s ra m i fi ed i n a n a pp r oxim a t ely r egu l a r m a nn er u n d er -

,
c onditi on s whi c h s u bj ec t thei r p arts to a pp roxim at ely r egu -
l ar dist r ibu ti on s of forc es S om e u n i ce ll u l a r A l yce b ecomi ng

.
,
e l a b ora t e ly b r a n ch ed a s su m e v ery m u ch th e a sp e ct s of s m a ll
,
t ree s ; a nd s h ow u s i n thei r b ra n ch es a na l og ou s rel a ti ons of

f or m s to f orces B ryops is p l u m os a m ay
.
be i nst anc e d .Fig 200 rep res ent s the

.
end of on e of its l a t er a l ra mifi c a ti on s ,
ab ov e an d b enea th whi ch c om e oth er s of

lik e ch ar a ct ers H er e it will be s een th at
.
the a tt a ch ed an d f ree en d s differ ; th a t

the two s id e s a re m u ch a lik e ; an d th a t th ey are u nlik e the
u pp er a nd u n d er s u r fa ces w hich rese m bl e one a n oth er
,
T he .
m ore highly d ev el op e d m em b ers of th e s a m e g rou p of A l gae ,
the S i p h on ece sh ow a m a r k e d r a di a l symm et ry c oexi sti ng

,
with v ery el a b ora t e b ra n chi ng e g N eo , . .

,
m eri s O m o ol i a a n d oth er s
, y p ,
.
§ 2 22 . Fig h ows u s h ow i n an
. 201 s ,
a ggr eg a t e of the sec on d or d er ea ch

p roxi ,
m at e c om p onent i s m odifie d by i ts rel a

ti on s to the rest ; j u st a s we b efor e s aw
a wh ol e fu ngu s Of th e s a m e typ e m odifi e d
56
1 46 MORP H OL OGI CA L D E V E L OP ME NT .
by its rel ati ons to envi roni ng obj ect s I f a b r a nc h of th e .
fu ngu s h er e figu re d be c omp a r e d w i th on e of th e fu n gi

,
cl u s t ered togeth er i n Fig 1 9 5 or s till b ett er wi th one of the

.
, , ,
l atera lly gr owi ng fu n gi s h own i n Fig 1 9 6 th ere will be per

-
.
,
ce i ve d a ki nd re d t r a n s i ti on f r om r a di a l to bil a t er a l s ymm etr y ,
occu r r i ng u n d er ki n d re d c on diti on s T h e p orti on of th e .
pil eu s next to the s id e of a tt a ch m ent i s u nde v el op ed i n this

b r anch e d f orm a s i n th e s impl er for m ; and i n the one case a s
i n the oth e r th e s t em i s m odifi ed t owa rd s th e sid e of a tt a ch
,
.
m ent A divi s i on i nt o s i m il ar h alv es whi ch a s s h own i n

.
, ,
Fig 1 9 6 e m ight be m a d e of the wh ol e fu ngu s by a v ertic a l

.
,
.
pl a n e p ass in g th rou gh the cent re of the pil eu s a nd the a xis

o f the s u pp orti n g b ody m ight h ere be m a d e Of th e b r a nc h
, ,
by a v erti ca l pl ane p a ss i ng th rou gh the cent re of i ts pil eus

a n d th e a xi s of th e m a i n st em A m on g a gg reg a t es of this .
or d er the A l yce fu r ni s h c a s es of ki n d re d n atu r e

, I n the .
b r a nch es of L ess oni a Fig 3 7 m ay be ob ser v e d a su bst anti ally

, .
,
s imil a r rel a ti on ship A s th ei r i nner p a r t s a re l ess d e v e l ope d

.
th an th ei r ou ter p art s whil e th e i r two sid es a re d evel op e d i n

,
a pp r oxi m a t ely e qu a l d egr ees th ey a r e ren d er ed bil a t e ra l

, .
223 . T h es e few c a ses i nt rod u c e u s to the m ore f a mili ar

bu t m ore c ompl e x c a s es which pl ants of the thi r d d eg ree of
aggr eg ati on p resent A t a .

, b, c, Fig 2 02 are sk etch ed th ree
.
,
h om ol ogou s p art s of th e s am e t ree : a be i ng the l ea di ng

T HE S H AP E S OF BRA NCH E S .
sh oot ; b a l a t er a l b ra nch n ea r the top a nd c a l a t era l b ran ch ,
l ower d own T here i s h ere a d ou ble exem pl i fi ca ti on Whil e

. .
the b r a n c h a as a wh ol e h a s i ts b ra n c hl et s a rr anged w i th
, ,
t ol era bl e reg ul a r ity a l l round i n corresp ond enc e wi th i ts ,
equ a l e xp osu r e on a l l s id e s e a ch b r a n chl et s h ows by its,
c u r v e a s m u c h bil a ter a l s ym m et r y a s its simpl e f or m p er mit s .
T he b ra n c h b di ss i m il a r ly ci r c u m st a n ce d on th e sid e next
,
the m a i n s t em a n d on the sid e a w a y f rom it h a s a n a pp roxi ,
m a t e bil at er a l ness a s a wh ol e w hil e th e bil a t er a l nes s Of i ts ,
b ra nchl et s v a ri es w ith the i r res pec tiv e p os i ti on s A n d i n the .
b ran ch 0 h a vi ng i ts p a rt s still m or e di ff ere ntly c ondi ti oned

, ,
th ese t raits of stru ctu re a re s till m ore m a rk ed E xt rem ely .
s t ron g c on t r a st s of t hi s ki n d occ u r i n t r e es h a vi n g v e ry
regu l a r m od e s of g r owth T h e u pp er m ost b r a n ch es of a

.
S p ru c e fi r h a v e ra di a lly a rra nge d b r a nchl ets : ea ch of th e m

- -
,
if g rowi ng vig orou s ly repea t s the typ e of t he l ea di ng s h oot

, ,
a s s h own i n Fig 2 03 a b . B u t i f w e e x a mi ne b ra nc h es
, , .
l ower a nd l ow er d own the t ree we fi nd th e v er ti ca lly growi n g ,

-
b ranchl ets be a r a l ess a nd l e s s ra ti o to the h ori zont a lly

growi n g ones ; u ntil tow a r ds th e b ottom the ra di a l a rran ge
, ,
m en t ha s wh olly m er ge d i nto th e bil a t era l S h a d ed a nd .
c on fined by the b r a n c h es a bo v e th em th es e eld es t b r anch es ,
d evel op the i r offs h oots i n those di re cti on s w h ere th ere i s

m ost s p a ce an d light : b ecomi ng fi n a lly qu it e fl a ttene d and
fan s h a p ed a s sh ow n a t Fig 2 03 0
-
,
A n d on r e m e mb er i n g
.
, .
th a t ea ch of th es e eld es t b ra nch es wh en fi rs t it diverge d ,
f rom the m a i n s tem w a s r a di a l we s ee not only th at b et ween

, ,
the u pp er a nd l ow er b r a n ch e s d oes thi s contr a s t i n s t ru ctu re

hold bu t a l s o th a t eac h b r an ch is t r an s form e d f rom th e
,
r a di a l to th e bil a te r a l by th e p rog r ess iv e c h a n g e i n i t s en
v i ron m en t . Oth er forc e s b es id es th ose w hich a i d or

hi nd er growth con spi re to p r od u ce this two s id e d ch a ra cter
,
-
i n l a t er a l b ra n ch e s T h e a nne x e d Fig 204 s k etch e d fr om

. .
,
a n ex a m pl e of th e P i nus C ou l teri i a t K ew sh ow s v er y cl ea r ly
,
h ow by m er e g r a vit a ti on th e on c e ra di a lly a r ran ged b ra n ch

, ,
-
l et s m ay be so be nt a s to p rod u ce i n the b ranch a s a wh ol e a

d ec id ed bil at era ln ess A f u ll grown A r au ca ri a too exhibits

.
-
, ,
i n i ts l ower b r an ch es m odifi c a ti on s s imil a rly c au se d ; a n d i n

e a c h of s u c h b r a n ch e s th er e m ay be r em a r k e d t h e fu r th er
f a c t th a t i ts u p w ar d b en di ng term i n ati on h a s a p ar ti a lly

,
-
m odi fi ed r a di a l n es s at th e s am e t im e t h a t i ts d roopi n g l a tera l

,
b ran chl ets giv e to the p ar t nearer the t runk a c ompl etely
bil a tera l ch a r a ct er .
N ow i n th es e few i nst a nc es typic a l of c ou ntl ess i nst a n c es ,
whi ch might be given we see a s we s aw i n the c a s e of , ,
t he fu n gi th at th e s a m e t hi ng i s t rue of the p a rt s i n th ei r
,
rel a ti on s t o t he w h ol e a n d t o on e a n oth er w hi ch i s t r ue of ,
the w h ol e i n i ts rel a ti ons to th e envi r onm en t at l a r ge E nti r e .
t rees b ecome bil a t er a l i n stea d of ra di a l wh en expo s ed to ,
f or ces th at a re equ a l only on opp os it e s id es of on e pl an e ; a nd

I n t h e i r b r a n ch es p ara ll el ch a nge s of f or m oc cu r u nd er
,
p a r all el ch a nges of con diti ons .
2 24 . T h er e s a id s om ethi ng resp ecti ng the

r em a ns i to be
di s t ribu ti on of l ea v es H ow a b r anc h c a r r i es i ts l e av es
.
c onsti tu t es one of i ts c h a r a cter s a s a b ran ch an d is to be ,
c ons id e r e d a p a rt f ro m th e c h a r a ct ers o f th e l ea v es th e m
s elv es T he p r i n cipl es hith ert o ill u s t ra t ed we s h a ll h ere find

.
ill u s t ra t e d s till f u rt h er .
T he l ea di n g s h oot a nd al l the u pp er t w ig s of a fi r t ree -

,
h av e th ei r p i n sh a p ed l ea v es e v enly di st r ib ut ed a l l r ou nd or
-
,
pl aced r a di a lly ; bu t a s w e d es cen d w e fi n d th e m b egi nnin g

to a s su m e a bil a t era l dist r ib u ti on ; a n d on the l ow er h or i zon ,
t a lly g rowi ng b r anch es th ei r di st r ibu ti on i s qu it e bil a tera l !

-
,
B etween th e I ri s h a n d E ngli sh ki nd s of Y ew th ere i s a con ,
t ra st of lik e s ignifi ca nce T he b ranch es Of th e one s h ooti n g .

,
u p as th ey do a l m os t v er tic a lly a r e c l oth e d w i t h l eav e s ,
He re a nd th ro u gh out th e word r di a l i s a pp l ied e q ual l y to th e s pi a l

,
a r
a nd th e wh o rl e d s t ru c t u re s . The s e, as b ei ng a l i k e on a l l s id e s , a re si m il a rl y
d i s ti ng ui s h e d from a rra ng em en t s th a t a re a l i k e on t wo s i d e s onl y .
+ It s h oul d be a dde d th a t th i s c ha n ge of d i s tri b u ti on i s no t du e t o
ch a nge i n the r el a t i ve p os i ti ons of th e i nse rt i ons of th e l e a e s but to th e i r v

t wi s t i ng s .
THE S H A P E S OF BR A N C H E S .
1 49
al l r oun d ; while thos e of the oth er whi ch s p r ea d l at era lly , ,
b ea r th ei r l ea v e s on the two s id e s I n trees w ith b etter .
d ev el op e d l ea v es the s a m e p r i nc ipl e is m ore or l ess m a ni fest

,
i n p r op orti on a s th e l e a v es a r e m ore or l ess en a bl e d by th e i r

s t ru ct u res t o m a i n t a i n fix e d p os iti on s Wh ere th e f oot s t a lk s .
-
a re l on g a nd sl en d er a n d wh e r e conseq u en tly
, ea ch l ea f
, , ,
a cc or di n g to i ts weight th e fl e xibili ty a nd t wi s t o f its f oot

,
s t a lk a nd th e di r e cti on of th e b ra nc h it g row s from

,
fa ll s ,
i nt o s om e i n defi n it e a ttitu d e the r el ati ons a re ob scu red B u t

,
.
wh er e the f oot s t a lk s a re sti ff a s i n the L a u re l it w ill be

-
, ,
fou nd a s b efor e th a t f rom th e t op m os t a nd u p w a r d g row i n g

, ,
-
b ra nch es th e l ea ve s div erg e on al l s id e s ; whil e th e u nd er

m ost b ran ch es g rowi ng ou t from the s h a d e of th ose a b ov e
, ,
h a v e th ei r l ea v es s o t u rne d a s to b r i ng t h e m i nt o r ow s h ori
zonta l l y s p r e a d ou t on th e two s id e s of ea c h b r a n c h .
A ki nd re d t ru th h a vi n g lik e i m plic a ti on s c om es i nt o Vi ew
, ,
wh en we Ob s erv e th e rel a tiv e si z es Of l ea v es on the sa m e

b r an ch wh ere th ei r s i zes di ffer
,
.
F i g 2 05 rep r esent s a b r a n ch of a
.
H orse—ch est nu t t a k en from th e

,
l owerm os t f ri ng e of th e t r ee w h ere ,
the light h a s b een to a g r ea t e xt en t

i nt erc ept ed f rom a l l bu t th e m ost
p r ot ru d ed p a rt s B evon d the f a c t
.
tha t the l ea v es b ecom e by a pp ro

pri ate g rowth s of th e i r f oot st a lk s
-
bil atera lly di s t ribu te d on thi s d roop

i ng b r a nc h i nstea d of b ei n g di st r ib u t ed symm et ri ca lly al l
,
rou n d a s on o n e o f t h e a s c en di n g s h o o t s
,
w e h a v e h ere t o ,
n ot e th e fa c t th a t th er e i s u n equ a l d ev e l opm en t on th e u pp er
a nd l owe r s id e s E ac h of the c om p ou n d l ea v es a cqu i r es a

.
foot stalk an d l ea fl et s th a t a r e l a r ge i n p rop or ti on to th e

-
s u pply o f light ; a n d h en ce a s w e d escen d t ow a r ds th e bot

,
tom of th e tr ee th e c l u s t er s of l ea v es di s pl a y i nc r ea s i n g
,
c on t r a st s H ow m a r k e d t h ese c ont ra st s b ecom e w ill be s een

.
on c omp ari n g a a n d b whi c h for m -
one p a i r of l ea v e s th a t
,
1 50 M ORP H OL OGI CA L D E VE L OP M E NT .
a re n or m a lly eq u a l or c a nd d whi ch f orm a nother p a i r nor

, ,
m a lly equ a l .
L et u s not omi t to not e w h il e we h a v e thi s ca s e b efor e u s

, ,
the p roof it a fi or ds th a t t h es e di ff eren c es of d ev el op m ent a r e

i n a c on sid er a bl e d egr ee d eter m i ned by the di fferent con
di ti on s of th e p art s a fter th ey h a v e b een u n fold ed T h ou gh .
th ose i n equ a liti es of di m en s i ons wh ence the di fferenti a t i ons

of f or m resu lt m ay be i n m a n y cas e s l a r g ely du e to th e
,
i nequ a liti es i n the c i r cu m st ances of the p a rts whil e i n th e

bu d ( whi ch a r e h owev er r ep res ent a tiv e of i nequ a liti es i n
, ,
a n c est r a l ci r cu mst a n c es ) ; yet th ese a re cl e arly n ot th e s ol e
c au s e s of the u nlik eness es w hich e ventu ally a ri se T his bi .
l a tera l ness r esu lti n g f rom th e u neq u a l s i zes of the l ea v es ,
m u st be c ons id ered a s du e to the di fi erenti a l a cti ons th at

c om e i nt o pl a y a ft er the l ea v es h av e a s su m e d th ei r typic a l
st ru ctu res .
2 25 . H ow, i n the gem ent of th ei r t wigs and l eav es

a rr an ,
b ra nch es t en d to l a p s e from form s th a t a re a pp roxim at ely

s ym m et ri c a l to for m s th a t a re q u i te a sy m m et ric a l n ee d n ot ,
be d e m ons t ra t e d : it i s su ffi ci ently c ons pi cu ou s Bu t it m ay

.
be w ell to p oi nt ou t h ow th e t end enc y to do thi s fu r th er

e n forces ou r a r gu m en t T he c om p a ra tiv ely r egu l a r b u ddi ng
.
ou t of s ec on d a ry a x es a n d t erti a ry a x es d oes not u s u a lly

,
p rodu ce a n agg reg a t e whi ch m a i ntai ns its reg ul a r i ty for ,
the s impl e rea son th a t m a ny o f th e a x es a b or t T erm i na l

.
b u d s a re s om e of th em d es t roy ed by bi rd s ; oth ers a re bu r

r ow e d 1 n to by i n s e c t s ; oth ers a r e n ipp ed by fro s t ; oth e rs
a r e b rok en off or i n j u re d d u r i n g g a l es of w i n d The c nvi .
r on m ent of ea c h b r a n ch a n d i ts b r a n chl et s i s th u s e v e r
b ei n g v a ri ed on a l l s id es : h ere sp a c e b ei n g l eft v a ca nt by
,
the d eath of som e s h oot th a t w ou ld ordi n a r ily h a v e occu pi e d

it ; a n d th ere sp a c e b ei ng trench ed on by th e l ate ra l gr ow th
,
o f s om e a dj a c en t b r a nc h th a t h a s h a d its m a i n a xis b rok en .
H ence th e a s ym m et ry or h et erogen eity of form a ssu m ed

, ,
b y th e b r a n ch i s c au s ed by the a symm e t ric a l di st rib u ti on

,
T HE S H A P E S OF B RA NCH E S .
1 51
of i nc id ent f orc es— a r es u lt a nd a ca u s e whi ch g o on v

e er
c om pli c a ti n g .
§ 22 6 . pi cu ou s t ra it i n the sh a p es of b ra nch es
On e c on s
h a s s till to be n a m e d T h ei r p roxi m a l or a t ta ch ed end s

.
di ffer f rom th ei r di s t a l or free end s i n th e s a m e w ay th a t ,
th e l ower en d s Of t r ees di ffer f r om th e i r u pp er en d s T hi s .
f act lik e the f act to whi ch i t i s h er e p a ra lle l ed h as h a d i ts

, ,
s ign ifi ca nce ob scu r e d by i ts e xt r em e f m ili a r ity B u t it

a .
s h ow s i n a s t r iki n g w a y h ow th e m os t diff e r en tly c on diti oned -
p a rt s b ec om e th e m ost st rongly con t r a ste d i n th e i r s t ru e

t u res A pha nog a m i c a xi s i s m a d e u p of h om ol ogou s s eg
.
m en ts m a r k e d off from on e anoth er by th e nod es ; and a

,
c om p ou n d b ra nc h c ons i s ts o f grou p s of s u c h seg m ent s T h e .
ea rli es t for m e d-
s egm en t s a lik e o f th e t r e e a n d o f ea c h
,
b ranch s er v e as m ech an i ca l s u pp ort s a nd ch a nnel s for s ap to

,
th e su ccess iv e g en er a ti ons of se g m ent s th a t g row ou t of

th em ; and b ecom e m or e a nd m ore sh a d e d by th ei r p rog eny
a s th es e i nc r ea s e H en ce th e p rog r es s iv e ly i nc r ea si ng con
.
-
tr a sts w hi c h whil e m a i nly du e to the u n lik eness es of b u lk

,
a cc om p a nyi n g difi e r en ces of a g e a r e i n p a r t du e to th e u n

,
lik en esse s of s t ru ctu re w hi c h differenc es of r el ati on to the

envi r on m en t h a v e ca u s e d .
22 7 th en it i s w ith the p r oxim a t e p a rt s of pl a nt s

. T h u s, ,
a s it is with pl a nt s as wh ol es T h e ra di a l s ym m e t r y the
.
,
bil at era l sy m m et r y a n d th e a symm et ry whi ch b r a nch es di s

, ,
pl a y i n di ff erent t rees i n di fferent p a rt s of th e s a m e t ree a nd

, ,
a t di fferen t s t a g e s of th ei r ow n g row th s p rov e t o be a l l con ,
s eq u ent on th e w a y s i n w hi c h th ey s t a n d t ow a r ds th e en ti re
pl ex u s Of su rrou ndi ng a c ti on s T h e p r i n cipl e th a t th e

.
g rowt hs ar e u nequ a l i n p r op orti on a s th e rel a ti on s of p a rts

to th e envi ron m en t a r e u nequ a l s er v es to expl a i n al l th e ,
l eadi ng t ra its of st ru ct u re .
C H A P TE R I X .
TH E SH A P E S OE L E AVE S .
228 . N E X T i n th e d es c en di ng or d er of c omp ositi on com e

c omp ou n d l ea v es .T h e r el a tiv e s i zes a nd di s t rib u ti on s of
th ei r l ea fl ets a s a ffec ti ng th ei r f orms a s wh ol es h av e to be
, ,
c on s id e r e d i n th e i r rel a ti on s to con di ti on s Fig s 2 06 207 . .

, ,
r ep r es en t l ea v e s of th e com m on Oxa l i s a nd o f th e M a rs i l ea ,
i n whi c h ra di a l s ym m etry i s a s c om pl et ely di s pl ay e d a s th e

s m a ll nu m b er of l e a fl et s p er m its T hi s equ a l d ev el opm ent
.
of th e l ea fl et s on a l l s id e s oc cu r s wh e r e th e foot s ta lk s g row
,
-
,
i ng u p v e rti ca lly from c reepi n g or u n d erg rou n d s t em s a re ,
s o l on g th a t th e l ea v es e ith er do n ot i n t er fere wi th one
a noth er or do it i n a n i nc onst a n t w ay : th e l e a fl et s a r e n ot
di ff eren tly con diti one d on di fferent s id es as th ey are wh ere ,
th e foot s t a lk s gr ow ou t i n the ordi n a r y m a nn er

-
H ow u m .
lik eness of p os i ti on i nfl u en ces th e l eafl et s i s cl e a r ly sh own i n

a C l ov e r l ea f
-
,
Fig 2 08 which d evi at es from the Ox a li s l ea f
.
,
-
bu t s ligh tly towa rd s bil a t er a ln ess a s it d evi a t es from i t bu t

,
s lightly i n th e a t tit u d e of i ts p eti ol e ; whi c h i s a li ttl e i n
c l i n ed a wa y from th e oth e r s b or ne by th e sa m e p r ocu m b ent
a xi s
. A f a m ili ar e x a mpl e of an a l m ost r a di a l sym m etry
a l on g w ith a lm os t e qu a l rel a ti on s t o s u r rou n di ng c on diti on s ,
oc cu r s i n th e r o o t l ea v es o f th e L u pi n
-
Fig 2 09 b
,
H ere
. .
th ou gh we h a v e l a tera l div ergence from a v ertic a l a xi s yet ,
th e l on g f oot s t alk s p res er v e nea rly er ect p os iti ons a n d

-
,
c a rry th ei r lea v es to s u ch di st ances f rom the a xis th a t the ,
d evelopm ent of the l ea fl ets on the sid e next it i s not m u ch

1 52
TH E S H A P E S OF L E A VE S .
1 53
hi n d ered S till the i nt er ferenc e of the l e a v es with one

.
a n oth er is on th e a v er a g e s om ew h a t grea t e r on th e p r oxi m a l

, ,
s id e th a n on th e di s t a l s id e ; a n d h en c e th e i n t e r i or l ea fl et s
a r e r a th er l ess t h a n th e e x t er i or l ea fl et s I n fu rt h er p roof of
.
w hi c h i nfl u enc e l et i t be a dd ed t h a t a s s h own i n th e fig u re
, , ,
at a ,the l ea v es g rowi n g ou t of t h e fl oweri n g—s te m d evi a t e

t ow a rd s the two s id ed f or m m ore decid edly T wo—s id ed nes s
-
.
i s m u ch g r eater w h ere th ere i s a g r eat er r el a tiv e p roxi m i ty

of th e i nn er l ea fl et s to th e a xis or w h ere th e foo t st a lk
,
-
a pp roac h es t ow a r d s a h or i z on t a l p os iti on T h e H orse ch es t

.
-
nu t Fig 2 05 a l re a dy i n st an ce d a s s h ow i n g h ow th e a rr a n g e
,
.
,
m ent s and s i zes of l ea fl ets a re d et er mi ned by the i nc id ence of

f orc es s erv e s a l so to s how how the i ncid enc e of forc es d et er
,
mi nes l tiv e si zes a nd a r rangem ents of l ea fl ets Fig

th e re a . .
2 1 0 whi ch s h ows a l ea f of th e B om ba x fu rth er ill u s t ra t es

, ,
thi s re l a ti on of stru ctu re to c on diti ons .
Comp ou n d l ea v es th a t a re c ompl et ely bil a t er a l p r esen t u s ,

'
wi th m odifi c ati ons of form exem pl i fyi ng the s a m e g enera l

t ru th i n an oth er way I n th em th e p r oxim a l a n d di st a l
.
p a rt s h a v e n one of th a t r esem bl a nc e which we see i n th ose

i nt erm edi a te for m s j u s t d es c r ib ed T h e p orti on next th e a xi s
.
a n d th e p orti on fu r th es t from th e a xi s a r e e n ti r e ly di f
feren t ;
a nd th e only lik en es s i s b et ween th e w i n g s or l ea fl et s on
o pp os it e s id e s o f th e m a i n f oot st a lk or mid r i b
-
On t u r ni ng -
.
b ac k to Fig 6 5 it will be s een th at th e c om p ou n d l ea f th ere

.
,
1 54 MORP HOL OGICA L D E VE L OP ME NT .
d rawn to ex em pli fy a noth er t ru th serv es a l so to ex em pli fy thi s

,
t ru th : the h om ol ogou s p a rt s a b c d whil e they a re u nlik e

, , , ,
o n e a n ot h er ,
a re i n t h ei r m a i n p rop ort i on s sev era lly lik e
, ,
th e p a rt s w i th w hi c h t h ey a r e p a i re d A n d h e r e l et u s n ot
.
ov e r l ook a c h a r a c te r i s ti c wh i c h i s l e ss c ons pi cu ou s bu t n ot
l ess s ign ifi ca n t . E ac h of the l a t e ra l w i ng s h a s wi ngl et s

t h a t a re l a r ger on the on e S id e th a n on th e oth er ; a n d i n
ea ch c a s e t h e two sid es a r e di ss i m il a r ly c on di ti one d E v en .
i n t h e s ev era l c omp on ents of ea c h wi n g m a y be t r a ce d a lik e

div ergence from s ym m et ry a l ong wi th a lik e i nequ a li ty i n
,
t h e rel a ti on s to th e rest : th e p roxim a l h a lf of each l ea fl et

i s h a bit u a lly l a rger th a n t he di s t a l h a l f I n t h e l ea v es of
.
t he B ra m bl e p revi ou sly fi g u re d ki nd red f a ct s a re p r esen t ed

, , .
H ow fa r su c h di ffe r ences of d ev e l op m en t a re du e to th e p os i
ti ons o f the p a rts i n th e bu d ; h ow fa r the res p ectiv e
s p a c e s a v a il a bl e for th e p a rts w h en u n f old e d a f fec t t h e m ;
a n d h ow far th e p art s ar e ren d e r e d u n lik e by u nlik en esses
i n th ei r rel a ti ons to light ; it i s di ffi cu lt to s a y P rob a bly

.
t h es e sev e ra l fa ct or s Op e r a t e i n a l l v a r i eti es of p rop orti on .
T h a t th e h a bit u a l s h a di n g of s om e p ar ts by oth e r s l a r ge ly
a id s i n c a u s i n g th es e div er g en ce s from s y m m etry is v ery ,
i nst ru ctiv ely sh own by th e com p oun d l ea v es of th e Cow

p arsnip Fig 2 1 1 rep resent s one of th ese Whil e th e l ea f a s a
. . .
wh ol e i s bil a t erally sym m et r ic a l ea ch of the w i ngs h as an u n

,
s y m m et r i ca l bil a t e r a l n e s s : th e s id e n ext th e a xi s b ei n g l a rg e r
t h a n th e rem oter s id e . H ow d oes this h a pp en ? Fig 2 1 2 .

,
which i s a di a g r a m m a ti c secti on d own t h e mid ri b of the -
l ea f sh owi ng its i ncli ned a tti tu d e a nd th e p os iti ons of the

,
THE S H A P E S OF L E A VE S .
1 55
wi ngs a b 0 will m a k e the c au s e cl ear A s th e wi ngs

, , , .
ov er l a p ,
lik e th e b ars of a Veneti an bli n d ea ch i nt ercep ts ,
s om e light from the one b el ow i t ; a nd the one b el ow it

th u s su ffers m or e on i t s di st a l s id e th a n on i t s p roxi m a l s id e .
H ence th e s m a ll er d e v el op m en t of the di st a l s id e T h a t this .
i s th e c au se i s fu rth er s h ow n by th e p r op orti on th a t i s m a i n
t a i ned b et ween th e d eg ree of ob scu r a t i on a nd th e d egree of
n on d ev el opm ent ; for thi s u n lik e n ess i s g rea t e r b et ween the
-
'
tw o s id es a a n d a th an b etw een b a n d b or c an d c a t the
’ '
, ,
s a m e ti m e th a t th e i nterfe r e n ce i s g rea t er i n th e l owe r w i n g s
th an i n the u pp er Of cou r se i n thi s c a se a nd i n th e ki nd red

.
c a s es h erea ft er s i m il a r ly i n t er p re t ed it i s n ot m ea nt th a t
,
thi s di fferenti a ti on i s c on s eq u ent s o l ely or even chi efly on , ,
th e di ffer enti a l a c ti on s e xp er i en ce d by th e i n divid u a l pl a nt .
T h ou gh th e r e i s g ood r ea s on t o b eli ev e th a t the ra t e of gr ow th

i n ea ch p a rt of ea ch l ea f i s a fi ected by th e i nc id enc e of light ,
et c ontra st s s o m a r k e d a nd so sy s t em a tic a s th e s e a r e not

y
e xplic a bl e wi th ou t t a ki n g i nt o a ccou n t th e i n h e r it a n ce of
modi fic a ti ons e ith er fu n cti ona lly c au s ed or c au s e d by s p on

t a neou s v a r i a ti on C l e a r ly th e t en d en cy w ill be t owa r ds
.
,
the p res erv a ti on of a pl a n t whi c h di s t r ib u t es i ts chl orophyl l

i n the m os t a dv a nt a geou s w ay a nd h ence th er e w ill a l w a y s be
a gr a vi t a ti on t ow a r d s a f or m i n w hich sh a d e d p a r t s of l ea v e s
a r e u n d e v e l op e d .
§ 229 F rom com p ou nd l ea v e s to s impl e ones w e fi nd

.
,
t ran s iti on s i n l ea v es of whi ch the divi si on s ar e p a rti a l i n

s t e a d of t ot a l ; an d i n th e s e we s e e w ith e qu a l cl ea r ne s s t h e
, ,
r el a ti on s b et w een form s a n d p ositi on s th a t h a v e b e en t r a ce d
th u s fa r Fig 2 1 3 i s th e l eaf of a Wi nter aconit e i n whi ch

. .
-
,
rou n d a v e rt ic a l p eti ol e th er e i s a r a di a l di s t r ib u ti on of h a lf
,
s ep a r a te d l ea fl et s T h e C ecr opi a —l ea f Fig 2 1 4 s h ow s u s a

.
'
,
.
,
t wo s id e d d e v e l op m en t Of th e p a rt s b egi nn i n g to m odi fy
-
,
bu t n ot obli t er a ti n g th e a l l sid e d a rr a ngem en t ; a n d thi s

,
-
mix ed s ym m et ry oc c u r s u n d er c on diti on s th a t a re i nterm e

di a te A m ore m ark e d d e gree of the s a m e rel a ti on is pre
.
s ent ed i n the l ea f of the L a dy

’
s Mantl e Fig ,
. 21 5 . A nd
then i n th e S yc a more a nd th e Vi n e, w e ha ve a cl e ft typ e of
l ea f i n w hi c h a d ecid e d bil a t era l ness of f or m co exi s t s w i th a -
d ecid e d bil a tera l ness of condi ti ons .
Th e qu it e s i m pl e l ea v es to w hi c h w e now d esc end exhibit , ,
v ery di sti n ctly a p a ra ll el s er i es of facts Wh ere th ey g row

,
.
u p on l on g a nd com pl et ely i nd ep end en t f oo t st a lk s

-
with ou t -
,
d efi n it e su b or di nati on to som e cen t r a l v e r ti ca l a xi s th e ,
l ea v es of w a t er pl ants a r e s ym m et r ic a lly p el ta te
.
-
Of this .
th e s a c re d I n di a n b ea n Fig 2 1 6 fu rni s h es a n e x a m pl e
-
,
.
,
H er e .
th ere i s only a trace of bil a t era l n ess i n the v enati on of th e

l ea f corres p ondi ng to the v ery s m a ll di fferenc e of th e con
,
di ti on s on the p roxim a l a n d di s t a l s id es I n th e Vi c tor i a .
r egi a ,
Fig 2 1 7 the foot sta lk s th ou gh ra di a ti ng a l m os t
.
,
-
,
h ori zont a lly f rom a c en t re a re s o l ong a s to k eep th e l ea v es

,
qu i te rem ote from on e a noth er ; an d i n it ea c h l ea f i s a lm os t

s y m m et r i ca lly p e lt a t e w ith a bil a t e r a l nes s i n dic a t e d on ly by
,
a s ea m ov er th e li n e of th e foot s t a lk T he l ea v es of th e
-
.
N ym p haea Fig 2 1 8 m or e c l os ely c l u s t ered a nd ha vi n g l es s

, .
, ,
r oom t ra n sv ers ely t h a n l ongit u dina lly exhibit a m a rk ed ,
a dv a n c e to th e tw o s id e d for m ; n ot only i n the e xc e ss of

-
the l ength ov er th e b rea dth bu t i n th e exi st ence of a cl eft

, ,
TH E S H A P E S OF L E A V E S .
1 57
wh ere i n th e Vi ctor ia r egi a th ere i s m erely s e a m A m ong

a .
l a nd —pl a nts simil a r form s a re fou n d u n d er a na l og ou s con di
ti on s . T he co mm on H ydr ocotyl e ,
Fig 2 1 9 which s ends
.
,
up di rec t from its root s a few a lm os t u p right l ea f s t alk s has -

,
thes e s u r m ou nt e d by p elt a t e l ea v es ; which l eav es h owev er , ,
diverge s lightly from ra di a l symm et ry i n c orresp ond ence w ith

th e s light cont ra st of c i r c u m st an c es w hi c h th ei r g rou pi n g i n
v olv es A noth er c ase i s su ppli ed by the N a stu r ti u m Fig
.
,
.
2 20 whi c h combi nes th e ch a ra ct ers— a creepi ng s t em l ong

, ,
l ea f s t a lk s growi ng u p a t right a ngl e s t o it a n d u n sym m e

-
,
t ri c a l l y p elt a t e l e a v e s of which the l ea s t dim ens i on i s on

, ,
the a v era ge t owa rds th e s tem

,
B u t p erh a p s th e m os t
.
st r iki ng ill u st ra ti on is th a t fu rni s he d by the C otyl edon u m bi

l i cus Fig 2 2 1 i n whi ch di ffer ent ki nds of symm et r y occ u '
,
.
,
r
i n th e l ea v es of th e s a m e pl a n t a l on g w ith di fferen c es i n th ei r
,
r e l a ti on s to c on di t i on s T he r oot —l ea v es a grow i n g u p on
.
, ,
v erti ca l p eti ole s b efore the fl ower st a lk m a k es i ts app ea r a nce

-
,
a r e s y m m e t r ic a lly p el ta t e ; w hil e the l ea v es w hi c h su bs e

qu ently g row ou t of th e fl ower s t a lk b a r e a t th e b ott om
-
, ,
t ra n s iti on a lly bil a t era l a nd high er u p compl et ely bil a t era l

,
.
T h a t the bil a t era l f or m of l ea f i s the ordi na ry form ,
c orres p on d s w ith th e f a ct th a t or di na r ily

,
th e c i r cu m ,
s t a nc e s O f th e l ea f a r e diff er en t i n th e di recti on of th e pl a nt s
’
a xi s fr om wh a t th ey a r e i n th e opp os it e di recti on whil e ,

1 58 M ORP H O L OGI CA L D E VE L O P ME NT .
t ra n sv ers ely the ci rcu m s ta nces a re a lik e I t i s n eedl es s t o

.
giv e di a g r a m s to illu s tr ate thi s ext rem ely fa m ili ar tru th .
Wh eth er th ey a r e b roa d or l ong ov a l or h ea r t sh a p ed p oi nte d

,
-
,
or obt u se th e l ea v e s of m os t t r ees a n d pl an t s w ill be re m e m

,
b ered by a l l a s h a vi n g the e nd s by whi ch th ey a re a tt a ch e d

u nlik e th e fr ee en d s w hil e t he tw o s id es a re a lik e
,
A n d it w ill .
a l s o be r e m e m b e r e d th a t th e s e e qu a li t i e s a n d i n eq u a liti es o f
d ev el opm en t corres p on d w ith the equ a liti es a nd i nequ a li ti es

i n the i nc id enc e Of forc es .
§ 23 0 A. c on fi rm a ti on th a t i nt eresti n g and imp ort a nt

is ,
is fu rni sh ed by th e ca ses i n which l ea v es p resent u n sym m e

t ri ca l for m s i n p os iti on s wh er e th e i r p a rts a r e u n sym m etri
c a lly rel a ted to the e nvi ron m en t A con s id e ra bl e d evi ati on
.
f rom bil a te ra l sy m m etry m ay be seen i n a l ea f whi ch h a bi tu

a lly s o c a r ri es it s el f th a t t h e h a l f on t h e o n e s id e o f t he m i d—
,
ri b
i s m ore s h a d e d t h a n th e ot h er h a l f T h e d r oopi n g b ra n ch es o f
.
th e L im e d e li n ea t ed i n Fig 2 22 sh ow u s l ea v es s o a rra ng ed
, .
,
a nd so m odifi e d . x a mi ni n g th ei r a ttit u d es a nd th ei r
On e
re l a ti on s on e to a n oth e r it will be f ou n d t h a t ea ch l ea f i s s o
,
i n cli ned th a t th e h a l f of i t n ext the s hoot g rows over th e

s h oot a n d g et s pl en ty of light ; w hil e th e oth e r h a lf s o h a n g s
d own th a t i t com es a good d ea l i nto th e s h a d e of the p re

c edi ng l ea f T h e res u lt i s th a t h a vi ng l ea v es w hi ch fa ll i nt o
.
th ese p os iti ons the sp eci es p rofit s by a l arge d ev el opm ent of

,
th e e xp osed h a lv es ; a nd by s u rviv a l of the fit test a c ti n g ,
a l on g w i th th e di rec t ef fec t of ext ra e xp os u re thi s m odi fi ,
ca ti on b ec om es e st a bli sh e d H ow u n qu es ti ona bl e i s the

.
.
c onn exi on be t ween th e re l a tiv e p os iti ons o f th e h a lv es an d
th ei r r el at i v e d ev el opm en t s w ill be a dmitt ed on ob servi ng a

,
THE S HA P E S OF L E A V E S .
1 59
c on v erse c a se . Fig . 2 23 re p res ents a s hoot of S tr obi l a n thes

g l o m er a tus . H ere th e
l ea v es a r e so s et on t he s t e m th a t t he
i nner h a l f of ea c h l ea f i s s h a d ed by the s u b seq u ently form ed -
l e a f whil e i ts ou t er h a l f i s not th u s s h a d ed ; an d h e re we fi n d
,
th e inn er h a l f l es s d ev el op ed th a n the ou t er h a lf Bu t th e .
m os t c on c l u s iv e evid en c e of thi s r e l a ti on b etwe en u n sym m e
t r i ca l f orm a nd u ns y m m et ri ca l di s t r ib u ti on of s u rr ou n di ng
forc es is su ppli e d by th e g enu s B eg oni a ; for i n it we h a v e
,
a m a n i fe s t p rop orti on b e t w e en th e d e g ree Of the a ll eg e d
e ffect a n d the d e g r ee o f th e a ll eg e d c au s e T h ese pl a nts .
p rod u ce th ei r l ea v es i n p a i rs i n su ch way s th at the c onna t e ,
l ea v e s i nt er fere with one a noth er m u ch or littl e a ccor di ng ,
a s th e f oot — s ta lk s a re s h ort or l ong ; an d th e res u lt i s a cor
re l a tiv e div er g en c e f rom sy m m e t r y I n B eg oni a n el u m bii .
fol i o whi ch h a s p eti ol es so l ong th a t the c onna t e l ea v es a re not

,
k ept cl ose tog eth er the re i s bu t li ttl e d evi a ti on f rom a bil ater
,
a lly p elt a t e fo r m ; w h erea s

-
a cco m p a n yi ng th e com p a r a tiv ely
,
m a rk ed a nd c ons t a n t p roxim i ty i n B p rui na ta Fig 224, we .

, .
se e a m ore d ecid edly u nsym m et ri ca l s h a p e ; a n d i n B .
m a hr i ngi i Fig 2 2 5 th e m odifi cati on th u s c au s ed is p u sh ed

,
.
,
s o fa r a s to d e s t r oy th e p el t a t e s t ru ct u r e
*
.
23 1 . A g a i n, th en we a re t au ght th e sa m e t ruth H ere

, .
,
as b efore ,
we s ee th a t hom ol ogou s u nit s of a ny or d er b ecom e
We m ay note th a t s om e of th e s e l ea ves ,
as th os e of th e L i m e, fu rni sh
i ndi c a t i ons of th e ra ti o wh i ch e x i s t s b e t w ee n t h e fects of
ef i nd i v i dua l c i rc um
s ta nces a nd thos e of t y p i cal t e ndenc i e s . On th e one ha nd, th e l ea v e s borne

by th ese droo p i ng br a nch e s o f t he L i m e a re wi t h h a r dl y an e cep t ion u ns vm x
v
m etri ca l m ore or l es s dec i ded l y, e e n i n p os i t i on s w h e re th e ca u s e s of u ns vm
m e try a r e not i n a c t i on : a fa ct sh ow i ng u s t h e re p et i t i on of t h e typ e i rr e s p e c
v
ti e of th e condi t i on s . On t h e o th e r h a nd, t he d egree of de vi a ti on from
s ym m e try x tre m el y v a ri a bl e e v en
is e ,
a fa ct p rov ing th a t
on t h e s a m e s h oo t :
t he ci rcu m st a nc e s of th e i nd i v i du a l l e a f a re i nfl ue nti a l i n m od ifyi ng i t s form .
B ut th e m os t s t ri k ing e v i dence of th i s di rect m odi fica ti on i s a fl orde d by t he ’
s u ck e rs o f t h e L i m e Gro wi ng a s t h e s e do in a pp rox i m a te l y u prig h t a t ti

.
, ,
tude s th e l ea v e s th ey b ea r do not s t a nd to one a noth e r i n th e wa y a b ov e

,
d e s c ri be d, a nd th e ca us e s o f un s v m m e try a re no t i n ac t i on a nd h ere, th ough

th e re i s genera l l ea ni ng t o th e uns ym
a m et ri ca l form ,
a l a rge proport i on of th e
l ea v es b ecom e q u i te s ym m e tri ca l .
1 60 MOR P H OL OGICA L D E V E L OP ME NT .
di fferenti at e d i n p rop orti on a s th ei r rel ati ons to i nc id ent

f orces b ecom e di fferent A nd h ere a s b efore w e see t h at i n
.
, ,
eac h u ni t c on s id e r e d by i ts el f th e di ffe r ences of di m en s i on

, ,
a r e g rea t es t i n t h os e di rect i on s i n which th e p a rt s a r e m o s t
di ffe rently c ondi ti on ed ; whil e th ere a r e n o di fferen ces be

tween th e dim ens i ons of the p a rt s th a t a r e n ot di fi er entl y
*
c ondi ti oned .
I t was by a n Ob s erv a t i on on th e form s of l ea v e s , th a t I w as fi rs t l ed to

th e vi e ws s e t forth i n th e p re ce di ng a nd s uccee di ng c ha pte rs on th e m or .
ph ol ogi ca l di fferent i a t i on of pl ant s a nd an i m a l s In th e yea r 1 8 5 1 , du ri ng .
a count ry ra m bl e i n whi ch th e s t ru c tu re s of
pl a nts h ad b ee n a top i c of c on
v ers at i on w i th a fri e nd— Mr . G . H L e we s— I h a p pened t o p i ck u p t he l ea f
.
of a b ut terc u p, d ra wi ng i t by i ts foot-s ta l k t h rou g h m y fi ngers s o as t o

a nd,
thr us t togethe r i ts d ee pl y-c l eft di v i s i ons , ob s e rv ed t h a t i ts p a l m a te a n d a l m os t

ra di a l form was ch anged i nto a bi l a te ra l one ; a nd th a t w ere th e di v i s i ons t o
grow t ogeth er i n t hi s new p os i t i on, an ord i na ry b i l a tera l l e a f w oul d res ul t .
Joi ni ng thi s o bs erv a ti on w i th t h e fa m il i a r fact tha t l ea v e s , i n c om m on wi th
th e l a rger m em ber s of pl a nt s , h abi tu a ll y t urn th em sel v es to th e l i ght, i t

occ urred to m e tha t a na t u ra l c h a nge i n th e ci rcu m s ta nce s of th e l ea f m i gh t
re a di l y c a u s e s u ch a m od ifica t i on o f form as th a t wh i ch I h a d p rod uced a rt i
fi ci a l l y . I f, as t he y o fte n do w i th pl a nt s , s oi l a nd c l ima t e were gre a tl y to

ch ang e th e h a b i t of th e bu tte rcup, m ak i ng i t b ra nch ed a n d sh rub-l i ke ; a nd if
t he s e p a l m ate l ea v e s w e re t hus v ; w ou l d
m uch o e rs h a d ow ed by on e a nother
v
not th e in ne r segm e nts of t h e l ea e s g ro w to w a rds th e p e ri ph ery o f t h e pl a nt
wh ere th e l i gh t w a s g re a test , a nd so ch a nge the p a l m a t e form i nt o a m ore

dec i ded l y bi l a te ral form Im m e di a t el y I be gan to l ook round for e v ide nce of
th e re l a t i on be t ween th e form s of l e a v e s a nd t he genera l ch a ra cte rs of th e
p l a nt s t h ey b el on g t o ; a n d soo n found s om e s i g ns o f conne i on C erta in x .
v
a no m a l i e s, or se em i ng a no m al i e s , h ow e er, p re ented m e fro m th en pu rs ui ng v
q
t he i n ui ry m u ch fur th e r B ut cons ide ra t i on c l ea red up th es e diffi cul ti es ;
.
and the i d ea a fte r wa rd s wi dened i nt o th e g ene ra l doct r i ne h e re el abora te d .
Oc cup a t i on wi th o th er v
t hi ng s p re ente d m e from gi v i ng e xp re ss i on t o th i s
d oc tri ne u nt i l J a n 1 8 5 9 ; wh en I publ i shed an o ut l i ne of i t i n th e

general .
C H A P TE R X .
TH E SH A P E S OF F L O WE RS .
23 2 FO L L O W I N G a n ord er lik e th at of p receding ch ap

.
t ers l et u s fi rs t not e a few typic a l fa ct s res p ecti n g the form s of

,
c l u st e r s of fl ower s a p a rt f r om th e f orms of the fl owers th e m

,
se lv e s . T wo ki n d r e d ki nd s of L ega m i nos ce s erv e to S h ow how

the m em b e r s of c l u s t er s a re di st r ib u t ed i n a n a l l s id e d m a nne r
-
or i n a two s id e d m a nn e r
-
a cc or di n g a s th e ci rc u m st a nc e s
,
a re a lik e on a l l s id es or a lik e on on ly two sid es I n H i pp o.
c r ep i s r ep r ese nte d i n Fig 22 6 th e fl owers g rowi ng a t the end

,
.
,
o f a v er ti c a l s t e m a re a r r a n g e d
,
r ou n d i t i n r a di a l s y m m et r y .
C ont r a r i wi s e i n M el i l otu s Fig , .
2 2 7 w h ere the a xill a ry s t em

,
b ea ri ng th e fl owers i s s o
pl a ce d i n rel a ti on to th e m a i n
s tem , th a t i ts ou t er and i nn er
fa ces a re di fferently c on di
ti oned th e fl ow ers a r e a l l on
,
the ou t er fa ce : th e cl u ste r is
bil at er a lly sy m m etri ca l s i n ce ,
i t m a y be cu t i nt o a pp r ox
i m a tel y equ a l an d s i m il ar
grou p s by a v erti ca l pl a ne p a s s i n g th r ou gh m a i n a xi s .
P l a nts o f thi s s a m e t r ib e fu r ni sh c l u ster s o f i nt er m edi a t e

ch a r a ct e r s h a vi n g i nte r m e di a t e con di ti on s A m on g th ese ,
.
a s a m on g th e c l u st ers w hi c h oth er typ es p r esen t m a y be

57
f ou n d s om e i n whi ch c on form ity to th e g ener a l l aw is not

obvi ou s T he di s cu s si on of th ese a pp a ren t a n om a li es wou ld
.
c a rry u s too m u c h ou t of ou r c ou rs e A cl u e to the expl a .
n a t i on of th e m w ill I b eli ev e he fou n d i n th e e xpl a na ti on

, ,
p r esen tly to be giv en of cert ai n ki nd red anom a li es i n th e

f orm s of i ndividu a l fl owers .
§23 3 .di a lly sy m m et r i cal for m i s com m on to al l

T he ra -
i n dividu a l fl ow ers th a t h a v e v ertica l a x es I n pl a nt s whi ch .
a r e p r ac ti c a lly if not litera lly u ni a xi a l a nd hear th e i r fl ow e rs ,
a t th e en ds o f u p r ight s t a lk s s o t h a t th e faces Op en h or i
,
z onta l l y th e p eta ls a re di s p os e d i n an a l l s id ed w a y
,
C ro -
.
c u ses T u lip s a nd P oppi e s are f a mili ar e x a mpl es of this

, ,
s t ru ctu re occu rr i ng u n d e r th ese c on di t i ons A Ra nu nc u l u s .
fl ower Fig 2 2 8 w ill s erve a s a typi ca l

,
.
,
one S i m il a r ly fl owers whi c h h a v e

.
,
p ed u n cl es fl exibl e en ou gh to l et th e m
h ang di rectly d ownw a rd s and a re not ,
l at era lly i n comm od ed a re al so r a di a l ; ,
a s i n the F u c hs i a Fig 2 2 9 a s i n Oyol a ,

.
,
m en H ya ci n th &c T h es e r el ati ons of

, ,
.
form to p os iti on a re I b eli ev e u n i form T h ou gh som e fl owers

, , .
ca rr i e d at th e en d s of u p right or d ow nr ight s tem s h av e

obliqu e s h a p es it i s only wh en th ey h a v e i ncli ned a x es or
,
a r e not eq u a lly c ondi t i one d a l l r ou n d N o s olit a ry fl ow er .
h a vi ng an a xi s h a bi tu a lly v ertic al p res ents a bil a t era l for m ,

.
T hi s i s a s w e s h ou ld exp ect ; s i n ce fl owers whi c h Op en ou t

th ei r faces h ori zont a lly wh eth er f aci ng u p w ar d s or d own
,
w a rd s a re on the a v era ge s imil arly a ffect ed on a l l Sid es

, , ,
.
A t fi rst it seem s th a t fl ower s th u s pl a ce d s h ou ld a l one be

r a di a l ; bu t f u rth er c on s id era ti on di sc l o s e s con diti ons u nd er
w hi c h thi s typ e of s ym m etry m a y e xi st i n fl ow er s oth erw i s e

pl aced Rem em b e ri ng th a t th e r a di a l f orm is the p r i m i tiv e
.
f or m — th a t m orph ol ogi ca lly s p ea ki ng it resu l ts f rom th e

, ,
c on tra ct i on i n to a wh or l o f p a rt s th a t a r e or igi n a lly a rr an g e d

,
i n the s a m e s pi ra l s u c c es s i on a s th e l eav es ; we m u s t e xp ect

T HE S H A P E S OF F L OWE RS .
1 63
it to ti nu e wh erev er th ere a re no for ces t endi ng to ch ange

c on
it . Wh at now m u s t be the forc es t ending to ch ange it ?

T h ey m u s t be forc es which do n ot simply a f fect di fferently
th e diffe rent p a rts of a n i n divid u a l fl ower T h ey m u s t be .
for c es whi c h a ffec t i n lik e contras ted w a y s the h om ol ogou s

p arts of oth e r i ndivid u a l fl owers both on the s a m e pl ant a n d ,
on s u rrou ndi n g pl ants o f the s a m e sp eci es A p erm a n ent .
m o difi c a ti on c a n be exp ec t e d on ly i n c a s es wh ere by i nh eri t ,
a nc e th e ef, f ects of the modi fyi ng c au se s ac cu m u l a t e T h at .
th ey m ay a cc u mu l at e the fl owe rs m u st k eep th em selv es so

re l a t e d to th e en vi ronm ent th a t th e h om ol og ou s p art s m ay
, ,
g enerati on a fter g enera ti on be s u bj ect e d to like diff erenti a ti ng

,
f orc es H ence am ong a pl ant s fl owers whi ch m ai nt a i n no

.
,
’
u n i formity i n th e re l a ti ons o f th e i r p a r t s to s u rrou n di ng i n
fl u ences th e ra di a l form will c onti n u e L et u s gl a nc e a t the

, .
s e v era l c a u s e s whi c h en t a il thi s v a r i a bility When .
fl ower s a re b orne on m a ny b r a nch es w hi ch h a ve al l in cli n a ,
ti ons f rom the v ertic a l to the h ori zon t a l— a s a re the fl owers

of th e A ppl e th e P l u m ,
th e H a w th or n— th ey a re pl ace d i n
,
c ou ntl es s di f f erent a ttitu d es Consequ ently any sp ont an eou s

.
,
v a ri a ti on i n s h a p e whi ch might be a dvan t ageou s were the

a tt it u d e c on st a nt is not lik ely to be a dv ant a geou s ; a nd a ny
,
fu ncti ona lly p rod u c e d m odifi c a ti on i n one fl ower is lik ely to

-
,
be n eu t r a li ze d i n offs p ri ng by som e opp os it e f u ncti on a lly pr o -
du ced m odifi c a ti on i n a noth er fl ower I t is q u it e c omp re .
h en s i bl e th e re fore th a t i rre gu l a r ly b r a nc h e d pl a n t s s h ou ld
, ,
-
thu s p reserv e the i r l a t erally b orne fl ower s f rom u nder -
goi ng p er m a n ent d evi a

ti on s f rom th ei r p rimi
tive r a di a l s ym m et ry .
Fig 2 3 0 r ep r es enti ng a
.
,
bl ossom i ng t wig of the

Bl a ckth orn ill u st r at e s
,
th i s . A ga i n u p r ight ,
p an icl es su ch a s th ose of th e S a xif ra ge ex em plifi ed i n

,
Fig 2 3 1 and i rregu l ar t er mi nal g rou p s of fl owers oth er

.
,
w i se n a m ed , fu rn i s h diti ons u nd er which th ere is s i m i

c on
l a rl y a n a b senc e of d eterm i na t e r el a ti ons b et ween the

p a rt s of th e fl owers a nd the i ncid ent forc es ; and h en ce an
a b s en c e of bil a t e ra l nes s . T hi s i ncon stancy of rel a
tiv e p os iti on is p rod u c e d i n v a r i ou s oth er wa y s— by ext rem e
fl exibility of the st em s as i n the B l u e—b ell ; by the ten
,
'
d eney of the p ed u ncl e s to c u r l to a g rea t er or l ess ext ent

i n div er se di r ecti ons a s i n P yr ol a ; by s p ec i a l t wi s ti n gs of
,
t he p ed u nc l es di ffer i n g i n d eg ree i n di fferen t i n divid u a l s

, ,
’
a s i n C on v ol vu l u s ; by u nu su a l l a xity o f th e p et a l s a s i n ,
L y thr u m . E l s ew h ere th e lik e gener a l r es u lt a r i s es f ro m a

p rogres s iv e ch a nge of a ttit u d e a s i n Myos oti s the st em of
, ,
which a s it u n fold s c au ses ea ch fl O wer to u nd ergo a t ra n s iti on

f rom an u p war d p os iti on of the m ou th to a l a te ral p os i ti on ;
or a s i n m os t C r u ci fer ce wh er e th e lik e e f
,
fec t foll ows f rom an
a lt ere d di recti on of th e p ed u nc l e .
T h ere a re h ow ev er cert a i n s eem i ngl y a n om a l ou s c a se s

, ,
-
wh ere ra di a l sy m m e try i s m a i n t a i n e d by l a t e ra lly pl a ced -
fl ow er s whi ch k eep th ei r p a rts i n rel a tiv e p os iti ons th at are

,
t ol era bly c ons t a n t

. The expl a n a ti on o f th es e e x cepti on s i s
n ot m a nif es t. I t i s only wh en we t a k e i nt o accou nt c ert a i n
i ncid ent a cti on s li a bl e t o be l eft u nr em em b ered th a t w e fi nd ,
a p rob a bl e sol u ti on I t w ill be m ost conv en i en t to p ostp one

.
the c ons id era ti on of th ese ca s es u n til we h a v e reacli ed th e

g enera l ru l e to whi ch th ey a re ex cepti on s .
§ 2 3 4 T ransiti ons v aryi ng i n d eg ree f rom the r a di a l to

.
wa rds the bil a t era l a re c o mm o n i n fl owe r s th a t a re b orne a t

,
th e en d s of b ra nch es or a x es w hi c h a re i nc li ne d i n t ol era bly

c onst an t w a y s .We m ay s ee thi s i n sund ry ga rd en fl owe rs
s u ch a s P etu nia or su ch as ,
I s ol om a a nd A c hi m en es ,
h own i n Fig s 2 3 2 a n d 2 3 3
s . .
I f th es e pl a nt s be e x a m i ned ,
it will be p erceive d th a t the

m o e of
d g rowth m ak es th e fl ow er u nfold i n a p a rti ally one
THE S H A P E S OF FL OWE RS .
1 65
sid e d p os iti on ; th a t i ts p a rt s of a tt achmen t h a ve r igidi ty

su ffici ent to p re v en t thi s a ttitu d e from b e i ng v ery m u c h i n
te r fered w i th ; a n d th a t th ou gh the i n divid u a l fl owe rs v a r y
som ewh a t i n th ei r atti tu d es th ey do not v a ry to the e xtent of
,
neu tra li z i ng the di fi erenti ati ng c on di ti on s — th e re re m a i ns a n
a v e r a ge div erg en ce f rom a h or iz ont a l un f oldi ng of the fl owe r ,
t o a c c oun t for i t s div er gence f ro m ra di a l s ymm et ry .
We p a s s i nsens ibly from for m s lik e the se to forms h a vi ng ,
bil a t era l symm et ry s t rongly p ronounced S om e s u ch f orms .
oc c u r a m ong fl ow er s th a t g row a t th e en ds of u p r ight s t em s ;
a s i n P i ng ui cu l a a n d i n th e Vi o l et t ribe
,
B u t t hi s h a pp en s .
on ly w h ere in su ccess iv e genera ti on s the fl ower u n fold s its

, ,
p a rt s s id ew ays i n const ant re l a tiv e p os iti ons A n d i n the .
i m m ens e m a j ority of fl owers h a vi ng well m a rk ed two s ide d - -
form s the h a bitu a l exp osu re of th e di fferent p a rt s to different

,
s et s of forces i s eff ect u a lly s ecu red by the m od e of pl ac i ng

, .
A S ill u s t ra t i on s I m a y na m e the gen er a— Orchi s U tri cul a ri a

, , ,
S a l via , S a l i x , D el p hi ni u m , M en tha , T eu cr i u m , A ju ga , Ba l l ota ,

Ga l e op si s , L a m i u m , S ta c hys , N ep eta , Ma rr u bi u m , C a l a m i n tha ,
Mel i ttis , P r u nel l a , S cu tel l a ri a , Ba r ts i a , E up hra s ia , Rhi na u
th us , Mel a m pyru m ,
P edi cu l a ri s , L i n a r ia , D i gi ta l i s , Or c
ba n che, F u m a ri a , rei n; t o w hi c h m ay be a dd ed al l th e Gra s s es
a nd a l l th e P a pi l i ona cece In
t of th es e ca s es t he fl owers
. m os ,
being s es s il e on th e s id e s
o f u p r ight s t em s a re k ept i n qu it e ,
fix ed a ttit u d es ; a nd in t he oth er c a ses the p ed u ncl es a re v ery

sh ort or e l s e sti f
,
f enou gh to s ecu r e gen era l u n i form ity i n the
pos iti ons A few of the m ore m ark e d typ es a re sh own i n
.
Figs 2 3 4 to 2 4 1
. .
23 5 23 7 24 0 241
Very i n st ru ctiv e evid enc es h er e m eet us . S om etim e s withi n

th e limit s of genu s we fi n d r adi a l fl ower s bil a t era l
one ,
fl owers , a n d fl ow ers o f i nt er m edi a t e ch a ra ct ers The g enu s .

1 66 M ORP H OL OGI CA L D E VE L OP ME NT .
B eg oni a m ay be i n s t a n ce d . In B r i gi da
fl owers v ar i ou s
. the ,
i n th e i r a tti tu d es a r e i n th ei r m or e c on s pi cu ou s c h arac ter s

,
r a di a l : th ou gh th er e i s a c ert a i n bil a t e r a l ness i n th e c a lyx ,
the fi ve p et a l s a r e sy m m etri ca lly di s p ose d a l l r ou n d B . .
Wa g en eri a na f u rni s h es two f orm s of fl owers On the sa m e i n .
divid u a l pl a nt m ay be f ou n d ra di a l fl ower s lik e Fig 2 4 2 and .

,
oth e r s ,
lik e Fig 2 4 3 whi c h a re m er gi ng i n t o the bil ater a l
.
, .
More d ecid e d i s the bil a ter a l n ess i n B a l bo cocci n ea Fig 2 44 ; .

-
, .
a n d s till m or e i n B ni ti da Fig 2 4 5 . Whil e i n B hera cl ei

, . . .
fol i a Fig 2 4 6 the ch ange rea ch es its ext rem e by the di s

,
.
,
a pp ea r a n c e of t h e l a t era l p eta l s On e x a m i ni ng th e m od es of
.
g rowt h i n th ese sev era l s p ec i es th ey will be seen to expl a i n

,
th es e ch an ge s i n the m a nner a ll ege d E v en .
m ore c oncl u s iv e a re the nea rly a lli e d t ra ns for m a ti ons occu r

-
r i n g i n a rti fi c i a l l y p rod u ce d v a r i e ti es o f th e s a m e s p ec i es
-
.
Gl oxi ni a m ay be n a m e d i n ill u s t ra ti on I n Fig 2 4 7 i s repre . .
s ent ed one of the or di na ry forms w hi c h sh ow s u s bil at era l ness

,
of sh a p e a l ong w ith a m od e of g r owth th a t ren d e rs th e c on di
ti ons a lik e on th e two sides whil e di fferen t a b ov e and b el ow .
Bu t i n G er ec ta Fig 2 4 8 we .
, .
,
h a v e th e fl ower a ssu m i ng a n
u p r ight a tti tu d e a n d a t th e ,
s a m e tim e a ss u mi ng the r a di a l
typ e T hi s i s not to be i nter

.
r ete d as a p r od u cti on o f ra
p
di a l symm et ry ou t of bil a t era l s y m m et ry u n d er the a cti on of ,
the a pp r op ri a t e con di ti on s I t i s ra th er t o be t a k en a s a ca s e
.
of wh a t is t er m e d p el ori a —a r ev ers i on to the p ri m itiv e

“
r a di a l t yp e f rom whi ch the bil a t era l m odifi ca ti on h a d b ee n

,
d eriv ed T he sign ifi cant i n fer ence to be d rawn f rom it i s

.
,
T HE S H A P E S OF FL OWE RS .
1 67
th a t this p r i m itiv e ra di a l typ e h a d a n u p r ight a ttit u d e ; a nd

th a t the d eriv a ti on of a bil a ter a l ty pe f rom it occ u rre d a l ong ,
with the a ssu m pti on of a n i ncli ned a ttit u d e .
We com e now to a grou p of c a ses a b ov e referred to i n ,
which ra di a l symm et ry c onti nu es to co exi s t w ith th a t con -
s ta n t l a t era l a ttit u d e or di n a r ily a cc omp a ni e d by t h e two
s id e d f orm . T wo e x a mpl es w ill su ffi ce : on e a v er y l ar ge

fl ower the H ollyh oc k and the oth er a v ery s m a ll fl ower the
, , ,
A gr i m ony Why d oe s th e ra di a l for m h ere rem a i n u nch a nged ?

.
a nd how d oes i ts c onti nu a nc e c ons i s t w i th th e a ll eg ed g e nera l
l aw ?
Until q u it e c tly I h a ve be en u n abl e to fi nd a ny p rob
r e en
a bl e a nswe r s t o t h e s e q u es ti on s Wh en the di ffi cu lty fi r st

.
p resent e d i ts el f I c ould thi nk of no oth er p oss ibl e ca u s e for

,
th e a n om a ly t h a n th a t the p a r ts of th e H ollyh ock fl ower

,
-
,
u n foldi n g s pi r a lly as th ey do might h a v e di ffer e nt d eg ree s

,
o f s pi r a l t wi s t i n di f ferent fl ow e rs a nd might thu s not be

,
u n fold e d i n su f fi ci entl y con sta n t p os iti ons

-
B u t this s eem e d
.
a q u es ti on a bl e i nt erp r eta t i on ; a nd on e which did not Oh
vi ou sl y a pply to th e c a s e of th e A gr im ony I t wa s only on

.
i nq u i ri ng wh a t a re the sp eci a l c au s es of m odific a ti ons i n the

form s of fl ower s th a t a m ore fea s ibl e expl an a ti on su gg est e d
,
i tsel f ; a nd thi s wou ld p rob ably n ev er h a ve s u gges t e d it sel f ,
h a d not Mr D arw i n s i nv e stig a ti on s i nt o th e f ertili z a ti on of

’
.
Orchid s l ed m e to t a k e i nt o a c cou nt a n u nnotic e d a gency .
T he a cti ons whi c h a ff ect th e for m s of l ea v es a ffect m u ch ,
l ess d ecid edly the f or ms of fl ow er s ; a n d th e forms of fl owers

a r e i nfl u e n c ed by a cti on s whi c h do not i n fl u en ce th e f or ms of
l e a v es P artly th r ou gh th e di rect a cti on of i ncid en t f orces

.
a n d p a r tly th rou gh th e i n di re c t a cti on of na t u r a l s el ec ti on

,
l ea v es g et th ei r p art s di st ribu t ed i n wa ys th a t m ost f ac ilit a t e

th ei r a ss imil a tive fu ncti on s u nd er the ci r cu m st a nces i n whi c h
,
th ey a re pl aced ; an d th ei r s ev era l typ es of s ym m et ry a re thu s

expli ca bl e. B u t i n fl ow ers the p et a l s a nd fr u c tifyi n g orga n s
,
o f whi c h do not c on t a i n c hl or ophyll th e t en d ency to g row

,
m os t wh er e th e s u pply of light i s grea t es t i s l es s d ec id ed if

, ,
1 68 MOR P H OL OGI CA L D E V E L OP ME NT .
n ot a s en b t ; an d a s h a p e oth erw i s e d et ermi ne d i s h ence l ess

li a bl e to a lt er i n c onseq u ence of a l tere d r el ati ons to su n a n d
ai r. Gr a vity too m u st be c om p a r a tiv ely i neffec tiv e i n c a u s
, ,
i ng m odifi c a ti ons : th e s m a ll er s i zes of the p a rt s a s well a s ,
t h ei r m od es of a tt a ch m ent givi ng th em greater r el a tiv e

,
g i di ty N
. o t i n d ee d t
,
h a t th e s e i n c id
,
ent f o r c e s of th e i no r
g ani e wor ld a r e h er e qu i te i nop era tiv e Fig . .
2 4 9 rep res en ti ng a sp e ci es of C a m pa n u l a
, ,
sh ows th a t the d ev el opm ent s of i ndivid u a l

fl owers a re s om ewh a t m odifie d by the rel a
ti ons of th ei r p a rts to genera l condi ti ons B u t .
th e fa ct to be ob s erv e d i s th a t th e ext rem e ,
t ra ns form a ti on s w hich fl ow ers u nd ergo a re

n ot lik e ly to be thu s c au sed : s om e fu rth er
c au se m u s t be s ou ght A n d if w e be a r i n
.
mi n d the functi on s of fl owers w e sh a ll fi nd i n ,
th ei r ad a pt a ti ons to th es e fu n cti ons u n d er conditi ons th a t a re

,
ext rem ely v a ri ed a n a d eq u a t e ca u se for t h e diff erent typ es

,
of sy m m et ry a s well a s for th e e x cepti ons to th e m

,
Fl owers .
a re p a rt s i n w hi c h f ert ili za ti on i s eff ect ed ; a n d the a ctiv e
a g ents of thi s f ert iliz a ti on a r e i ns ec t s b ees m oths b u tt er

-
, ,
fli es &c,
Mr D a rwi n h a s sh own i n m any c a s es th at the
. .
,
f orm s a nd p os iti on s of the es s enti a l org a ns o f fru cti fi ca ti on ,
a r e s u ch a s t o fac ili ta t e th e a cti on s of i n s ect s i n t ra n s
ferri ng p oll en f rom th e a nth ers of one fl ow er to the pi stil of

a n oth er— a n a rra n g em en t p rod u ced by na tu ra l s el ecti on .
A nd h ere we sh a ll fi nd rea s on for c oncl u di ng th a t the for ms ,
a n d p os iti on s of th o s e s u b s idi a r y p a rt s which giv e th ei r
sh a p e s to fl owers s imil ar ly a r i se by the s u rviv al of i ndi

,
vi d u a l s w hi c h h a v e the s u b s idi a ry p arts s o a dj u st ed a s to a i d
thi s f ertili zi n g p rocess— th e d evi ati ons f rom ra di a l symm et ry

b ei ng a m ong su ch a dj u stm en ts The r easoni ng i s a s fol .
l ows . S o l ong a s th e a xis of a fl ower is v er tic a l an d

t he conditi on s a re s imil a r a l l r ou nd a be e or b u tt erfly a light
,
i ng on it will be as lik ely to c om e f rom one sid e as f rom

,
a noth er ; and h enc e hi nd ra nc e r a th er th an f a cilit a ti on w ou ld

,
T HE S H A P E S OF F L O WE RS .
1 69
resu lt if the v
s e era l s id es of the fl ower did not a ffor d it equ a lly
free access . I n lik e m a nner fl owers w hi ch a r e di s t r ib u t ed
,
o v er a pl a n t i n s u c h wa y s th a t t h ei r di s c s o p en ou t on
pl a nes of a l l di recti on s a nd i nc li na ti ons will h av e no t end ,
e ncy to l os e th ei r ra di a l symm e t ry ; s i n ce on th e a v era g e , ,
n o p ar t of the p er iph ery i s di ffer en tly rel a t e d t o i n s ec t
a ge nc y f rom a ny oth er p a rt B u t fl owers s o fi x e d a s to

.
Op en ou t s id ewa y s i n t ol er ably const ant a ttitu d es h a v e -

,
th ei r p eta l s di fferently r el a t ed to i nsect a gency A bee or -

.
b u tt erfly c om i ng to a l a t era lly g row i ng fl ower d oes not s ettl e

-
,
on it i n one way a s r ea dily a s i n a n oth er ; bu t a l m os t of
n ecess i ty s et t l e s w ith th e a xi s o f i ts b ody i n c li n e d u p w a rd s
t ow a rds the st ern of the pl ant H en ce the s id e p et a l s of a

.
-
fl ower s o fix ed h a bit u a lly s t an d to the a lighti ng i ns ec t i n

,
r e l a t i on s di ffe r e nt f r o m th ose i n whi c h the u pp er a n d l ower
p et a l s st a nd ; a n d the u pp er an d l ower p e t a l s di ff er from one

a noth er i n t h ei r r el a ti on s to it I f th en th ere so a r i ses a n
.
, ,
h a bit u a l a t titu d e of the i nsect t owar d s th e p et a l s th ere i s ,
lik ely to be s om e a rra ngem en t of the p et a l s th a t will be

m ost c onv en i ent to the i n s ect— will m os t f a cilit a t e its ent r a nc e
i nt o th e fl ow e r T h u s we s ee i n m any c a s es th a t a l ong
.
,
u n d erm ost p e t a l or lip by ena bli ng the i n s ect to s ettl e i n

,
s u ch w a y a s t o b r i n g i ts h ea d Opp os it e to th e Op eni n g of th e
t u b e a id s i ts fertili zi ng a g ency B ut wh a tev er he th e sp eci a l

,
.
m odifi ca ti ons of the coroll a w hi c h fac ili ta te the ac ti ons of

th e p a r ticu l ar i n se ct s c oncer ned a l l of th em w ill c on d u ce to
,
bil atera l s ym m et ry ; s i nce th ey w ill be a lik e for th e two s id es

but u nlik e for th e t op a n d b ot to m A nd n ow we
.
a re p r ep a r e d for u n d er st an di ng th e ex cepti ons Fl owers .
g row i n g s id ew a ys ca n b ecom e thu s a d apt ed by su r viv a l of

the fi ttest only if th ey a r e of su ch s i zes a nd s t ru ct u res th a t
,
i n sect a gency c a n a ff ect th em i n th e w a y d escrib ed

- Bu t .
i n the pl a nt s na m e d a b ov e thi s c onditi on i s not f u lfill ed

,
A .
H ollyh ock fl ower i s so Op en a s w ell a s s o l a r g e th a t i ts p et a ls

-
, ,
a r e not i n a n a pp re ci a bl e d e gr ee di ffer ently r el a t e d to th e

y
i ns ect s whi ch vi s it it On th e oth er h a nd the fl ow er of th e
.
,
A gr im ony i s ll th a t u nl es s vi s i ted by i ns ects of a

so sm a ,
c orr es p on di ng s i z e w hich settl e a s b ees a n d b u tte r fli es s et tl e ,
i t s p a rt s w ill n ot be a ffect ed i n th e a ll ege d m a nner T ha t .
a l l a n om a li es o f thi s ki n d c a n a t on ce be s a ti s f a c t or ily ex
pl a i ned is s c a rcely t o be exp ect ed : the ci rcu m s t a nces of ea ch

,
c a s e h a v e t o be s t u di e d B u t it seem s n ot imp rob a bl e th a t

.
th ey are du e to c au ses of the ki nd i di cat ed

n *
.
23 5 . We h a v e a l rea dy gl a nc e dt fl ow er s for a t c u s ers of

l
th e p u r p ose of c on sid e ri ng th ei r s h a p es a s c l u ster s We m u s t .
now retu rn to th em to ob s erv e the m odific a ti ons u n d er gone
by th e i r com p onent fl owers A m ong th ese occu r illu st rati ons .
o f g rea t s ig n ifi ca n ce .
A n ex am pl e of t r a nsiti on f rom th e ra di a l to the bil a t er a l

f or m i n c l u stered fl ow er s of the s a m e s p eci es i s f urn i sh ed by ,
the cu ltiv a t e d Gera ni u m s c a ll e d by fl or i sts P el a rgoni u m s,

.
S om e of th es e b ea r i ng som ewh a t sm a ll t ermi n a l cl u st ers of

,
fl ow er s whi ch a r e cl osely p a ck ed t ogeth er w i th th ei r f a c e s

,
a l m os t u p w a r d s h a v e ra di a lly s ymm et r ic a l fl ow er s
,
B ut -
.
a m ong oth er v a ri e ti es h a vi n g t er mi n a l c l u st e r s of w hi c h th e
m e mb ers a re m u tu a lly th ru st on one s id e by c row di n g th e ,
fl owers d ep a rt very cons id era bly f rom the r a di a l s h a p e

It i s j
ob e c te d to th e abo e v i nte rpre ta ti on tha t “
m a ny fl ow e rs o f s i z es
i nterm edi a te b et w een th e Hol l yh ock a nd t h e A gr i m ony a re ra di a l l y sy m .
m e tri ca l a nd yet g row s i de ways . I m a y m enti on v a rious L i l i a ceaz, e .

g.Ch ’o .
rop hyt a m ,
E l wom is , M us ca ri , A nt/
aeri cm n . S a gi tta fi a
,
a l s o, has m a ny o f i ts
fl owe rs i n t hi s pos i ti on Furt her, i f t he hi gh e r i nse ct s a l ight on o we rs i n
. fl
a d efini te wa y a s th ey do, th e p a rt s of t he ow e r m u st be a r di ffe re nt re l a
. fl
t i ons to the v i s i tin
g i ns e ct , h owe v er l a rg e, s o th a t owers un v i s i t ed ough t fl
al l t o be zyg om orp hi c
”
.My re pl y i s th a t i n the s e nse wh i ch he re conce rns
us , t h e di ffe rent pe t a l s of t he Hol l yhoc k fl ow er do not be a r di fferent rel a
t io ns to th e v i s i ti ng
i ns e c t ; s i nce , p ra c ti ca l l y, th e up p er a nd l a te ra l pe ta l s
b e a r no phys i ca l rel a ti ons a t al l : i n s o fa r a s t he v i s i ti ng bee i s concern ed
t h ey a re non-e xi s tent . The a rgum en t i m pl ie s th a t c h a nge i n t h e form of a
fl o we r from th e ra d i a l to t h e bi l a te ra l i s l i k e l y t o t a ke p l a ce onl y whe n the
c onta ct-re l a ti ons o f t h e p e ta l s to t h e v is i ti ng i ns e ct, a re s uch a s to m ak e s om e
form s fa ci l i ta te i ts a c t i on m ore t h a n ot h e rs ; a nd t h e l a rge p e ta l s of th e

Hol l yhock ca nnot fa ci l i ta te i ts act i on at al l . In res p ec t of th e L i l i a ccce
i nst a nced, i t i s ne ed ful to i n q ui re whethe r th e s t ruc ture s a re s uch th at thi s
a l l eged c a us e o f b i l a tera l sym m e try ca n co m e i nto pl a y .
THE S HA P E S OF FL OWE RS .
t owa rd s th e bil a t era l sh a p e A lik e resu lt occu rs u nder lik e

.
c ondi ti ons i n Rhod od en d rons a nd A z a l ea s T he Ver bena too .

, ,
fu rni s h es a n ill u stra ti on of ra di a l fl ow ers ren d ere d s ligh tly

t wo s id e d by the s light tw o s id e d n ess of th ei r rel a t i ons to
- -
o th er fl ow ers i n th e cl u s t e r A nd a m ong th e C r u ci ferce a

.
ki nd red case oc cu r s i n the cu ltiv a t ed C andyt u ft .
E vid e nce of a s om ewh a t difi erent ki n d i s offe re d u s by

cl u s tere d fl ower s i n whi ch the p eriph era l m em b er s Of the
c l u ste r s di ffe r from the cen tr a l m e mbe rs ; a nd thi s e vid en ce
i s es p ec i a lly s ignifi ca nt wh ere w e fi n d a lli e d s p ec i es th a t do
n ot exhibit the d evi a ti on at th e sa m e ti m e t h a t th ey do not
,
fu lfil the c ondi ti ons u nd er w hi c h it m ay be ex pe c te d T hu s .

,
i n S c a bi osa s u ccis a Fig 2 5 0 whi c h b e a r s i ts n u m er ou s s m a ll

,
.
,
fl owers i n a h em i s ph eri ca l k nob the com p onen t fl owers , ,
s i m il a r ly c i rc u m st a n c e d a r e a l l eq u a l a n d a l l ra di a l ; bu t i n
,
S ca bi osa a rv ens i s Fig 2 5 1 i n w hi c h the n u m erou s sm a ll

,
.
,
fl ower s form a fl a t ten ed di s k

onl
y t h e c onfi ne d c ent ra l one s
a re ra di a l : rou n d the e dg e th e
fl owers a re m u ch l arger and

cons pi cu ou s ly bil at era l .
Bu t th e m os t re m ark a bl e
a n d m os t c on cl u s iv e p r oofs of th es e r el a ti ons b et ween f or ms
a n d p os iti ons a r e th os e giv en by th e c l u s t e r ed fl ow er s c a ll e d

,
Um bel l i fer ce . I n som e c a ses a s wh ere th e com p on ent fl owers

,
h a v e a l l pl enty of room or wh ere the s u r fa c e of the u mb el i s

,
m ore or l ess gl ob u l a r the m odifi cati ons a re not cons pi cu ou s ;

,
but wh ere a s i nVi bu r n u m C hcer 0p hyl l u m A n thri s cus T ori l i s

, , , , ,
C a u ca l i s D a u cus T or dyl i u m &c we h a v e fl owers c l u s t ere d

, , ,
.
,
i n su ch wa y s a s t o be differ en tly con diti one d w e fi nd a nu m ,
be r of m odifi ca ti ons t h a t a re m ark e d a nd v a ri e d i n p rop or

ti on a s th e di fferen ces of c onditi ons ar e m ark ed a n d v a ri ed .
I n C ha r op hy l l u m wh ere th e fl ow ers o f ea c h u m b ell u l e a re

,
c l os e ly pl ace d s o a s to for m a fl a t su r fa ce bu t w h ere th e ,
u m b e ll u l e s a r e w id e a p a rt a nd f or m a di s p ers e d u m b e l the ,
u m b ell u l e s do n ot di ffer from one a noth er ; th ou gh a m on g th e

1 72 MORP HOL OGICA L D E V E L OP ME NT .
fl owers of each u m b e ll u l e th ere a re d ec id ed di fferences : th e

cen tr a l fl owe r s b e i n g s m a ll a n d ra di a l w hil e th e p eriph era l
,
one s a re l a r g e a n d bil a ter a l B u t i n oth er gen er a w h ere n ot

.
,
on ly t h e fl ow e r s of e a c h u m b e ll u l e bu t a l s o th e u m b e ll u l e s
them s elve s a re cl osely cl u st ere d i nto a fl at su r face th e u m b el

, ,
l u l es th em selv es b ecom e c on t ra s t ed ; a nd m any rem a rk a bl e

s e con d a r y m odifi ca ti on s a r i se I n an u m b el of H era c l eu m
.
,
for i nst a nce th ere a r e to be n ot ed the fa ct s — fi rst th a t th e

, ,
e x tern a l u m b ell u l e s a re l a rger th a n the i nte rna l ones ;

s ec on d ,
th a t i n ea ch u m b ell u l e the cent ra l fl owers a re l ess
d e ve l op e d th an the p eriph era l ones ; thi r d th at thi s gr eat er ,
d ev elopm en t of the p eriph era l fl owers i s m ost m ark ed i n th e

o u te r u mb e ll u l e s fou rth th at it i s m ost m a rk ed on th e ou ter
,
s id es of the ou te r u m b e ll u l es ; fi fth th a t w hil e the i nter i or

,
fl owers of ea ch u m bellu l e a re ra di a l the ext eri or ones a re,
bil a t era l ; s ixth th a t thi s bil a t er a l nes s i s m ost m a rk ed i n

,
the p er iph e r a l fl ower s of th e p er iph era l u m b ell u l es ; s ev enth ,
th a t the fl owers on the ou t er s id es of these p eriph era l

u mb e ll u l es a r e th os e i n w hi c h th e bil a t era l nes s r ea ch e s a
m aximu m ; a nd eighth th at wh ere th e ou t er u m b e ll u l es

,
t ou ch one anoth er the fl ow ,
ers b ei n g un sy m m et ri ca lly
,
pl a c ed a re u n sy m m etric a lly
,
bil a ter a l *
T he lik e m odi fi
.
c ati ons a r e d i spl aye d though ,
not i n so c l e a rly t ra cea bl e a -
wa y i n a n u mb el of T or dy
,
l i u m Fig 2 5 2 C ons id eri n g

,
. .
how obvi ou sly th e s e v a r i o us
f orms are r el a t ed to the v a r i

ou s c on diti ons w e sh ou ld be ,
s c ar cely a bl e even i n the ,
ha d i nte nded h ere to i ns e rt a figu re exh i b i ting the se di fference s ; bu t

a s t he Cow -p ar s ni p doe s not fl o we r t i l lJ ul y, an d a s I c a n fi nd no d ra wi ng
o f t h e u m be l wh i ch ade q ua tel y rep rese nts i ts det a il s , I a m ob l ig ed to ta k e
a noth er i ns ta nce .
THE S H A P E S OF F L OWE RS .
1 73
a b s ence of a l l other fa ct s to res i st the c oncl u s i on th at the ,
di fferen ces i n the c on diti ons are the cau s es of the differenc es
i n the f or m s .
Com p os i te fl ower s fu rn i s h evid enc e s o nea rly a lli ed to th a t

w hi c h c l u s t er e d fl ower s f u r ni s h th a t w e m ay fi tl y gl an ce ,
a t th e m u nd er the s a m e h ea d S u ch a co mm on typ e of .
thi s or der as the S u n fl ower ex em pli -

,
fi es th e ext rem ely m a rk e d diff eren c e

w hich a r i ses i n m a ny of th es e pl a nts
b etween the cl ose ly p a ck ed i nt erna l -
fl oret s each s imil ar ly c i rcu m st a nced on

,
a l l s id es a n d th e e xt e r n a l fl or et s
,
not ,
s i m il a r ly ci rcu m st anc e d on a l l s id es .
I n Fig 2 5 3 r e p r es enti ng the i nner a n d

.
,
ou t er fl or e ts of a D a i s y th e con t r a s t is ,
m ark ed b etween the s m all r a di a l c oroll a of the one and the

l a rg er bil a t er a l coroll a of the oth er I n m a ny c ases how .
,
ev er ,
this c ont ras t i s l ess m a rk ed : th e i nne r fl or ets a l s o
h a VI ng th e i r ou t w a r d g row i ng p r ol onga t i on s— a difference
-
p oss ibly rel at ed to s om e differen ce i n th e h a bit s of th e i nsec t s

th a t f ertili ze th em N ev er th e l es s th ese c omp os it e fl owers
.
,
whi ch h a v e i nn er fl oret s w ith st r a p s h a p e d c oro ll a s ou t -
w a r dl y di re ct ed equ a lly c on form to th e g en era l p r i nc ipl e ;

,
b oth i n the r a di a l a rra ng em ent of the a ssem bl a ge of fl orets ,
a n d i n th e bil a t era l s h a p e of eac h fl or et ; w hich h as i ts
p a rts a lik e on the two s id es of a li ne p as si ng f rom th e c ent re

of th e a s s em bl a g e to th e c i r cu m fe r en c e C e r ta i n .
oth e r m e m b e r s of thi s or d er fu lfil th e l a w som ew h a t di f fer

e ntly . I n C en ta u r ea for i nsta nce th e i nn er fl orets a r e s m a l l
, ,
a n d v er ti c a l i n di r ect i on whil e th e ou t e r fl or e t s a r e l a r g e a n d
,
l a t er a l i n di recti on A n d h e re m a y be re m a rke d i n p a ss i ng
.
, ,
a c l e a r i n di ca ti on o f th e e f fec t wh i ch g r ea t fl e xibili ty of th e
p et a l s h a s i n p r ev en t i n g a fl ow er f rom l os i n g i ts origi n a l
r a di a t e for m
’
; for wh il e i n C cya n us th e l a r g e ou t w a r d grow
-
,
.
i ng fl or et s h a vi n g s h ort s ti ff divi si ons of th e c or oll a a re

, , ,
d ecid edly bil a ter a l i n C s ca bi os a w h ere the divisi ons of the

'
, .
,
coroll a a re l ong and fl exibl e the ra di al f orm i s s c ar ce ly at ,
a l l m odifi e d On b ea ri ng in mi nd the p rob a bl e rel a ti ons of

.
the forms to i ns ect a g ency the m eaning of this diff e renc e

-
,
will not be difficu lt to u nd erst and *

.
§ 23 6 . xt r emIn
ely v a rei e d w a y s th e re a r
-
e th u s re ill u s -
tr a ted a mong fl owers th e genera l l a w s of form which l ea v es

,
a nd b ra nch es a n d en ti re pl a nts di s cl os e t o u s Composed as .
e a c h cl u st er of fl ow e rs is of i n divid u a ls th a t a re or igi na lly
s i m il a r ; a nd c omp os e d a s e ach fl owe r i s of h om ol og ou s f oli ar
o rg a n s ; we see b oth th a t th e lik e fl ow ers b ecom e u nlik e a nd
the lik e p art s of ea c h fl ower b ec om e u n lik e wh ere the p os i ,
ti ons inv olv e u nlik e i ncid ence of for ces T he s ymm et ry .
r em a ins ra di a l wh ere t he c onditi on s a re eq u a l al l roun d ;
s h ow s d evi a ti on t ow a r d s two s id e d ne ss wh er e th ere i s s light

-
t wo sid ed n es s of c onditi on s ; be com es d ecid e dly bil a tera l

-
wh ere the conditi ons a re d ec id edly bil a tera l ; and p ass es i nto
a n u ns ymm e t ric a l f or m w h ere the rel ati ons to th e en vi ron
m ent ar e u n sym m etr ic a l .
I t h a s be en pointed ou t to m e th a t “ th e e xt re m e d ev el opm e nt o f the

c oro l l a s o o ft en fou nd i n t h e o u te r fl
o we r s or on t h e o ut e r s i de o f t he o ut e r
fl o w ers in cl os e l y p a ck e d i nfl o re s c en c e s , a s s o c i a te d as it o ft en i s wi th d i sa p
p ea ra nc e o f s ta m ens or ca rp e l s o r both, i s u s ua l l y p ut d o wn to s p e ci a l i za t i on
of th ese out e r fl o we rs for a ttra ct i v e p u rp os e s . Since th e wh ol e i n ore s ce nc e fl
i s i nc rea sed i n by s u ch a m odi fi ca t i on, i t is s u ppos e d t h a t
co ns p i c uou sne s s
v
n a t u r a l s e l e ct i on fa oure d th o s e pl a nt s w h i ch s a cri fice d a po rt i on o f the i r
s ee d b ea ri ng ca p aci ty fo r th e s u p po se d g rea te r a d a nt a ge o f s e curi ng m o re

-
v
i ns ec t v i s its .
”
But gr ant i ng t hi s i nte rp re ta ti on, i t m a y s ti l l be he l d th a t
i nc rea se of a tt rac ti v ene s s d ue t o i nc rea s e o f a rea m u s t be ach i e ed v by fl ore ts
a t th e pe ri ph e ry, and v
th a t th e i r a bi l i ty to a ch i e e i t d e pe nds on v
t he i r h a i ng
a n ou te r, u noc c up i ed, s p a ce w hich th e i nner fl ore t s ha v e not ; so th a t, th ough
in a m o re i ndi rec t w a y, th ei r di fferent dev el op m ent is determ i ned by di ffe re nt

e x posure to condi ti ons .
C H A P TE R X I .
TH E S H A P E S or VE GE T AL CE L L S .
§23 7 . WE c om e gg re g a t es of th e l owes t or d er
now to a .
A l rea dy s om ethi ng h a s b een s a id 2 1 7 ) c onc er n i n g th e

f or m s of th ose m orphol ogi c a l u n i ts w hi c h e xi s t a s in depen
d ent pl ants Bu t it is h e re requ i s it e b ri e fly to not e th e
.
m odifi ca ti ons u nd ergone by th em wh ere th ey b ecome comp o

nents of l a rge r pl a nt s .
Of the nu m erou s c ell f or m s w hi ch a re fou n d i n th e ti s su es

-
of the high e r pl a nt s it will s u ffic e to giv e i n Fig 2 5 4

,
re ,
.
,
p res enti ng a secti on of

a l ea f ,a s i n gl e e x a mpl e .
I n u se it will be seen
th a t the cells for m i ng
the u pp er a n d l ower su r
f a ces a a n d b h a v e di f
, ,
ferences of sh a p e r el a te d
to di fl e rences i n th e i nc i
d ence of for c es : t h ey a re
m ore or l ess fl a tt en e d i n
r el a ti on to th e e n vi r onm ent T he u n d ernea th c ell s a t 0
.
,
form a c l a ss which s imil arly exp ose d to light at th ei r

,
ou ter e n d s ,
an d a s we m a y a s s u m e
,
l a r gely d e vel op e d i n
,
a dj u s t m ent t o th ei r a ctiv e a s s i m il a tiv e fu n c ti on s a re by , ,
mu tu a l p r e ss u r e m a d e to g row m or e i n th e di recti on of th ei r
,
l en gt h s th a n i n th e di recti on of th ei r b rea d th s T h en on .
the oth er s id e we s ee t h a t th e cell s d nex t a bov e th e ou ter ,
l ayer w hil e a pp roxi m ately si m il a r b ecom e m ore a nd m ore

, ,
dissimil ar a s th ey div er ge f rom th e su r fa ce a n d a re quit e ,
1 75
1 76 MOR P H OL OGI CA L D E V E L OP ME NT .
i rre gu l ar i n the i nt er i or 3 wh ere th er e i s no defin i ten ess i n

,
th e conditi on s to w hi c h th ey a re exp osed T hu s th e div er

.
g ences of th ese c ell s f rom p r i m ordi a l s ph er i city a re s u c h a s

corres p on d w ith u n lik ene ss es i n th e i r c i r cu m s t a n ces A nd .
th r ou gh ou t th e m ore c om pl e x m odific a ti ons w hi ch the c ell s

of oth er ti s s u e s e xhibit th e lik e c orres p on d en ce s h old
, .
A m ong pl a nts of a l ow er or d er of a ggr ega ti on we h a v e ,
a l r ea dy s ee n h ow c e ll s be com e m et a m or ph os e d as th ey b eco m e
i nteg ra t ed i nt o m asses h avi n g d efi n i te or g an i zat i on s T he .
high er A l g ae ex emplifi e d i n Fig s 3 2 3 4 3 5 s h ow th i s ve ry

,
.
, , ,
c l ea r ly . H ere th e d e p a rtu re from

the s i m pl e cell form to the fo m
- r
o f a n el on g a t e d p r i s m i s m a ni ,
festl y su b ordi na ted to th e c on

tr asts i n the r el a ti ons o f th e
p a rts A nd it i s i nteres ti ng to oh
.
s er v e h ow i n on e of th e b r a nch e s
,
of Fig 3 2 w e p ass from th e s m a ll

.
, ,
a l m os t s ph e r ic a l c ell s w hich te r
-
mi na te th e b r a n chl ets to t he ,
l arg e m u c h m odi fi ed c e ll s whi ch

,
-
j oi n the m a i n s tem th rou gh g ra ,
d a ti on s obvi ou sly rel a t ed i n th ei r ch an ged f orm s to the

a lt er e d a c ti on s th e i r p os iti on s exp os e th e m to .
More si m ply bu t qu it e a s con cl u s iv ely do th e i n fe ri or

, ,
A l gae of whi c h Fig s 1 9 — 2 3 a re ex a m pl es sh ow u s how

, .
,
cells p ass f rom th ei r origi n a l s ph eri ca l s y m m et ry i nt o r a di a l

s ymm et ry a s th ey p as s from a s ta te i n which th ey a r e s imi
,
THE SH A P ES OF VE GE TA L CE L L S .
1 77
l arl y c onditi one d on al l s id es to a st a t e i n whi ch two of th ei r

-
,
opp os i te s id es or e n d s a r e c on diti on e d i n w a y s th a t are lik e
one a noth er bu t u n lik e th e w a y s i n whi c h a l l oth er sid es a re

,
c onditi on e d .
S till m or e i n s t ru ctiv e a r e th e m or ph ol ogi ca l di ff erenti a

ti on s of th os e p r otophyt es i n whi ch the fi rst s tep s t owa r d s a
higher d eg ree of i nt eg r a ti on a r e s h own I n Fig 1 0 rep r esent . .
,
i ng on e of the t r a ns iti on a l f orm s of D es m i di a cew it i s to be

l
,
note d t h a t b e s id es th e di ffe r enc e b et w een th e t ra nsv er s e a n d
l ongitu di na l di m ens i ons w hich the c omp onent u n i ts di s pl a y

,
i n c om m on th e two end un its di ffer f rom th e re st : th ey h a v e

,
-
a pp end a g es whi ch th e r est h a v e not Onc e m ore wh ere the .

,
i nt eg ra ti on is c arr i e d on i n su ch wa y s a s to p rodu ce not st ri ngs

bu t cl u s ters th ere a r i s e c ont r a s t s a n d c orresp on d en ces j u s t
,
su c h a s m ight be l ook e d for A l l th e fou r m em b er s of th e

.
g rou p sh own i n Fig 1 2 a re s imil arly conditi on e d ; a nd ea ch

.
,
of th em h a s a bil a te r a l sh a p e an sw er i n g t o i ts bil a t era l r el a
ti on s . I n Fig 1 4 we h av e a nu m b er of s imil a r ly bil a t er a l

.
-
i ndivid u a l s on th e ci rcu m feren ce i ncl u di ng a c ent ra l i n ,
divid u a l di fi er i ng f rom th e rest by h a vi n g th e bil at er a l

c h a r a ct e r n ear ly oblit er a t ed A nd th en i n Fig 1 5 we h a v e
.
,
.
,
two centra l comp onents of th e grou p d evi a ti n g m ore d e ci ,
dedl y from th os e th a t su rrou nd th e m * .
One of m y cri ti cs w ri tes Thi s ch a pter m i gh t of c ou rs e be enorm ous l y
ex te nded . not onl y a s i n the p rec eding o ne s b y c i ta t i on of qui te si m il a r cas es ,

”
but by th e i ntrod uc ti on of fre sh grou ps of ca ses .
CH A P T E R X I I .
C H A N GE S OF S H A P E O T H E RW I S E CA U SE D .
23 8 . B E S I D E S th e
m ore sp eci a l c a u s es of m odifi ca ti on i n
th e s h a p es of pl a nt s a nd of th ei r p ar ts c ert a i n m ore genera l
,
c au ses mu st be b r i efly notic ed T h ese m ay be d escr ib ed as

.
c onsequ enc es of v a ri ati ons i n th e t ot a l qu antiti es of th e

m a tters an d forces fu rni sh ed to pl ant s by th ei r envi ronm ent s .
S om e of the c h a nges of form so p r od u ce d are di s pl a y ed by

pl a nt s as wh ol es a nd oth er s only by th ei r p art s
, We w ill .
gl anc e a t th em i n thi s or d er .
§23 9 . It is f a mili a r f a c t th a t l u x u ri a n t s hoot s h a v e

a
r el a tiv ely l on g
-
i nt ernod es ; a nd c onv ers ely th a t a s h oot
, ,
d wa r fe d from l a c k of s a p h as its nod es c l osely cl u stered : a

,
c on com it ant resu lt b ei ng th a t the l a t era l a x es wh er e th ese ,
a r e d e v el o p e d b ecom e i n th e one c a s e far a p a rt a nd i n the

,
oth er c a s e n ea r t og eth er Fig 2 5 5 rep resent s a b ra nch to

. .
th e p art s of whi ch the l onger a n d

sh ort er i nt ernod es so resu l ti ng giv e
di fferenti a l ch ara ct ers A wh ol e t re e .
b ei ng i n m a ny c a se s s imu lt aneou sly

th u s a ffect ed by st at es of the ea rth or
th e a i r a l l p a rt s of it m ay h a v e s u ch
,
v a ri ati ons imp r ess ed on th em ; a nd ,
i nd eed s u ch v ari a ti ons f oll owi ng m ore

, ,
or l es s re gu l a rly th e ch a ng es of the
s ea s ons giv e to m any t rees m anifes t

,
CH A N GE S OF S H A P E OT H E RWI S E CA U S E D .
1 79
t r a its Fig 2 5 6 a s h oot of P hyl l oca ctus

of s ru c u re t t . In .
,
cr ena tus we a n i n t er es ti n g e x a m pl e of a v a r i a ti on
,
ha ve
ess ent i a lly o f th e s a m e n a t u r e li ttl e a s i t a pp ea r s to be so ,
.
F or ea ch of the l a tera l i n d en ta ti on s i s h er e the s eat of a n

a xill ar y bu d ; a n d th ese we s ee are s ep a r a t e d by i nt ern od es
whi ch be c om i ng b roa d er a s th ey b e com e l ong er a nd n arrowe r

, ,
a s th ey b ecom e s h ort er p rod u ce ch a nges of for m th a t c or r e ,
spon d w ith ch a ng es i n th e l u x u ri a n c e of g r owth .
T o c om pl et e the s t a te m ent it mu s t be a dd e d th a t th ese

v ari a ti on s of nu t r iti on oft en d et erm i ne the d ev el op m en t or
non d ev e l opm ent of l a t er a l a x es ; a nd by s o d oi n g c au s e still
-
m ore m a rk ed st ru ctu ra l differenc es T he Fox gl ov e m ay be .

-
*
n a m e d a s a pl an t whi c h ill u s t ra t e s this t ru th .
F rom th e m orph ol ogic a l di fferenti a ti on s c au sed by

24 0 .
u n lik enes ses of nu t r iti on f elt by th e w h ol e pl a n t w e p a ss ,
now to th ose w hi ch a r e th u s c au s e d i n s om e of its p a rt s a n d
n ot i n oth ers A m ong su ch a re th e c ont ra s t s be t ween

.
fl oweri ng a x e s a nd th e a x es th a t b ear l ea v es only I t h as

,
.
a l r ea dy be en s h own i n
§ 7 8 th a t th e be li e f e xp r ess e d by ,
Wol ff i n a di rec t conn exi on between fru cti fi c ati on and i nnu
t ri ti on i s j u s tifi ed i n d u ctiv ely by m a ny f a ct s of m a ny ki nd s
, .
D ed u ctiv ely too i n 7 9 we s aw rea son to c oncl u d e t h a t su c h

, ,
a re l a ti on wou ld be e s t a bli s h e d by s u r viv a l of the fitt es t ; .
s eei n g th a t i t wou ld p rofit a s p ec i es for i t s m em bers to b e gi n '
s endi ng of f migr a ti n g g erm s from the ends of th ose a x es

whi ch i nnu t r i ti on p rev ent ed f rom fu rth er a g a m ogenetic m u l
t i pl i c a ti on Once m ore wh en c ons id e r i ng th e na t u re of th e
.
,
pha n oga m i c a xi s we f ou n d su pp ort for thi s b eli ef i n the fa c t

,
Natura l se l ec t i on m ay ha v e ope ra ted in e s tabl i sh i ng a const i t u t i ona l
te nde nc y to o th e r s u dde n a b ri dg m ent s . Mr Ta nsl ey

. a l l e g es th a t thi s i s a
p a rt-c au s e of v a rying di st ri bu ti on of l ea v e s He s ays — I h a v e m y se l f

th e .
“
m ade s o m e ob s erv a ti ons on t h e l en g th o f i nt ernod es i n th e B eec h a nd a m

,
s a ti s fie d th a t i t fo l l o w s q u i te oth e r l a w s connec te d wi th th e s ui t a b l e d i s
,
po si ti n o i th e l e a v e s on th e b ra nch Al th ough I h a v e not h ad the opp er

'
o .
tuni ty of fol l owi ng up thi s l i ne of w ork so a s in a ny w a y t o gene ra l i ze th e

r e s u l ts
,
I s us pe ct t ha t
‘
in di re ct e q ui l i bra t i on ’
is a wi de s p rea d ca us e of s uch
v a ri a ti on .
"
1 80 MOR P H OL OGI C A L D E V E L OP M E N T .
th a t the comp on en ts of a
fl ow er exhibi t a rev ers i on to th a t
typ e from w hi c h t he ph aen o ga m i c typ e h a s p r ob a bly a ri sen
a r e v er s i on w hi c h th e l aw s of e m b ry ol ogy wou ld l e a d u s to
l ook for wh ere i nnu t r i ti on h a d a r re sted d evel opment .
Hence th en we m ay p r op er ly c ou nt those devi a ti ons of

‘
, ,
s t ru c t u r e whi c h c on stit u t e i n fl or es c enc e a s a m on g th e m or ,
l
“
p h ol og i ca di f
f eren ti a t i on s p rod u c e d by l oca l i nnu tr i ti on I do .
not m ean th a t th e d et a il e d m odifi c a ti ons w hi ch th e e ss ent i a l
a n d s u b s er vi ent or ga n s of fru cti fi cati on di s pl ay are th u s ,
a ccou n t e d for : we h a v e s een r ea son to t hi n k th e m oth e rw i s e
c a u sed Bu t I m ea n th a t th e m or ph ol ogi ca l ch a ract er s whi c h

.
di sti ngu i s h g a m og en eti c a x e s i n g enera l f rom a g a m ogeneti c

a x es s u c h a s n on d e v el opm en t of th e i nt ern od es a n d d w a r f
,
-
i ng of th e foli ar or ga ns a re p r i m a r ily r esu lt s of f a il u re i n ,
*
the s u pply of s om e m at er i a l requ i r ed for f u rth er g r ow th .
§ 2 4 1 A noth er t ra it whi ch h a s to be n oti ce d u n d er thi s

.
h ea d i s the s pi ral or r a th er the h eli cal a rr a ngem ent of

, , ,
p art s T he su ccess iv e n od es of a ph aenog a m h abi tu ally b ea r

.
th e i r a pp en d a ge s i n w a y s i m plyi ng m or e or l es s t wi s t i n the
s u bst a n c e of t h e a xi s ; a n d i n c li m bi ng pl a n t s th e t wi s t i s s u c h
It i s but j u s t t o th e m em o y of W ol fl he e t o po i nt ou t th a t h e wa s
’
r r
,
im m e ns el y in ad v ance of Go e the i n h i s ra t i ona l e of th e s e m e t a m orp h os e s .
What ev er gre a t e r e l a b ora ti on Goe th e ga v e to th e t heory cons i der e d as an
i nduct i on, s ee m s to m e m ore . th a n c ounte r-b al a nce d by th e i rra ti ona l i ty of hi s
d ed u cti v e i n te rp ret a ti on ; whi ch uni t e s m edi ae a l v phys i ol ogy wi th P l a toni c

p h i l os ophy . A dom i na nt ide a w i th h i m i s th a t l e a v e s ex i s t for th e p u rp os e o f
c a rr y ing o ff cru de j u i c es -
th a t as l ong a s t h e re a re c rude j u i ce s t o be c a r ri e d
o v
i , th e p l a n t m u s t be p ro i de d w i th org a ns co m pe te nt to
’‘
e ffec t the t a s k
h
t at w i
hi l e the l es s p u re u ids a re got ri d of, pu rer one s
‘
fl a re i ntrod u ce d
a nd tha t
“
if nour i s h m ent i s w i thh el d , t h a t op e r a t i on o f na tu re .
( fl o we ri ng) i s
fa cili t a t e d a nd h a s t e ne d ; th e or
g an s of t he node s ( l ea v es ) be c om e m ore
r e fi ned i n t ex ture , th e a ct i on of th e p u ri fie d j ui c e s b e c om es s t ro ng e r , a nd th e
t ra ns form a t i on of p a r ts h a v i ng now b ecom e po s s ibl e t a kes pl a ce wi tho ut

.
del a y Thi s be ing th e prox i m a te e x

.
pl a na ti on, the ul t i m a t e e pl a na t i on i s , x
tha t Na ture wi s hes t o fo rm fl o w ers— t h a t w h en a pl a nt fl o we rs it a t ta i ns th e
”
Na tu re l ength a t ta i ns h er
“
t o i t by n a tu re a nd tha t at
e nd p re s cribed ; so o
bj v i ti a ti ng
h i s i nd uct i on by a tel e ol ogy th a t i s a s u nwa r
”
o ect . I ns t e a d of
r a nt ed i n i t s as s i gned o b e c t a s i n i t s a s s i gn e d m e a ns , Wol f
j f a s c ri b e s t he
'
u s e w hi c h , w he th e r s u fl i c i c nt or not , i s s t ri ct l y s c i ent i fic i n
‘
p h e n om e n a t o a ca
C H A NGE S O F S H A P E OT H E R WISE C A US E D .
1 81
as to p rodu ce c ork sc re w s h a p e T his s t ruct u re i s a scri b

a .
a bl e to di fl e r en ces of i n te rs ti ti a l nu t r i ti on T a k e a s hoot .
w hich i s g row i ng v erti ca lly I t i s c l ear th a t if the m ol ecu l e s

.
a re a dd e d w i t h p e r fec t eq u a li ty on a l l s id e s th e r e will be n o ,
t end ency tow a rd s a ny ki n d of l a ter a l d evi at i on ; a n d th e

su cces s iv ely p r o d u c e d p a rt s will be p e rp en di cu l a r ly ov er o ne
-
a noth er Bu t a ny i neq u a li ty i n the r a t e of g rowth on the

.
di fferent s id es of the s h oot w ill d es troy t hi s s t ra ight ness i n

.
,
t he li nes o f gr ow th I f the g rea t es t a n d l ea s t r a t es o f m ol e

.
c u l a r i ncr ea s e h a pp en to be on oppos i te sid es , th e s h oot m u s t

‘
a s su m e a c u r v e o f si ngl e c u rv a tu r e ; bu t i n e v e r y oth e r c a s e
of u nequ a l m ol ecu l a r i n c r eas e a cu r v e of d ou bl e c u rv a t u r e,
m u s t res u lt N ow it i s a coroll a ry f r om t he i n s t a bility of th e

.
h om ogeneou s th a t the r a t es of g rowth on a l l s id es of a s hoot

,
ca n n e v er be e x a ctly a lik e ; a nd it i s a l s o t o be i n f e rr e d f r om
the sa m e gene r a l l a w th a t th e g r ea test a n d l e a s t r at es of

,
g rowt h will not occu r on e x actly opp os i te s id es of th e sh oot ,
a t th e s a m e ti m e th a t e qu a l r a te s of g r ow th a r e p r es er v e d by
t he two oth er sid es H en ce th er e m u s t a l m os t i n evit a bly

.
,
a r i se m or e or l ess of tw i s t ; a n d th e a pp en d a g es o f th e i nt e r
n od es w ill s o be p re v ent e d f rom occ u rr i ng p er p en di cu l a rly
o ne ov er a n oth er .
A d evi a ti on of thi s ki n d nec e ss a r ily i n iti at e d by phy s i c a l

,
c a u s es i n c onform ity w i t h th e g ene ra l l a w s of e v ol u t i on is ,
lik ely t o be m a d e r egul a r a nd d ecid e d by n a t u ra l sel ec ti on .
For u nd e r or di n a ry c i rcu m sta nces a pl a nt p rofi ts by h a vi n g ,
i ts a xi s so tw i s te d a s to b r i ng the a ppe n d e d l ea v e s i nto p os i

ti ons whi ch p rev en t th em f rom s h a di ng on e a noth er A nd .
,
m a nifestly m odifi ca ti on s i n th e for m s si ze s a nd i n serti ons of

, , ,
th e l ea v es m a y u n d er th e sa m e a genc y l ea d to a d a p te d
, , ,
modifi ca ti ons of the tw i s t We m u st th erefor e a scr ib e thi s .
c om m on c h ara cte r i sti c o f ph aenog a m s p r i m a r ily t o l oca l di ffer ,
e n ce s of nu t ri ti on a n d s e con d a r ily to s u rviv a l o f th e fi ttes t

, .
i ts cha ra cte r . Va ri a t i on of n ut ri t i on is un q ues ti ona bl y a tru e ca u s e

”
of
va ri a t i on
i n pl a nt s t ru ct u re-
. We ha ve he re no i m agi na ry a ct i on of a fict i tious
agency ; b ut a n a scert a i ned ac t i on o f a known ag ency .
I t is p rop er to a dd s om e Monocotyl ed ons

th a t there a re ,
as Ra vena l a m a da gas ca ri ens i s i n whi ch thi s ch a ract er d oes

,
not occu r Wh a t conditi ons of exist enc e th ey a re th at here

.
*
h old this n atu ral t end ency i n ch eck it is not ea sy to see , .
The N a tu r a l H i s tory Revi ew for J ul y

, 1 8 65 , cont a i ned an a rt i cl e on
the doc t ri ne of m orphol ogi ca l compos i ti on set fort h i n th e foregoing Ch ap s .
I t o I II
. . I n thi s a rt i cl e, whi ch uni te s ex p os i t i on a nd c ri t i ci s m i n a way th a t
is unha p
p i l y not com m on wi th re v i ewe rs , i t i s s ugges t ed th at the s pi ra l s t ru e
t u re m a y b e ca u s e d by na tur al s el ec ti on When thi s a rt i cl e ap peare d, t he
.
foregoi ng fi ve p age s were v

s ta ndi ng o e r in type , a s surpl us from No 1 4 , i ss ued
.
J
in une, 1 8 6 5 .
CH A P TE R XI I I .
M O RP H O L O GI CA L D I F F E RE N T I A T I ON IN I
A N MAL S .
2 42 . genera l c ons id era ti ons which p rel u d ed ou r i n

TH E
qu i ry i nt o th e s h a p es of pl ants and th ei r p a rt s equ a lly serve , ,
so far as th ey g o to p re l u d e a n i nqu i ry i nto th e s h a p es of

,
a n im a ls a n d t h e i r p arts A m ong a n im a l s a s a m ong pl a nt s

.
, ,
the form a ti on of a ggr eg a tes g r ea ter i n b u lk or high er i n de

g ree of c om p os iti on or b oth ; i s a cc om p a ni ed by ch an ges of
,
for m i n the a gg reg a t es a s wh ol es as well a s by c h a n ge s of

f or m i n th ei r p a rt s ; a nd the p roce s ses of m or ph ol ogi ca l
d i fferenti a ti on con for m to the s a m e genera l l a ws i n th e one
ki ngd om a s i n the oth er .
I t is needl es s to rec a pitu l a t e the s ev era l ki nd s of m odi fi

c a t i on to be expl a i n e d a n d th e s ev era l
, fa ct or s th a t co
op era t e i n w or ki n g th e m I n so fa r a s th ese a r e c om m on
.
to pl an t s a n d a n i m a l s the p r ece di ng ch a pt ers h a v e s u f

,
fi ci entl y fa m ili a ri ze d th em N or i s it need fu l to s p ec i fy

.
a fres h the s ev era l typ es of s ymm et r y a nd th e i r d esc r ip t iv e
na m es ; for w h a t i s t ru e of th em i n t he on e c a s e i s t ru e o f
th em i n the oth er T h ere i s h owev er one n ew a nd a l l

.
, ,
i m p ort a nt f act or whi ch we s h a ll h a v e n ow to t a k e i nt o

a ccou n t ; a nd a b ou t thi s a few p r elimi n a ry r em a r ks are
r equ i s it e
.
2 4 3 T hi s n ew
. f act or i s m oti on— m oti on of th e org an i s m
in l ti on to
re a s u r r ou n di ng obj ec t s o r o f th e p a rt s of th e
,
1 83
or g a nism i n rel a ti on to one a noth er or b oth T h ou gh th ere

, .
a r e pla nt s e s p ec i a lly of th e s i m pl er ki n d s whi c h m ov e a nd

, , ,
th ou gh a few o f the s im pl er a n i m a l s do not m ov e ; yet m ov e

m ents a re s o e x c ep ti on a l a n d u n ob tru s iv e i n th e one ki n g
d om w hil e th ey a re s o gener a l a n d c on s pi cu ou s i n th e oth er

, ,
th a t the b roa d di sti n ct i on com m on ly m a d e i s w ell w a rr a n te d .
Wh a t a m on g pl a nts i s a n i n a pp reci a bl e cau se of m orph o

, ,
l ogi ca l di fferen ti ati on b ecom es a m on g ani m a l s the c hi e f

, , ,
c a u s e of m or ph ol ogi ca l di ffe r enti a t i on .
Root ed a n i m a l s or a n im a l s oth erw i se fix e d of c ou rse p r e ,
s en t t r a its of s t ru ctu r e n ea r es t a ki n t o th os e we h a v e l a tely
b een stu dyi n g T he m oti ons o f p arts i n r el a ti on to one anoth er

.
a n d t o th e e n vi ron m en t b ei n g g ov erne d by th e m od e of a
, ggr e
g a ti e n a n d m od e of fixi ng we a re p res ent ed wi th m or p ho
,
l ogi ca l di iferenti ati on s s i m il ar i n th ei r gen era l ch ar acter s to

th os e of pl a nt s and sh owi ng u s p ar al l el ki nd s of s ymm etry
,
u nd er p ara ll e l c on di ti on s B u t a n i m a l s whi ch m ov e from

.
pl a ce to pl a ce a re su bj ect to a n a ddi ti on a l c l a ss of a cti ons

a n d rea cti ons . T h ese a cti on s a n d rea cti ons a ffect th em i n
v a ri ou s way s a ccor di ng to th ei r v a ri ou s m od es of m ov e m e nt .
L e t u s gl a nce a t the s ev er a l l ea di n g rel a t i ons b et w een s h a p e

a n d m oti on w hi ch w e m ay exp ect to fi nd .
I f a n org a n i s m a dv anc es th r ou gh a h om ogeneou s m edi u m

w ith one en d a l w ays f ore m os t t h a t en d b e i ng e xp ose d to
, ,
f orces u nlik e th os e to whi ch the oth er e nd i s e xp os ed m ay ,
be e xp ec te d to b e c om e u nlik e it ; a n d su pp os i ng t h i s to be
th e on ly c ons ta nt c ont ra s t of c on diti on s we m a y e xp ect a n ,
eq u a l di s t rib u ti on of the p a rt s r ou n d th e a xis of m ov e m en t
a ra di a l s y m m e try . I f i n a ddi ti on to t h i s h a bi tu a l
,
a tti tu d e of th e e nd s one s u r fa ce of th e b ody i s a l wa y s u pp er

,
m os t a nd a noth er a l w a y s l ow e r m ost th er e a r i se b etw een th e

,
t op a nd b ott om di s s imil ari ti es of conditi on s whil e the two ,
sid es rem ai n s i m il a rly condi ti oned H ence it i s i nf era bl e

.
th a t su ch a n org a ni s m w ill be divi s ibl e i n to simil a r h a lves

by a v erti ca l pl a ne p ass i ng th o gh t xi
r u i s a s o f m ti
o on — w ill
h av e a bil a t era l symm et ry We m ay p resu m e th at this
.
MORP H O L OGICA L D I FFE RE NTIA T ION I N A NI MA L S .
1 85
sym m et ry w ill d evi a t e bu t littl e from d ou bl e bil a t er a l nes s

wh e r e th e u pp er a nd u nd er p a rts ar e not exp ose d to stron gly
c on tr ast e d i n fl u en ces ; whil e we m ay ra ti on a lly l ook for
si ngl e bil at era l sy m m et ry of a d ecid e d ki nd i n crea tu r es
,
h a vi ng d ors a l a nd v ent ra l p art s con v ers an t w i th v ery u n lik e

regi ons of th e en vi r onm en t : a s i n a l l c a s es w h e re th e m ov e
m ent i s ov er a s olid su r fa c e . I f the m ov em ent ,
t h ou gh ov er a s olid s u r f ace i s not c on st a n t i n di r ecti on bu t

, ,
t a k es p l ace as oft en on one s id e a s on a noth er ra di a l sym

,
m et ry m a y be a g a i n l ook ed for ; a nd if the m ot i ons a re s till

m or e v a r i ou s ly di rec t e d— i i th ey a r e n ot limit e d to a p prox i
m a t ely pl an e su r faces bu t ext end to su r f aces th a t are d i s

-
,
tr ibu t ed a l l a r ou n d w i th a r egu l a r i rregul a r ity— a n a pp roach

o f th e r a di a l t ow a r d s th e s ph e r i ca l s ymm et ry i s to be a nti c i
p a te d
. Wh er e th e h a bit s a re s u ch th a t th e i nt er
c ou r se b etw een t h e or g a n i s m a n d i ts envi r on m ent d oes n ot
,
i nv olv e an a v era g e equ a lity of a cti ons and rea cti on s on a n y

two or m ore s id es th ere m ay be exp e ct e d eith er t ot a l i r regu
,
l ar i ty or s om e div ergence from regu l a r ity .
T he lik e gen era l rel a ti on s b et ween for m s and i nc id ent

force s a r e i nfer a bl e i n th e com p on en t p a rt s of an i m a ls as ,
we ll a s i n the a ni m a l s a s wh ol es I t i s n eedl ess h ow ev er to

.
, ,
oc c u py s p a c e by d e s c r ipti on s of th es e . L et u s n ow p a s s to
th e fa cts a nd s ee h ow th e y c onfi rm d p os teri ori the c oncl u
, , ,
si ons here rea ch ed d pr i ori

. .
C H A P TE R X I V
TH E GE N E RA L I
S H A P E S OF A N M A L S .
§ 2 4 4 CE RT A I N of the P rotozoa are qu it e i n d efinit e i n th ei r

.
sh a p es and qu it e i n con st ant i n th ose i nd efi n it e sh ap es whi c h

,
th ey h a v e — the rel ati ons of th ei r p a rt s a r e i nd et er mi na t e

b oth i n s p ac e an d ti m e I n one of th e s i m pl er Rhi 2 0pods at
.
,
l ea st d u ri ng the a ctiv e s ta ge of i ts exi st enc e no p er m anent ,
di s ti ncti on of i nsid e and ou t s id e i s est abli sh ed ; a nd h ence

there can a r i se no est a bli sh e d c or res p ond enc e b etween the
sh a p e of the ou t sid e a n d the di st r ibu ti on of envi ron i ng
a cti on s . Bu t wh en th e r el a ti on of i nn er and ou t er b ec om es
fix ed e ith er ov er p art of the m ass or ov er the wh ol e of it we
, ,
h a v e ki nd s of sym m et ry th a t corres p ond with the h abitu a l

i n cid ence of forces A n A m ccba i n b ec omi n g en cy s t ed
.
,
p a sses from an i n d efi nit e ever ch an gi ng for m i nt o a sph eric a l

,
-
form ; a nd the or d er of s ymm et ry whi c h it thu s a ssu m es is ,
I n h a r m on y wi t h th e a v era ge e u a lity of th e a cti ons on al l

q
I n D i fil u gza Fig 1 3 4 a nd s till b ett er i n A r c el l a
’
i ts s id es .
, .
, ,
w e h a v e a n i n defi ni tel y ra di a l sym m et ry occ u rr i n g wh er e th e

-
conditi on s a r e di f ferent a b ov e a nd b el ow bu t a lik e al l a roun d .
A m on g the Gr ega r i ni da th e s ph er ic a l s y m m et ry an d sym

m et ry p ass i ng f rom th a t i nt o the r a di a l are s u ch as a pp ear
,
to be c ong ru ou s with the s i m pl e c i rcu m st a nc e s of th ese

c r ea t u re s i n the i nt esti n es of i ns ect s B u t th e r el a ti ons of
.
th es e l owes t typ es to th ei r envi ronm ents are comp a ra tiv ely

s o i nd et er m i n a t e a nd ou r k n owl e dg e of th e i r a c ti ons s o
,
T HE GE NE RA L S H A P E S OF A NI MA L S .
1 87
sc an ty th a t littl e b ey ond
, ne a g tiv e evid enc e ca n be e xp ect ed
f rom the s t u dy of them .
T he lik e m ay be s a id of th e I nfus oria T h ese ar e m or e or .
l ess i rregu l ar I n s om e c a ses w h er e the li n e of mov em ent

.
,
thr ou gh the wat er is t ol erably d efi n i te a n d c on s t ant we h a v e ,
a f orm th at is a pp roxim a t ely ra di a l— e xt er na lly a t l ea st .
Bu t u su a lly a s s h own i n Fig s 1 3 7 1 3 8 1 3 9 th er e i s e ith er

,
.
, , ,
a n u n sy m m et r i ca l or a n a sy m m et r i ca l sh a p e A n d wh en one .
of th e se c rea t u r es i s w a tch e d u n d er th e m i cr oscop e th e con ,
ru i t of this h a p e w ith t h e in c id e n c e of f or c es i s m a n if es t
g y s .
For th e m ov e m ents a re conspi cu ou s ly v a r i e d a n d i ndeter m i

n a t e —m ov em ents w hi ch d o not exp os e a ny tw o or m ore sid e s
*
of the m a ss to a pp r oxim a t ely e qu a l s ets of a cti on s .
§ 24 5 . A m on g
gg r e g a t e s ofa th e s ec on d or d er as a m on g ,
a gg re g a t e s of th e fi r s t or d e r we fi nd th a t of th os e p os s ess i ng
,
.
a ny d e fi n it e sh a p es th e l ow es t a re s ph er ic a l or s ph er oid a l .
Su ch a r e som e of the Ra di ol a ri a a s C ol l ozou m i n er m e T h ese , .
bo di es which fl oa t p a s s iv ely i n the s ea a nd p res ent i n t u rn ,
a l l th e i r s id es to th e sa m e i n fl u enc es h a v e th ei r p a rt s di s ,
p osed with app roxi m a t e r egu l ar ity rou n d a c ent re— a pproxi
m a te be cau s e i n th e a bs ence of l ocom oti on a slight i rregu
,
l a r i ty of gr owth a lm os t c ert a i n to t a k e pl ace m ay c a u se a

, ,
fix ed a ttit u d e an d a resu lti ng d evi ati on from s phe r ic a l sym

m et ry T h e b es t c a ses i n ill u st r a ti on of th e t ru th h er e
.
na m e d a r e fu rn i s h e d by rot a ti n g a n d l ocom otiv e or g an i s ms

,
res p e cti ng whi c h th ere i s a di s p u t e wh eth er th ey a r e a n i m a l
or v e g et a l— th e Vol voci n ew T h es e a l rea dy i ns t anc e d u n d er

.
,
“
A v eri fying com m e nt on thi s p ara g ra ph r un s a s fol l ows I n th e
Hypot ri ch a I nfus ori a , whi ch c re ep o v
er sol i d s u rfa ce s , t h ere i s a di ffere n
ti a ti on be t we en v entra l a nd dors a l s u r fa ce a nd a n a p p r oa ch to bi l a t era l s ym
m e t ry . The v ent ra l s u rface i s p ro v ided wi t h v

m o a b l e c il i a , th e dors al w i th
”
i m m obi l e setae.
1 88 MOR P H OL OGI CA L D E V E L OP ME N T .
th e on e h ea d in h ere be i n s t an ce d a f resh u nd er
m ay
th e oth er F u rth er a m ong th ese sec on d a ry a ggr eg a tes i n

.
,
w hi c h th e u ni ts on ly phy s i c a lly i nt eg r a t e d h a v e not h a d th ei r

, ,
i ndivid u a li ti es m erg ed i nt o an i n divid u a lity of a high er

or d er m u s t be n a m e d th e c omp ou n d I n us ori a
, f T h e c l u s ter
.
of Vor ti ce l l ce i n Fig 1 4 4 w ill s u ffi c i en tly ex empli fy th em ;

.
,
a n d the s t r iki ng res e m bl a n c e b orn e by its i n divid u a l s t o
thos e of a ra di a lly a r r ange d c l u ster of fl ow ers wi ll s h ow how

-
, ,
u n d er a n a l og ou s c on di t i ons th e g en era l p r i ncipl e s of m or

,
p h ol og i c a l diff erenti a ti on a re simil a r ly ill u st ra t e d i n th e two
ki ngd om s .
§ 24 6 . Ra di a l
ym m et ry i s u s u a l i n l ow a gg r eg at es of
s
th e s e c ond or d er which h a v e th e i r p a rts su ffi ci ently differen

ti s t e d a n d i nt eg ra t ed to giv e i ndivid u a li ti es to th e m a s wh ol es .
T h e Cccl cn ter a ta offer nu m erou s e x a m pl es of thi s S oli tary .
p olyp es — hyd roid or h el i anthoi d— m ostly s t ati onary a nd ,
w h en th ey m ov e m ovi n g w i th a ny s id e fore m o s t do n ot by
, ,
l oc om oti on s u bj ect th ei r b odi es to h a bit u a l c ontr as ts of con

di ti on s .S eate d w ith th e i r m ou th s u p w a r d s or d ow nwa rd s ,
or e l s e a t a l l d eg r ee s of i nc li n a ti on the i ndivid ua l s of a
,
s p ec i es t a k en t og e th er a r e s u bj ec t to n o m ech a ni ca l ac ti ons
,
a ff ecti n g s om e p a rt s o f th e i r di scs m ore th a n oth e r p a rt s .
A nd thi s i n d et ermi n a t en es s of a t titu d e s i m il ar ly p r ev ents

th ei r r el a ti ons to p r ey f rom b ei ng su ch a s su bj ec t s om e of
th ei r p r eh en s il e or g a ns to forces u nlike th os e to whi ch the
r es t a r e s u bj e ct T h e fix e d end i s di ffe r ently con di ti oned
.
f rom the free en d and th e two a re therefore di fferent ; bu t

,
a rou n d th e a xi s ru nni n g f rom th e fix e d to th e fr ee en d th e
c onditi on s a re a lik e i n al l di recti ons a nd th e for m th ere fore

,
i s ra di a l . A g a i n a m ong m a ny of th e s i m pl e free
,
s wimmi n g H ydrozoa th e sa m e g enera l tru th i s ex e m p l ifi ed

,
u n d er oth e r c i r cu m s t a n c e s I n a c omm on M edus a a dva nc

.
,
i n g th r ou gh the w at er by th e r hythmi ca l c ont rac ti ons of i ts

di sc th e m ech ani cal r ea cti on s are the s a m e on a l l s id es ; a nd
,
a s f ro m a c c id ent a l c a u ses e v ery p ar t of th e e dge of the di s c

, ,
T HE GE NE RA L S HA P E S OF A NI MA L S .
1 89
m es u pper m ost i n i ts t u rn no p a rt i s p erm ane ntly a ffect e d

’
co ,
i n a di fferen t way f rom the re s t H ence th e r a di a l for m c on .
ti nu es .
In th ers
o m e g rou p h ow ev er th ere oc cu r form s
of this sa , ,
w hi ch s h ow u s a n i n c ipi en t bil a ter a l nes s ; a n d h e lp u s t o s ee

h ow a m ore d ec id ed bil a t e ra l ness m a y a ri se S u n d ry of th e .
M edus i daz a r e p rolife rou s givi n g origi n to g em m ae from th e

,
body of the cen tr a l p olypi te or f rom c ert a i n po i nts on th e

e dg e o f th e di s c ; a nd thi s b u ddi ng u n l ess it occu r s eq u a lly ,
on a l l sid e s whi c h it d oe s not a nd i s u n lik ely to do m u s t

, ,
t en d to d es troy th e b a l anc e of the di s c a nd to m a k e i ts ,
a tt it u d e l e s s ch a n g ea bl e I n oth er ca s es th e g row th of a
.
l a rg e p roces s [ a m u ch d ev el op e d t ent a cl e ] f rom the edge of

-
the di s c on one s id e a s i n S teens tru p i a Fig 2 5 7 c ons tit u t es

.
, ,
.
,
a s i m il a r m odifi ca ti on a n d a c a u s e of fu r th er m o difi ca ti on
,
.
T he a n i m a l i s no l ong er divi s ibl e i nto a ny t wo qu i te s i m il a r

h a lv es ex cept th os e for m ed by a pl a n e p a s s i ng th rou gh th e
,
p rocess ; a nd u nl ess th e p roces s i s of th e sa m e s p ecific g ra vi ty

a s the di s c i t m u s t t en d t ow ard s e i th er the l ow es t or th e
,
high est p oi nt an d m u st so s erv e to i n c rea se th e bil at era l nes s

, ,
by k eepi ng the tw o sid es of the di s c s imil a r ly c onditi one d

whil e the top an d b ott om a re diffe ren tly condi ti one d Fig 2 5 8 . .
r e p r esent s th e u n d e rs id e o f a n oth er M edus a i n w hi ch a m or e ,
d ec id e d bil a tera l ness i s p rodu c ed by th e p r es enc e of two s u ch

p rocesses . A m ong th e
s i m pl e f ree swi m mi ng A cti-
n oz oa ,
oc c u r lik e d evi a ti on s
from r a di a l s y m m etry a l on g ,
wi th lik e m oti ons th r ou gh th e

wa t er i n bil a ter a l a t tit u d es .
Of t hi s a Cydi p p e i s a f a m ili a r
‘
ex a m pl e T h ou gh r a di a l i n
.
som e of i ts ch ara cters as i n ,
the di str ib u ti on of i ts m er idi

.
on a l b a n d s of l ocom ot iv e p a ddl e s with th e i r a cc omp a nyi n g
c a n a l s thi s c r ea tu r e h a s a tw o s id ed dist r ib u t i on o f t en t a cl e s
,
-
a nd v ar i ou s oth er p arts corr esp ondi ng wi th its two sid ed

,
-
a ttit u d e i n m ovi n g th r ou gh th e w a t er A n d i n oth er g en era

.
o f thi s g r ou p a s i n C es i u m
,
E ur ha m p hwa a n d C a l l i a ni ra
, , ,
th a t a l m os t equ a l di st ribu ti on of p arts whi ch ch ara cteri zes

the B er oe i s qu it e l os t .
H er e s eem s a fit pl a c e to m eet th e obj ecti on which s om e

m ay fee l to this an d oth er s u ch ill u st ra ti on s th a t th ey a m ou nt ,
v ery m u c h to phy s i ca l t ru i sm s I f th e p a rts of a M edus a a re

.
di s p osed i n ra di a l s y m m et ry rou n d th e a xi s o f m oti on th rou gh

t he wa t er th ere w ill of cou r s e be no m ea ns of m a i nta i ni n g
,
on e p a rt of its edge u pp er m os t m or e th a n a noth er ; a nd th e
e qu a lity of conditi ons m ay be a scr ib e d to th e r a di a t eness as ,
m u ch a s th e r a di a t eness to the equ a lity of con diti ons Con .
ve rse l y wh en th e p a rt s a re not ra di a lly arra n g e d a r ou n d th e

,
a xi s of m oti on th e y m u s t g ra vit a t e t owa r d s s om e on e a tti

,
t u d e implyi ng a b a l a n ce on the two s id es of a v ertic a l pl an e

,
— a bil a ter a l nes s

; a nd th e two s id ed c on diti ons s o n ec es si
-
t a ted m ay be a s m u c h a sc r ib e d to the bi l atera l nes s a s th e

,
bil atera l nes s to the two s id ed con diti ons

-
D ou bt
.
l es s th e f orm a n d th e conditi ons a re i n the w ay all eged , ,
n ec essary c or r el a t es ; a nd i n so far a s it a ss erts thi s the oh ,
jecti on h a rm on i zes with the ar gu m ent T o th e di ffic u lty

.
which it a t the s am e ti m e ra i s es by the i m pli ed qu esti on

Why m a k e th e for m the resu lt of the c onditi ons ra th er th an ,
th e con di ti on s the r esu lt of the for m ? the reply i s thi s

.
T h e ra di a l typ e b ot h a s b e i ng the l east di ff erenti a t e d typ e

,
a n d as b ei n g th e m os t obvi ou s ly re l a t e d to l ow er typ es m u s t ,
be t a k en a s a nt ec ed ent to th e bil a t era l typ e T h e i ndi .
vid u a l v ar i a ti ons whi ch i nc id ent al c i r cu mst a nces p rod u c e i n

t he ra di a l typ e will not c au s e div erg ence of a s p ec i es from
,
th e r a di a l typ e u nl es s s u ch v a r i a ti on s giv e a dv a nt a ges to th e

,
i ndivid u al s di spl ayi ng th em whi ch th ere i s no r ea son to su p

p ose th ey w ill a l wa ys do T h ose occ a si onal d evi ati ons f rom
.
th e ra di a l typ e w hi ch th e l a w of th e i ns ta bility of th e h om e
,
gen ee ns w a rr a nt s u s i n exp ecti ng to t a k e pl ace will h owev er , , ,
i n s om e c a s es be b enefici al ; a nd w ill th en be lik ely to e st a b

T HE GE NE RA L S H A P E S OF A NIMA L S .
1 91
li s h th ems elv es S u c h d e vi a ti on s m u s t t end to d es t roy th e

.
or igi n a l i n defi n i ten ess a n d v a r i a bili ty o f a ttitu d e— m u s t c au se
g ra vi tati on t ow a r d s a n h a bit u a l a tti tu d e A n d g ra vit ati on .
t ow a r d s a n h a bi tu a l a ttit u d e h a vi ng once c om m ence d will ,
c onti nu a lly i nc rea s e where in crea s e o f i t i s not neg a tiv ed by

,
a dv er s e a g en c i e s : e a c h f u rt h er d eg ree of bil a t e r a l n ess r en d e r
i ng m ore d ec id ed th e a cti ons th a t c on d u c e to bil at er a l ness If .
this reply be th ou ght in s uffic i ent it m a y be en for c ed by the ,
f u rth er one th at a s a m ong pl ant s th e i ncident force s a re the

, , ,
a n t eced en ts a nd th e f or m s th e co n seq u en t s ( c h a n g es o f forc es
be in g i n m any ca s es vi s ibly foll owed by ch a nges of for m s ) we

a r e w arr ant e d i n c onc l u di n g th a t th e lik e ord er of c au s e an d
*
ef f ec t h olds am ong a nim a l s .
§ 24 7 . Keepi n g to the s a m e typ e bu t p a s s i n g to high er a
d egr ee of c omp os iti on w e m eet m or e ,

c om pl ex a n d v a ri e d
illu strati on s of th e s a m e genera l l aws . I n th e c omp ou n d
C i ti ci sm s on th e a bo v e p as s a ge h a v e s h own
r the need for na m i ng s un
dry co m pl i c a ti ons The s e com pl i ca t i on s chi e y, i f not wh ol l y, ari s e from

. fl
ch a ng e s i n m od e s of li fe—c ha nge s fro m th e l oco m ot i v e to th e sta ti ona ry, a nd
fro m t h e s t a t i ona ry to t h e l oc om ot i ve . Re fe rri ng t o y s t a t em ent th a t (ignor
m
i ng th e S ph eri ca l ) th e ra di a l t ype i s th e l ow es t a nd m us t b e t a ken a s a nte

h
ce dent to t e b i l a t e ra l t e
yp , i t i s a ll eged t h a t a l l e x i s t i ng
“
ra di a l a ni m a ls
a bov e P roto zoa a re p rob a bl y d e ri v e d from free s wi m m i ng, b il a te ral l y-sym -
”
m e t ri ca l a ni m a l s . If thi s i s i nt ended to i ncl ud e th e p l a nul as of th e hydroi d
pol yps , t he n i t s ee m s rat h e r a s tra i ni ng of th e e vi denc e Th es e l oc o m oti v e .
e m bryos , des cri bed as se v era l l y h a v i ng th e s t ru ct ur e of a gas t ru l a wi th a

c l os ed m ou th , c a n be s ai d to s how bi l a te ral ne ss onl y b eca us e th e fi rs t t wo t en
t a el e s m a ke t h ei r a pp e ara nce on Op po si t e s i des of th e m ou th — a bi l atera l nes s
w hi ch l as t s onl y ti l l t w o othe r ten ta cl es m ake th e i r a pp e a rance in a pl a ne a t
ri gh t a ngl e s , so gi vi ng th e ra di a l s t ructu re I thi nk th e cri ti ci s m . a pp l i es onl y
to cas es furn i s h ed by E chi noderm s Th e l a rv ae of th e s e c rea tures h a v e bil a t

.
eral l y-s ym m e t ri ca l s tru c tu re s , wh i ch th ey r e t a in as l ong as th ey s wi m a bo ut
a nd wh i ch s u ch of th em as fi x th e m s el v es
l os e by bec om ing s im il a rl y rel a te d
to co ndi ti ons al l round : th e ra di a l s t ru c ture be i ng ret a i ned by th ose typ es
whi ch , be com i ng s ub s e q u ent l y det a ch ed , v

m o e a b out m i s c ell a neous l y . Bu t, as
h a ppe ns i n som e o f th e Se a -u rch i ns a nd s t i l l m o re a m on g th e Hol oth uri a ns ,

th e s t ructu re is aga i n m a de bil a tera l l y-sym m e tri ca l by a l ocom ot i v e l i fe pu r
s ue d wi t h one end forem os t . Shou l d i t be contende d t h a t the condi ti ons a nd
the for m s a re re c i pr oca ll y i nfl uenti a l — th a t e i th er m a y i ni ti a te th e oth e r, it
s til l rem a i ns u n q ues t i ona bl e tha t ordina r il y the condi ti ons are th e a nteced ents ,
as is s o a b unda ntl y s ho wn by pl a nts .

C ael en tem ta , p res enti ng c lu s t er s of i ndividu a ls whi ch a re

s ever a lly h om ol ogou s with th e s olit ary i n divid u a l s l a s t d ea lt
w ith we h a v e to n ote b oth th e S h ap es of th e i ndivid u a l s t hu s
,
u n i te d a nd th e s h a p e s of th e a gg r e g a te s m a d e u p of th e m
, .
S u c h of th e fix e d H ydr ozoa a n d A c ti n ozoa a s for m b r a n ch e d

s oc i et i es c ont i nu e ra di a l ; b ot h b eca u se t h ei r v a r i e d a tti tu d es
,
do not exp ose th em to a pp r ec i a bl e differe nc es i n th ei r rel a

ti ons to those s u rrou n di ng a cti ons whi ch chi efly c onc ern
th em ( th e a cti ons of p rey ) an d b ecau s e su ch di ffer en ces ev en
, ,
i f th ey wer e a pp rec i a bl e w ou ld be s o a v era ge d i n th e i r effects

,
on the di s s imil a r ly pl ace d m emb er s of e ac h g r ou p a s to be

-
n eu tr a li ze d i n th e r a c e A m on g the t ree
.
like cora l pol ypedom s a s well as i n s u ch

-
,
r a m i fi ed a s s e m bl a g es of S i m pl er p olyp e s
a s are S h own i n Fig s 1 4 9 1 5 0 we h a v e

.
, , ,
i nd eed c a s es i n m a ny res p ec t s p a r a ll el
,
to th e c a ses of s c attere d fl owers

whi c h th ou gh pl a ced l a t era lly rem a i n
r a di a l b ec au s e no differenti a ti ng agency
,
c an a ct u n i formly on a l l of th em Meanwhil e i n the

.
,
g rou p s whi c h th ese u n it e d i ndividu a l s com p os e we see the ,
sh a p es of pl a nt s fu rth er s i m u l a t ed u nd er a f u rth er p a ra ll eli s m

of c on diti on s T he a t ta ch e d end s di ffer f rom the free end s
.
a s th ey do i n pl a nt s ; a nd th e re gu l a r or i rre gu l ar b ra n c h es
o bvi ou sly st a n d to e n vi ron i n g a cti on s i n r e l a ti ons a n a l og ou s
to th ose i n w hi ch th e b ra nch e s of pl a n t s s t a nd .
'
T h e m e mb er s of th ose com p ou n d Cazl en tera ta whi ch m ov e
th rou gh the wa t er by th ei r own a cti ons i n a tti tu d es th a t a re
,
a pp r oxim a t ely c on s t ant S h ow u s a m ore or l ess di sti n c t t wo

,
s id ed nes s D i p hyes Fig 2 5 9 fu rni sh es an e x a mpl e

.
,
.
,
E ach .
of th e l arg e ly d e v el op e d a nd m odifi e d p olypit es for m i ng its

-
s wi m m i ng s a cs i s bil a t era l i n c orresp on d ence with the bi

,
l a ter a l nes s of i ts c onditi on s ; a nd i n ea ch of the a pp end e d

p olypit es the i ns er ti on of the s olit a ry t ent a cl e p rodu ces a
ki nd re d div erg en c e from th e p rimitiv e radi a l typ e T he .
a gg reg a t e too which h ere v er y m u ch s u b or di na t es i ts m em

, ,
TH E GE NE RA L S HA P E S OF A N IMAL S .
93
bers ,
e xhibits s a m e conf or mity of st ru ctu re to ci rcu m
th e
s t a n c es I t a dmits of sym m et ric a l bi s ecti on by a pl a ne p a ss
.
i ng th rou gh its two c ont ra ctil e s a c s or nect oc a ly ces bu t not , ,
by any other pl ane and the pl ane which thu s symm et r ic ally
bi sec ts it i s the v erti ca l pl ane on the two sid es of whic h its
,
r e s i m i l ar l y c on di t i one d a s it p r op els it s el f t h rou gh

p
'
a rt s a
th e wa t er .
A n oth er g rou p of th e oc ea n i c H ydrozoa the P hysop hori dce , ,
fu rn i s h es i nt er esti n g e vid en c e — not s o m u c h i n resp ec t of th e

f orm s of the u n i ted i n dividu a l s whi ch we m ay p a s s ov er a s
, ,
i n r esp e ct of th e for m s of th e a gg r eg a t es S om e of th ese

.
a r e with ou t s w immi n g or g an s a n d h a v e th ei r p a rt s s u s
,
p end ed from a i r v e s sel s w hi c h h a bi tu a lly fl oa t on the su rfa c e

-
of th e w a ter H en c e th e di st r ib u ti on of th ei r p ar t s is a sym
.
m et ri cal T he P hys a l i a Fig 1 5 2 i s an ex a mpl e H ere the

.
,
.
, .
r el a ti on s of th e i nt eg r a t e d g rou p of
i ndivid u a ls to th e envi ron m en t are i n

d efi nit e ; a nd th er e i s thu s no a gency
t en di n g to ch a nge t h at c omp ara tiv ely
i rre gu l a r m od e of growt h whi ch i s pro
b ably d er iv ed f rom a p r i m or di a l typ e
of th e b ra nc h e d H dr ozoa
y .
S o v ar i ou s a r e th e m od es of u ni on
a m on g th e c omp ou n d C wl en tem ta th a t ,
i t i s ou t of th e qu es ti on to d ea l w ith
t h em a l l E v en did s p a c e p erm it it
.
,
wou ld be i m p r a ctic a bl e for a ny one bu t

a p r ofes s e d n a tu r a li s t to t r a ce th rou gh
,
59
1 94 MORP H OL OGI CA L D E V E L OP M E NT .
ou t thi s g rou p th e
l ti ons b etw een sh a p es a n d c ondi ti ons of
re a
e xi ste n ce T he ab ov e m u s t be t ak en s i m ply a s a few of the

.
m os t s ig nifi can t an d ea s ily i nt erp reta bl e ca ses -

.
24 8 . I n th e s u b-
ki n gd oms P ol yz oa an d T u ni ca ta w e m eet
w ith ex a m pl es n ot w h olly u n lik e th e for egoi ng A m on g th e .
typ es assem bl ed u nd er th ese n a m es th e re a re s impl e i n di vi

d u a l s or a ggreg a tes of th e s ec on d ord er a nd soci eti es or ,
t ert i a ry a ggreg ates p r odu ced by th ei r u ni on T he rel a ti ons .
of for m s to f or ce s h a v e to be t ra c e d i n b oth .
S oli tary A sc idi a ns fix e d or fl oa ti n g c a r ry on a n i na ctiv e

, ,
a n d i nd e fin i te c onv e r se wi t h th e a cti on s i n th e e nvi r onm en t :
Wi th ou t p ower to m ov e a b ou t viv a ci ou sly a nd u n abl e to ,
ca tc h a ny p rey bu t th a t c onta i ne d i n th e c u r r ents of w a ter
they a b s orb a nd exp el th ese c rea tu res a re not exp ose d to

,
s et s of for c es which are eq u a l on two or m ore S id es an d th ei r

sh a p e s consequ ently rem a i n v a gu e T h ou gh i nt ern ally th ei r .
p a rts h a v e a p a r ti a lly sy m m et r i ca l a rrangem ent du e to th ei r

-
,
d er iv a ti on yet th ey are su b s t anti a lly u n sy m m etric a l i n th a t

,
p a rt o f the b ody whi ch i s concer ne d w ith th e en vi ronm ent .
Fig 1 5 6 i s a n ex ampl e
.
*
A m on g th e c omp os it e
.
A sc idi a n s fl oa ti n g an d fix ed th e sh a p e of the a gg rega t e

, , ,
p artly d ete rmi ned by th e h a bi tu a l m od e of g em m ati on a nd

p artly by the s u rrou n di ng conditi ons i n each c a s e i s i n ,
g rea t m ea s u re i nd efi n it e We ca n s ay no m or e ab ou t it th an
.
th a t it i s not obvi ou sly a t v ar i an ce with the l aws a ll eged .
E vid en ce of a m ore p os i tiv e ki n d occu r s a m on g th os e com

p ou n d M ol l us coi da whi ch a re m os t lik e th e c omp ou n d
C ael en ter a ta i n th ei r m od es of u n i on— th e P ol yzoa Ma n y o f .
th ese form g rou ps th a t a re m ore or l es s i rregul a r— s p re a d

i ng a s film s ov er s olid s u r f a c es combi ni n g i nt o sea weed ,
-
lik e frond s b u ddi n g ou t from c reepi ng s tol ons or g rowi n g

, ,
up i nt o tfee sh ap e d s oci eti es ; and b es id es a gg reg a ti ng

-
Shoul d i t be p ro v ed t h a t the A scidi a n i s a degra ded v er teb ra te , th en th e

a rgum e nt w i l l b e s t rengt h ened ; s i nce l oss of bi l a t era l sym m etry has gone
a l ong wi th ch a nge to a sym m et ri ca l co nd i t i ons .

T HE GE NE RA L S H A P E S OF A NIMA L S .
1 95
i rre gu l arly th e y a re i rre gu l a rly pl a c ed on su r fa c es i ncli ned

i n a l l di re c t i ons Merely not i ng th a t thi s as ym m et ri ca l
.
di s t r ibu ti on of th e u ni te d i n divid u a l s is e xpl a i ne d by the

a b s e n ce of de fi n i te n es s i n the re l a t i on s of t he a gg re g a t e to
i n cid en t forces it concerns us chi efly to ob ser v e th a t th e

,
u n i te d i n divid u a l s s ev e r a lly e x em pli fy the s a m e t ru th a s do
s imil a r ly u n it e d i n divid u a l s a m on g the C ael en tera ta

-
A vera g .
i ng the m e mb ers of ea ch s oci e ty the c ili a te d t ent a cl e s th ey ,
p rotru d e ar e S imil a rly rel a te d to p rey on al l sid es ; and

th ere for e re m a i n the s am e on a l l s ides T hi s di st r ibu ti on of .
t e nta cl es i s not h ow e v er wi thd u t e x cepti on

, ,
A m qn g th e .
fre s h w a ter P ol yzoa th ere a r e s om e gen er a a s P l u m a tel l a an d

-
,
Crys ta tel l a i n w hi ch th e a r ra ngem ent of th es e p a rt s is v ery

,
d e cid edly bil a t er a l S om e s p ec i es of th e m S h ow u s s u ch

.
r el a ti on s of th e i n divid u a l s to on e a noth er a n d to th e i r su r
f ace of att ach m en t as giv e a clu e to th e m odific a ti on ; bu t

,
i n oth er s p ec i es th e m ean i ng of this d evi a ti on f rom the

ra di a l typ e is not obvi ou s .
§ 24 9 . good e x a mpl es of the con

I n th e P l a tyhel m i nth es
n ex i ons b etw een f or m s a nd f orc es occu r T he P l a na ri a .
e x em plifi es t he s i ng l e bil a t er a l sym m e t r y which e v en i n ,
v ery i n fer i or form s a ccom p a n i es the h abit of m ovi ng i n one

,
di recti on ov er a solid s u rface H u m bly or g a ni ze d a s a re .
th ese c rea tu r es an d th ei r a lli es th e N em er ti dce w e see i n ,
th em j u s t a s c l ea rly a s i n the high est ani m a l s th at wh ere

, ,
the m ov em ents s u bj ect the b ody to di fferent f or c es a t its two

end s ,
differ en t for ces on i ts u n d er an d u pp er su r fa ces a n d ,
lik e for c es a l on g i ts two S id es th ere a ri s es a c orres p ondin g

,
for m u nlik e a t i t s e xtrem i ti e s u n lik e a b ov e a nd b el ow bu t

, , ,
h avi ng i ts tw o s id es a lik e .
The E c hi noder m a ta fu rn i sh u s with i n st ru ctiv e ill u st r a

ti on s — i n st ru ctiv e b ec au s e a m on g t yp es th a t a r e nea r ly a lli ed ,
w e m eet w i th wid e d evi a ti on s of f orm a n s w er i n g to m a rk e d

contr a s ts i n th e r el a ti on s to th e e n vi r on m en t The f a ct s f a ll .
i nto fou r g rou p s T he C ri noi dea onc e s o a b u nd a nt

‘
.
,
a nd n ow so p res ent a r a di a l sy m m etry answeri ng to

r a r e,
a n in c id en ce o f for ce s th a t a r e e qu a l on a l l s id e s I n th e .
g enera l a tti tu d es of th ei r p ar ts t owa r d s s u rrou ndi ng a cti ons ,
t h ey a re lik e u ni a xi a l pl a n ts or lik e p olyp es ; a n d s h ow a s ,
the se do m a r k e d di ffe r ences b etw een th e a tta ch e d en d s a nd

,
t he free en d s a l ong w i t h ev en di str ib u ti on s of p a rt s a l l rou n d

,
t h ei r a x e s . I n th e Op hi u ri dea a n d i n the S ta r ,
fi shes we h a v e r a di a l sym m et ry co e xi s ti ng wi th v ery di ffer

,
-
e nt h a bi ts ; bu t h a bits w hich n ev erth el es s a c cou n t for th e
m a i nt enance of th e f orm H oldi n g on to roc k s a nd weed s

.
by i ts S i m pl e or b ra nche d a rm s or by the s u ck er s b orn e on ,
t h e u nhe r su r f ace of i ts ray s one of th ese c r ea tu r e s m ov es ,
a b ou t n ot a l w ay s w i th on e s id e f or e m os t bu t w i th a ny s id e ,
fore m os t .Cons equ ently a v e ra gi n g i ts m ov em ents i ts a rm s

, ,
or r ay s a re equ a lly a ffec te d a n d th erefore re m a i n th e s a m e

,
on a l l sid e s . On w a tch i n g the way s of th e com

m on S ea u r chi n -
w e a re s i m il arly fu rn i s h e d w i th a n e x
,
pl a nati on of i ts s p h eri ca l or ra th er i ts s ph eroid a l figu re

, ,
.
H er e th e h abit is not to m ov e ov er any on e a pp roxim a tely

fl a t su r fa ce ; bu t th e h a bit i s to h old on by s ev era l su r f a c es
on di ffe r ent s id e s a t th e s a m e time F requ enti n g c revi ces .
a n d t h e i nt ers ti ces a m on g s t ones a n d w eed s th e S ea u rchi n ,

-
p rot ru d es the su ck ers a rr an ge d i n m eridi ona l b and s ov er i ts

s h e ll l ayi ng h old of obj ec ts n ow on thi s S id e a n d n ow on th a t
, ,
n owa b ov e a nd now b el ow : th e resu lt b e i n g th a t it d oes n ot
m ov e i n a l l di r ec ti ons o v e r on e pl a n e bu t i n al l di r ecti on s
th rou gh s p a c e H ence the a pp roach i n gen era l for m t owa rd s

.
S ph e r i c a l s y m m e t ry — an a pp roa ch w hi c h i s h owev er re , ,
s tr a i ne d by the r el a t i on s o f th e p a rt s to th e m ou th a n d v ent
th e c on diti on s not b e i ng e x a ctly th e s am e a t t he t wo p ol es a s

a t oth er p a rt s of th e s u r fa c e S till m or e s ig nific a nt i s
.
th a t d evi a ti on from thi s sh a p e which occu r s a m ong s u ch of

th e E chi ni dca a s h a v e h a bi ta ts of a di fferent ki nd a nd c on ,
s eq u e ntly di f ferent h a bi ts
,
T he genera E chi u ocya m us S p a
.
,
ta n g us B r i ss us a n d A m p hi d o tus div er ge m a rk edly t owa rd s

, , ,
a bil a t era l s t ru c t u re T h es e c r ea tu res a re f ou n d n ot on rocky

.
T HE G E NE R A L S H A P E S or A NI MA L S .
1 97 .
shores bu t on fl at s ea - b ottom s, a nd
th e m only on s om e of
b ott oms of sa n d or m u d H er e th ere i s n one of th a t di str i

.
,
bu ti on of su r f a ces on a l l S id es w hi ch m a k es th e s ph er oid a l
for m c ong ru ou s with th e c on diti on s H a vi ng to m ov e a b ou t
.
ov er a n a pp roxi m a t ely h or i z onta l pl an e a ny d evi ati on of

-
,
st ru ct u r e a ri s i n g a cc id e nt a lly w hi c h l ea d s to on e s id e b e i n g
k ept a l wa ys forem os t will be a n a dv a n t a ge : g rea t er fit n es s

,
to f u ncti on b ecom i n g p oss ibl e i n p r op or ti on a s f u n cti on

b ec om es fix ed S u r viv a l of the fitt est will th erefore t end to
.
e s t a blish u n d er s u c h c on diti on s a for m th a t k eep s th e s a m e

, ,
p a rt i n a dv a nc e — a for m i n w hi ch consequ en tly the or igi na l , ,
r a di a l s ymm et ry div er g es m or e a n d m or e t owa r d s bil a t er a l
sym m et r y .
§ 2 5 0 Very d efi nit e a nd comp arativ ely u niform a r e the

.
,
rel a ti on s b etween S h a p es and c i rcu m sta nces a m ong the

A nnu l os a : inclu di ng u nd er th a t titl e th e A nn el i da a n d th e
’
A r thr op oda T he a g reem ents a nd th e di s a g reem ents a re
.
equ a lly i nst ru ctiv e .
A t on e ti m e or oth er of i ts li fe i f not th rou gh ou t i ts li fe , ,
e v e r y a nnu l os e a n i m a l i s l ocom otiv e ; a nd i ts t e mp ora ry or
p erm a nent l oc om oti on b ei ng c a rr i ed on w i th one en d h a bi tu

,
a lly fore m os t an d on e su r face h a bit u a lly u pp e rm os t i t fu lfils

,
th ose c on diti ons u n d er whi c h bil a t er a l s ym m et ry a r i ses .
A c c ordi ngly bil a t er a l sy m m et ry i s t ra cea bl e th rou gh ou t th e

,
wh ol e of this su b ki ngd om T r a c ea b l e
-
w e m u s t s ay
.
, ,
b ec au se thou gh it is ext rem ely cons picu ou s i n th e i m m ense

,
m a j or ity of a nnu l ose typ es it i s to a cons id era bl e e xt en t

,
ob scu re d wh er e obsc u r a ti on is to be e xp ec t ed T h e e mb ry os .
o f th e T u bi c ol ae a ft er s w immi ng a b ou t a whil e
, s ettl e d ow n ,
a nd bu ild th ems elv es t u b es from whi ch th ey p rotru d e,
th ei r h ea d s ; a nd in th em or i n s om e of th em the bil a t er a l
, ,
s ym met ry is di s g u is e d by the d ev el opm en t o f h e a d a pp e n d
a ges i n an a l l S id ed m a nn e r
-
T he t ent acl es of T er ebel l a a r e
.
di st ribut ed m u ch I n the sa m e way a s th ose of a p olyp e T h e .
b reathi ng org ans i n S abel l a u m s pi m , Fig 2 6 0, do not c orre .

s pond on Opp os it e s ide s of a m edi an pl an e E ven h ere how .

,
e v er th e b o dy r et a i n s i t s p r i m itiv e bil a t era l n es s

, a n d it i s
furth er to be r em ar k e d th a t thi s l oss of bil a t e r a l ness i n the

ext er na l a pp en d a g es d oes not occu r wh er e the r el a ti ons to
,
e xt ern a l c onditi ons con ti nu e bil a t era l : w it ness t h e S er u l a

p ,
Fig 2 6 1 whi ch ha s its r espi ra tory t u fts arranged i n a two

.
,
sid ed way u n der the two sid ed c onditi ons i nv olved by th e

,
-
h abit u a l p ositi on of its t u b e .
T he c omm u nity of s ymm et ry a m ong the high er A n nu l osa ,
h a s an u nob ser v e d S ign ifi c an c e T h a t Fli es B eetl e s L ob .

, ,
ster s Centip ed es S pid ers Mit es h av e i n c om m on th e c ha

, , , ,
ra cter s th a t the end which m ov es i n a dv a n c e di ffer s f rom

,
th e hi nd er end th a t th e u pp er su r fa c e differs f rom the u n d er

,
s u rf ac e a nd th a t the two S id es a re alik e i s a t ru th rec eiv ed

, ,
a s a m a tt er o f c ou rs e A ft er a l l th at h a s b een s a id a b ov e
.
,
h owev er it will be s een to h a v e a m eani ng not to be ov er

,
l ooked ; S i nce it s u ppli es a milli on fold illu st rati on of th e l aw s -
w hi ch h a v e b een set forth I t is needl es s to giv e di a gr ams

. .
E v ery re a d e r ca n c all to mi n d th e u n ity i ndi ca t ed .
While h owever a nnu l ose a ni m al s rep ea t so u niformly

, ,
th es e t ra it s of st ru ct u re th ere a re c ert a in oth er t ra it s i n

,
.
whi ch th ey a re v ar i ou sly c ont ra st e d ; and th ei r c ont ra sts

h a v e to be h ere not ed as s ervi ng fu rth er to b u ild u p the
,
g enera l a rgu m ent I n th em we s ee the st a ge s th rou gh which

.
T HE GE NE RA L S H A P E S OF A NIMAL S .
1 99
bil at era l s ym m et ry b ecom es gr a d u a lly m ore m a rk ed a s the ,
con diti on s it re s p on d s to b ec om e m or e d ecid e d . A

c om m on E arth w or m m ay be i n s t a nc e d
-
as a m e mb er of
this su b ki ngd om th a t is a m ong th e l ea s t c onspicu ou sly
- -
bil a tera l
. T h ou gh i nt erna lly its p a rt s h a v e a tw o S id e d -
a rra n g em e nt ; a n d th ou gh the p os iti on s of its or ific e s giv e it
a n e xt e rn a l two s id e d nes s a t the s a m e ti m e th a t th ey e st a b

-
,
li s h a di fference b etween the two en ds ; yet i ts two S ide dnes s -
i s n ot st r ongly m a rk e d
-
T he f orm d evi a t es bu t littl e f ro m
.
wh a t we h a v e di s ti n gu i s h e d a s t ripl e bil a t er a l sym m et ry : i f

th e c rea t u re i s cu t a cr oss th e middl e th e h ea d a nd t a il ends
,
a r e v ery m u c h a lik e ; if cu t i n two a l ong i ts a xi s by a h or i
zonta l pl ane th e u n d er a n d u pp er h a lv e s a re v ery mu ch

,
a lik e ext er n a lly if not i n t er n a lly ; a n d i f ou t i n two a l on g its

,
a xi s by a v e rtic a l pl a ne th e two s id es a re qu it e a lik e

,
.
Fig s 2 6 3 a nd 2 64 will m a k e this cl ear

. S u c h crea tu r es
.
a s th e J u l us a nd th e C en tip ed e m a y be t a k en a s sh ow i n g
,
a t ra n s iti on t o d ou bl e bil a t era l s ymm e t ry Besid es b ei n g

.
divi s ibl e i nt o ex a ctly s imil a r h alv es by a v ertic a l pl ane p a s s

i ng th rou gh i ts a xi s on e of th ese a nim a l s m ay be bis ect e d
,
t ra ns vers ely i nt o p arts th a t di ffer only slightly ; bu t if ou t i n
two by a hori zont al pl an e p ass i ng th rou gh its axis the und er ,
a n d u pp er h a lv es a re d e cid e dly u nlik e Figs 2 6 5 2 6 6 . .

, ,
e xhibit th es e t r a i t s A m ong th e i sop od ou s c ru st a c eans

.
,
th e d ep art u re f rom th ese l ow typ es of symm et ry i s m ore

m arked . As
h own i n Figs 2 6 7 and 2 6 8 the c ont ra st
s .
,
b et ween th e u pp er a n d u n d er p a rts is gr ea ter a nd th e h e a d ,
a nd t a il en d s di ffer m or e obvi ou s ly I n a l l th e high er

.
A r thm p oda th e u nlik en e s s b et ween th e fron t h a lf a n d th e

,
hi nd h a lf h as b ecom e cons pi cu ou s T h ere i s i n t h em s i ngl e .
bil a t er a l symm et ry of s o p r on ou nced a ki n d th a t no ot h er ,
re s embl ance i s su gg es t e d th a n th a t b e t w een th e t wo s id e s .
B y Figs 2 6 9 a nd 2 70 re p re s enti n g a d ec a p od ou s c ru sta cea n

.
,
divid e d l ongitu di nally an d t rans vers ely thi s t ru th i s m a d e ,
m a ni fest . On c a lli ng to mi n d the h a bit s of th e

c r eat ur e s h ere d rawn a nd d es c rib e d it will be s een th a t ,
they expl a i n th es e f orms Th e i ncid ence of forc es i s th e

.
s a m e a l l a rou nd th e E arth wor m a s it bu r rows th rou gh th e

-
c om p a ct g r ou n d T he Centip ed e cre epi ng a m id l oose s oil or

.
,
débri s o r b enea th s t ones i ns i nu a t es it s el f b et ween s olid su r

,
f aces— the i nter s ti ces b ei ng m ostly g rea t er i n one di m ens i on

th a n i n oth ers A n d al l th e high er A nnu l osa m ovi ng a b ou t
.
,
as th e y do ov er e xp os e d obj e ct s h a v e th e i r d or s a l a n d v en t ra l
,
p a rt s a s di s simil a rly a ct e d u p on a s a re th ei r two ends .
One oth er fact only r es p ec ti ng annu l os e a nim a l s n eed s to

be notic e d u nd er thi s h ea d — the fa ct n a m ely th at th ey , ,
b ecom e u nsymm et ri cal wh ere th ei r p art s a re u nsymm et ric

a lly r el a te d to th e envi r on m ent T he c omm on H erm it cra b
.
-
s erv e s a s a n i n st a n ce H ere i n a ddi ti on to th e u nlik eness o f

.
,
th e tw o s id e s i m pli ed by th a t c u r v atu re of th e b ody w hi c h

fi t s th e crea t u re to the s h ell it i nh a bit s th ere i s a n u nlik enes s ,
du e to th e grea t er d e v el op m ent of the li m bs a nd esp eci ally ,
the cl a ws on the ou t er sid e

, A s i n th e e mb ry o of the
.
H ermit cra b th e two s id es a re a lik e ; a n d as b oth the emb ry o

-
a nd th e a nc e st or liv ed i n s u c h a w a y b ei ng f ree , ,
th at the conditi ons w ere a lik e on th e two s id es ;

a nd a s th e e mb ry o m a y be t a k en to r e pre
s ent the typ e f rom which the H er mi t cra b h as -
b een d eriv ed ; we h a v e i n thi s c a se evid ence

th at a symm et ric ally bil at era l form h as b een -
m ou ld ed i nt o an unsymm et ric ally bil at era l -

TH E GE NE RA L S H A P E S OF A NI MA L S .
01
f orm by the acti on of u nsy m m et r i ca lly bil a t era l condi ti ons

,
-
.
A fu rth er illu s tr ati on i s su ppli e d by B opyr us Fig 2 7 1 : a ,

.
p a ra s it e whi ch liv es i n the b ra nchi a l ch a mb er of p ra wns a nd ,
wh os e h a bit s s imi l a rly a cc ou nt for i ts di s t ort ed s h a p e .
25 1 . A m ong the M ol l us ca we find m ore v a r i e d l i

re a t ons
b et ween sh a p es a nd ci rc u mst ance s S om e of th es e

. re l a ti ons
ar e highly i n s t ru c tiv e .
Moll u s k s of one ord er the P terop oda s wim i n the sea

, ,
mu ch i n the s a m e wa y th at b u t te rfli es fly i n
the a i r a nd h a v e sh a p es not a lt oge th er u n lik e
,
th os e of bu tt erfli es Fig 2 7 2 rep resent s on e

. .
of th es e c r e a tu res T h a t i ts bil a t e r a lly s ym

.
-
m et ri ca l s h a p e h a rm oni zes with i ts bil a ter

a lly s ymm et ric a l
-
c onditi on s is s u ffici ently
obvi ou s .
A m ong the L a m el l i bra nchi a ta we h a v e ,
diverse f orms a c comp anyi ng div ers e m od es of li fe Su ch .
of th e m a s f requ ently m ov e a b ou t like the f resh wa t er ,

-
Mu s sel h a v e th ei r two v a lv es a nd th e c ont a i ned p a rt s

,
a l ik e on th e opp os it e S id e s o f a v ert ic a l pl a n e : th e y a re
bil a tera lly symm etr ic a l i n conformity w ith th ei r m od e of

m ov em ent T he m ari ne Mu s s el
. too th ou gh h a bi tu a l l y
, ,
fix ed a nd th ou gh not u su a lly s o fix e d th a t i ts two v a lv es a re

,
s i m il ar ly c on diti oned s t ill ret a i ns th a t bil a tera l s y m m et r y

,
whi c h i s c h a r a c ter i s ti c of t h e ord er ; a n d it d oes thi s b eca u se

i n th e s p eci es c ons id er ed a s a w h ol e th e two v a lv es a re n ot
,
di s s i m il a rly c onditi oned I f th e p os iti ons o f th e v a ri ou s

.
i ndivid u al s a r e a v e ra ged it will be s een th a t the di fi eren

,
t i a ti ng a cti on s neu tr a li ze one a noth er I n c e r ta i n

.
o th er fix ed L a m ellib ra nc h s h ow ev er th ere i s a con s id era bl e

, ,
d evi ati on from bil a t era l s ymm et ry ; an d it is a d evi a ti on of

the ki nd to be anticip a t ed und e r the c i rc u m sta nce s Wh ere .
one v a lv e is al w ays d own w ar ds or n ext to t he su r f a c e o f

,
att a chm ent whil e the oth er v a lv e is a lw ays u pw a r ds or n ext

, ,
to the envi roni ng w at er, we m ay exp ect to fi nd th e two

2 02
'
MORP H OL OG IC A L D E VE L OP ME NT .
v a lves b ecom e u n lik e T hi s w e do fi nd : w it ness the Oy ster

. .
I n the Oy s ter too we s ee a fu r th er i rre gu l a r i ty

, , T h ere i s a .
grea t i ndefi ni teness of ou tli ne b oth i n the s h ell a nd i n the

,
a n im a l— a n i n defi ni teness m a d e m a n i fe st by c om p ar i ng di f
fer ent i n divid u a l s We h a v e bu t to re m em ber th a t growi n g

.
c l u s ter ed t ogeth er a s Oy s t e r s do
, they m u st i nterf ere w i th
,
on e a noth er i n v a r i ou s w a y s a n d d eg rees to s ee h ow th e ,
i n d et ermi n a tenes s of for m and the v a r i ety of form a re

a c c ou nt ed for .
A m ong th e Ga st erop od s m odifi c ati on s of a m ore d efi n it e

I n a l l Moll u s ks
”
ki n d occu r . s ay s P rofes sor H u xl ey
, ,
th e a xi s of the body is a t fi r s t s t r a ight a nd i ts p a rt s are ,
a r r a n g e d s ym m et r i c a lly w i t h reg a r d to a l ongi tu di na l v e rt i c a l

”
pl a ne j u s t a s i n a v ert eb ra t e or a n a rt icu l a te emb ryo
,
In .
s om e Ga s t er op o d s a s th e C hi ton
,
thi s bil a t era l s y m m et ry
,
i s ret a i ned — th e re l a ti ons of the b ody to su rrou ndi ng a c ti ons

not b ei n g s u c h a s to di s tu rb it B u t i n th os e m or e nu m erou s
.
typ es whi c h h av e s pi ra l s h ell s th ere i s a m ark e d d evi a ti on

,
f rom bil at era l s ymm et ry as might be exp ect ed

,
T his .
a s ymm et ric a l ov er —d ev el op m ent nev er a f fec ts the h ea d or

”
f oot of the m ollu s k : only th ose p art s w hi ch by i ncl osu re ,
i n a s h e ll are p rot ect ed f rom envi ron i ng acti on s l ose th e i r

, ,
bil a t er al ness ; whil e the e xt erna l p a rts s u bj ecte d by the ,
m ov em ents of th e c rea tu res to bil a t era l conditi ons r em a i n ,
bil a t era l H ere h ow ever a di ffi cu lty m eet s u s Why i s i t

.
, ,
.
th a t the na k ed Ga s terop od s s u ch a s ou r c om m on s l u gs
,
‘
,
d evi at e from bil a tera l s ym m etry th ou gh th e i r m od es of

,
m ov em ent a re th os e a l ong w i th w hi ch c om pl ete bil a t era l

s ymm et r y u s u a lly o c c u r s ? T h e reply i s th at th ei r d evi a
ti ons from bil a t era l s ym m et ry are p rob ably i nh eri ted a nd ,
th a t th ey a re m a i n t a i ned i n s u ch p a rt s of th ei r org ani za ti on

as a r e n ot e xp os e d to bil a t e ra lly sy m m etric a l condi ti on s
- .
T h ere i s reason to be li ev e th a t th e n a k ed Ga st erop od s a r e

d escend ed f rom Gast erop od s which h a d sh ell s : the evid enc e
be i ng th a t the n a k ed Ga st erop od s h a v e sh ells d u ri ng the
ea rl y st a g es of th ei r d ev el opm en t a n d th a t s om e of th e m
,
T HE GE NE RA L S H A P E S or A NIMA L S .
203
t i
r e a n ru di m en t a ry s h ell s th rou gh ou t li fe N ow the s h ell e d .
-
Ga s t erop od s d evi a t e fr o m bil a t er a l s y m m e t ry i n t he di s p os i

t i on of b ot h th e a li m en ta ry sy s t em a n d th e rep r od u c tiv e
sy st em .T he na k e d Ga sterop od s i n l os i n g th ei r s h ell s h a v e , ,
l ost th a t i m m ens e one s id e d d ev el op m en t of the a li m enta ry

-
sy s tem whi c h fi tted t h em to t h ei r s h ell s a n d h a v e ac qu i re d ,
t h a t bil a ter a l sy m m et ry of e x tern a l fi g ur e w hi c h fi ts th e m

for th ei r h a bit s of l oc om oti on ; bu t th e r ep rod u c tiv e s y st e m
re m a i n s on e S id ed b eca u se i n res p ec t t o it t h e r e l a ti on s t o
-
, , ,
e x tern a l con di ti on s r em a i n one s id e d -

.
T h e C eph a l op od s S h ow u s bil a t era lly sy m m et ri ca l e xt erna l -
f orm s a l ong w it h h a bi ts of m ov em en t th rou gh th e w at er i n

t wo s id e d a ttit u d es
-
A t th e s am e tim e i n th e r a di a l di s tri
.
,
bu ti on of the a rm s ena bli n g one of th ese c r ea tu res to t ake

,
a n a l l s id ed g r a s p o f i t s p rey w e see h ow r ea dily u p on on e

-
,
ki n d of sy m m et ry th ere m ay be p a rti a lly d ev el op ed a noth er

ki n d of symm etry wh ere th e r el ati on s to conditi ons f a v ou r it
, .
§ 25 2 T he Ver tebr a te ill u st r a t e a fres h th e t ru ths which

.
we h a v e a l rea dy t r a ced a m on g the A n nu l osa Flyi n g th rou gh .
th e a i r s wi m m i n g th r ou gh th e w a ter a n d r u nn i ng ov er th e
, ,
ea r th a s v er t eb ra t e a nim a l s d o i n c om m on w ith a nn u l os e,
a n i m a l s th ey a re i n com m on w ith a nn u l os e a n i m a l s di fferen t

, , ,
a t th ei r a n te r i or a n d p os t er i or en d s differen t a t t h ei r d o rs a l ,
a n d v en t r a l s u r fa c es bu t a lik e a l on g th ei r tw o S id es
, T hi s .
S i ngl e bil a ter a l sy m m etr y r em a i n s con s t a n t u n d er the e x
tr em es t m odifi c a ti on s of f orm A m on g fi sh w e s ee it a lik e

.
i n the h or iz on t a lly fl a ttened S k a t e i n t he v e rti c a lly fl a tten ed

-
,
-
B r ea m i n th e a l m os t sph er ic a l D i odon an d i n th e g r ea t ly
, ,
el ong ate d S n na thus A m on g reptil es th e T u r tl e th e S n a k e

y g .
, ,
a n d th e C r oc o dil e a l l di s pl a y i t A nd u n d er th e cou ntl es s .
modifi ca ti on s of s t ru ctu re di s pl ay ed by bi rd s an d m a mm a l s ,
it rem a i n s c on s pi cu ou s .
A l es s obvi ou s fa ct whi ch it c on cer ns u s t o n ot e a m on g th e

Ver tebr a te p a r a ll el to one whi c h w e n ot e d a m on g th e A n
,
u u l os a i s t h a t wh erea s th e l ower v ert eb ra t e f or m s d evi a t e

,
2 04: MORP H OL OGICA L D E VE L OP ME NT .
bu t littl e f rom t ripl e bil at era l s ym m et ry the d evi ati on be

,
c om es g rea t as we ascend . Fig s 2 7 3 a nd 2 74 Sh ow how

.
,
b esid es b ei n g divi s ibl e i nto s imil ar h a lv es by a v ertic a l pl ane

p a ss i ng th rou gh i ts a xis a Fi s h is di vi s ibl e i nt o h alves t h a t
,
a re not v ery d i ss i m il ar by a h or i z ont a l pl an e p a ss i n g th r ou gh
i t s a xis a n d a l so i n t o o th er n ot v ery di ss imil a r h a lv es by a

,
pl an e cu tti ng i t t ra ns v ersely I f a s s h own in Figs 2 7 5 a nd

.
, .
2 76 a n a l og ou s secti ons be m a d e of a su peri or Reptil e th e

, ,
divid ed p arts di ffer m ore d ecid edly Wh en a Ma mm a l a nd a

.
Bi rd t reat ed in the s am e way as sh own in Figs 2 7 7,

a re , .
2 78 a nd Figs 2 7 9 , 2 8 0, the p a rt s m a rk ed off by th e dividi ng

, .
pl a nes are unlike in far g reater d egrees On c onsid eri ng

.
THE GE NE RA L S H A P E S OF A NIMA L S .
2 05
th e m ech ani ca l c on v ers e b et ween org an i s m s o f thes e s ev er a l

typ es a nd th ei r envi ronm ent s — on rem emb er i ng th a t th e
fi s h h abit u a lly m ov es th rou gh a h om ogeneou s m edi u m of
n ear ly th e sa m e S p ecifi c g ra vity a s it s el f th a t th e t erres t r i a l
,
reptil e e i th er c r a w l s on th e su rfa c e or r a i s es i ts elf v ery i n
c om pl etely a b ov e i t th a t th e m ore a ctiv e m am m a l h avi ng

, ,
i ts su pp ort i n g p a rts m ore f u lly d ev e l op ed th ereby h a s th e,
u n d e r h a l f o f i ts b ody m a d e m ore di f fer en t f rom the u pp er

h a lf a nd th a t th e bi rd i s s u bj ect by i ts m od e of li fe to yet
,
a n oth er s et of a cti on s a n d r e a c ti on s ; we s h a ll s ee th a t th es e
f act s are qu it e c on g ru ou s with the gener a l d oct r i ne a nd ,
f u rni s h fu rth er su pp ort to it .
One oth er s ignifi ca nt pi ece Of evid en c e m u s t be n a m ed .
A m ong the A nn u l os a we fou n d u nsy m m etr i ca l bil a t era l n es s

i n c rea t u res h a vi n g h a bi ts e xp osi ng th em to u n lik e conditi ons
on t h e i r tw o S id es a nd a m on g th e Ver tebra ta w e fi n d p ara ll el
c ases T h ey a re p res ented by th e P l eu r onecti dw— the or d er

.
Of di s t ort ed fl a t fi s h es t o whi ch th e S ol e a nd th e Fl ou n d er
b el ong On the hyp oth es i s Of ev ol u ti on w e mu st concl u d e

.
,
th a t fi s h es Of this ord er h a v e a ri s en from a n or di n ary bil a

fera lly—sym m et r i c a l typ e Of fi sh w hi c h f eedi n g at the b ott om
, ,
of th e s ea g a i n e d s om e a dv a n t ag e by pl a c i ng it s elf w ith on e
,
Of i ts S id es down w ar d s i nst ea d of m a inta i ni ng the v ertic a l

,
a ttit u d e B es id es th e g ener a l rea s on th er e a re S p ec i a l

.
r ea son s for c on c l u di n g thi s I n th e fi rs t pl a ce the y ou ng

.
,
S ol e or Fl ou nd er i s bil a t er a lly s ymm et r i ca l— h as i ts ey es on

Opp os it e S id es of i ts h ea d an d swim s i n th e u su a l w ay I n .
the secon d pl a c e th e m et am orph osi s whi ch p rod u ces th e u n

,
sy m m etr i ca l s t ru ctu re s om eti m e s d oes n ot t a k e pl ace —th er e
a r e ab n or m a l Fl ou n d er s th a t s wi m v erti ca lly lik e oth er fi sh es

,
.
I n th e thi r d pl a c e th e t ra n s iti on fr om th e s ymm et r i ca l s t ru e

,
t u re to th e u ns ymm et r i c a l st ru ct u re m ay be tra ced A l m ost .
i ncredibl e th ou gh it seem s one Of th e ey es i s t rans f erred

,
from the u n d er s id e o f the h ea d to t he u pp er sid e : th e

- -
t rans fer b ei ng eff ected by a di st ort ed d ev el opm en t Of th e

cran i al b on es— a t rophy of som e a nd hy ert rophy of oth er s
p ,
l
a on g with a genera l tw i st T h i s m et a m orph os i s f u rni sh es
.
s ev era l rem a rk a bl e ill u stra ti ons of th e w ay i n which f orm s

b ec ome m ou ld ed i nto h arm ony w ith i nc ident forc es For .
b es id es th e div ergen ce from bil a t er a l sym m etry i n v olv ed by

p res ence of b oth ey es u p on th e u pp er s id e th ere i s a fu rth er ,
div erg ence from bil atera l s y m m etry i n volv ed by di fferenti a ti on

of th e two s id es i n r es p ec t t o th e c ont ou rs of th e i r s u r f aces
a n d th e s i z es Of th ei r fi n s A n d th en wh a t i s still m ore
.
,
s ig n ific ant th ere i s a n e ar a pp roac h t o lik enes s b etween th e

,
h alv es th a t w ere origi n a lly u n lik e bu t a re u n der the new

, ,
c i r cu m st a n c es exp ose d
,
t o lik e c on diti ons T h e b ody is .
divi s ibl e i nto si m il a rly Sh a p ed p arts by a pl an e cu tti ng it

-
“
a l on g th e s id e from h ea d to ta il th e d ors a l a nd v entra l
i nstea d Of the l a ter a l h a lves b ecom e symm et ric a l i n ou tli ne
”
a n d are e qu ip oi s ed .
253 . T h u s, th ere s eems i n common b etween th e

li ttl e as
sh ap es Of pl an t s an d th e S h a p es Of an i m al s we yet fi nd on , ,
a na ly s i s t h at th e sa m e g en era l t r u th s are d i s pl a y e d bv
,
b oth T he one u lti m a t e p ri ncipl e th a t i n a ny org a ni s m equ a l

.
a m ou nt s Of g r ow th t a k e pl a ce i n th os e di rec ti on s i n which
th e i n c id ent forc es a re equ a l s erv es a s a key to th e ph en om ena

,
of m or ph ol ogi c a l di f ferenti a ti on By i t w e a r e f u rn i sh e d
.
w i th i n ter p ret ati on s Of th os e lik enesse s and u n lik enes s es of

p a rts whi ch a re exhibit ed i n th e sev era l ki n ds of s ymm et ry ;
,
a nd wh en w e t a k e i n to a cc ou nt i nh er i te d eff ec ts w rou ght ,
u nd er a n c es t r a l c on diti on s c on t ra s t e d i n v a r i ou s w a y s w ith
p res ent conditi on s we a re ena bl ed t o c om p reh end i n a g en

, ,
e r a l w a y t h e a ct i ons by w hi ch a n i m a l s h a v e b een m ou ld ed
,
i nt o the s h a p es th ey p os s ess .
T o fill u p th e ou tli ne of th e a r gu m en t S O a s to m a k e it ,
corres p on d th r ou gh ou t w i th th e a r g um en t r es p ecti n g v eg et a l
form s i t wou ld be p rop er h ere to dev ot e a ch a pt er to the

,
di fferenti ati ons of th ose h om ol ogou s segm ent s ou t Of whi ch

an im al s Of cert a i n typ es a r e com p os ed T h ou gh a m ong m os t
.
,
a n i m a l s of th e thi r d d eg ree of c om p ositi on su ch as the ,

T HE GE NE RA L S H A P E S OF A NI MAL S .
2 07
roo et d H ydrozoa the P ol yzoa an d th e A s ci di oi da the u nit ed

, , ,
i n di vid u a l s a re not red u ce d to th e con di ti on of seg m en ts of a

com p os i te i n divid u a l ,
a n d do n ot di s pl a y a ny m a r k e d di ffer
e nti a ti ons ; y et t h e re a re s om e a n i m al s i n w h i ch su c h
s u bordi na ti on s an d c on sequ ent het erogenei ti es oc cu r T he

,
.
oc ea ni c H ydr oz oa f o r m o ne gr ou p O f th em ; a n d w e h a v e see n
reason to c on c l u d e th a t t h e A nnu l os a for m a noth er g rou p :
I t i s not w orth whil e h ow ev er to occu py sp a c e i n d eta ili ng

, , ,
these u nlikenesses of hom ol ogou s seg m ents an d s ee ki ng ,
sp ecific expl an a ti ons of th em A m ong the ocean i c H ydrozoa

.
th ey are ex trem el y v a ri ed ; an d the h a bit s a n d d er iv ati on s of

these creatu res a re so li ttl e kn own th a t th ere are n o a de
,
q u ate d a t a for i nt erp reti n g the f orm s Of th e p a rt s i n t erms

of th ei r rel a t i ons to th e envi ron m en t Con v er se ly a m ong
.
,
the A nn u l os a th os e di fferen ti ati ons O f th e h om ol ogou s s eg

m ent s whi c h a ccom p a ny t hei r p rog re ss i ng i nt eg r ati on h a ve ,
S O m u c h i n com m on ,
a n d h a v e g en er a l c a u s es w hich a re so
Obvi ou s th a t i t i s needl ess to d ea l w i th th em a t a ny l eng th

,
.
T h ey a re a l l expli c abl e a s du e to th e exp os u re Of di fferent

p art s of the ch a i n Of seg m en t s to diff eren t sets of a c ti on s and
r ea cti on s : th e m os t g en er a l c on tr a s t b e i n g t h a t b etwee n th e
a nt er i or s egm en t s a n d th e p os t eri or s egm en t s a n sw er i n g to

,
the m os t g enera l c on t ra s t Of c onditi ons to w hi c h a nnu l ose

an im a l s s u bj ec t th ei r s eg m en t s ; a n d t he m ore s p ec i a l con e
t ra s t s a ns werin g t o th e c ont r a s t s of c onditi on s en t a il e d by

th ei r m ore s p ec i a l h abit s .
Were an exh au s tiv e t rea tm ent of the su bj ect p ra cti c abl e ,
there s hou ld h ere al s o com e a ch ap ter d evoted to th e i n

, ,
t ern a l S t ru ctu res of a nim a l s— m eani n g m ore e sp eci a lly th e

, ,
S h ap es and arra n g em en t s of th e vi scera T h e r el a ti on s

.
b etw een form s a n d forces a m on g th ese i nc l osed p arts a r e ,
h owev er m os tly too ob scu re to a ll ow of i n ter p r et a ti on

,
.
P r ot ect ed a s th e vi scer a a r e i n grea t m ea su r e from th e i nc i

d en ce of extern a l forces we a r e not lik ely to fi nd m u ch
,
corres pon d en c e be tw een th e i r di s t r ib u ti on a n d th e di stri bu
ti on Of ext ern a l f orces I n thi s c ase the i nfl u en ces p artly

.
,
m echa nic a l p ar tly phy s i ol og i cal which the org an s exercis e

, ,
on one a noth er b ec om e t h e c hi e f c au ses of t h ei r c h a ng es of

,
fi g u re an d a rra ngem en t ; a n d th es e i n fl uences a r e com pl ex

a nd i n d efi n it e One genera l fa ct m ay i n d eed be n ot ed— the
.
, ,
fa ct n a m ely th a t th e di v ergenc e t ow a r d s a s ymm et ry whi c h

, ,
g enera lly ch a r acteri z es th e vi s cera i s m a rk e d a m on g th ose

,
o f th e m w hi c h a re m ost r em ov e d f rom m ec h a ni ca l conv er se

w ith th e en vi ron m en t bu t not s o m a rk ed a m ong th ose of
,
th em which a re l ess rem ov ed from su ch c on v ers e T hu s .
whil e th rou gh ou t the Ver tebra ta th e a lim ent a ry s y s t em

, , , ,
with the excepti on of i ts two e xt remi ti es i s a symmet ric a ll y ,
a rr a ng ed th e re s pi ra tory s y s tem w hi c h occ u pi es one end of

, ,
the b ody gen era lly d evi at es bu t li tt l e f rom bil atera l sym
,
m etry a n d th e rep rod u ctiv e sy s t em

,
p a rtly occu pyi ng th e
,
oth er end of th e b ody i s i n th e m a i n bil a t er a lly s y m m et r ic a l

,
s u ch d evi ati on from bil a t era l symm etry a s occu rs b ei ng ,
f ou nd i n i ts m ost i nt eri orly pl aced p a rt s the ov ari es J u st

-
,
.
i ndi cati ng th es e f a cts a s h a vi n g a c ert a i n s ignifi can ce it will ,
be b es t t o l ea v e thi s p a r t o f th e s u bj ect a s t oo i n v olv ed for

d et a il ed t reatm ent .
I n tern a l s tru ctu res of one c l ass h owev er n ot i ncl u d ed

, ,
a m on g th e vi s c e r a ad m it of gener al i n terp ret ati on— s t ru e

,
t u res whi ch thou gh i ntern a l a re b rou ght i n to t ol erably

, ,
di rec t rel ati on s w i th envi ron i ng forces a nd a re th erefor e ,
s u b ordi na t e i n th ei r fo r m s t o th e di s t rib u t i on of th ose for c es .
T h ese i nt ern a l s t ru ct u r es it w ill be d es i ra bl e t o d ea l w ith

a t s om e l en gth ; b oth b ec au s e th ey fu rn i s h i m p ort a n t ill u str a
t i on s en forc i ng th e g enera l argu m ent an d b eca u se a n i nter ,
r et a t i on Of th em w hi c h w e h a v e s een rea s on to r ej ec t ca n
p ,
not be r e j ected with ou t r a isi ng th e d em a n d for s om e oth er
i nt erp ret a ti on .
CHA P T E R X V .
TH E SH A P E S or VE RTE BRA T E S KE L E T O N S .
§ 2 5 4 WH E N a n el onga t ed m a s s Of a ny su b st a n ce i s
.
t ra nsvers ely st ra i ned different p a rt s of th e m a ss a re ex

,
p osed to f orc es of Opp os it e kin ds I f for ex a mpl e a ba r

.
, ,
of m et a l or wood is s u pp ort e d a t its two end s a s S h own i n ,
Fig 28 1 a nd h a s t o b ea r a w eight on its c ent re its l ower

.
, ,
p a rt is th rown i nto a s t a t e of t ensi on whil e its u pp er p a rt i s

,
th rown i nt o a s t a t e Of c omp ress i on A s wi ll be m a nifes t to

.
an
y one wh o ob s erves w h a t h a pp en s on b rea ki ng a sti c k
a c ros s h i s kn ee the g rea test d egree of t ens i on f a lls on th e
,
fi b res form i n g th e c onv ex s u r fa c e whil e th e fib res f or mi n g

,
th e c on c a v e su rfa c e a re s u bj ec t to th e gr ea t es t d eg ree of
com p re ss i on B et ween th es e e xtrem e s th e fib res a t differen t
.
d ep th s a re su bj ect to di ff erent f orc es P rogres m ng u p wa rd s

'
f rom th e u nd er su rfa c e of the ba r S h ow n i n Fig 2 8 1 th e .

,
t ens i on Of the fib res b ecom es l es s ; a nd p rogressi ng d own

wa r d s from the u pp er s u r fa ce th e comp ressi on of th e fib res
,
b ec om es l es s ; u ntil a t a c ert a i n dist a n ce b etween the two

,
su r fa ces th er e i s a pl a c e a t whi ch th e fib res a re neith er ex

,
t end ed nor c omp ressed T hi s s h own by th e d otted li ne i n

.
,
60 209
“
th e fi gure c a ll ed i n m ech a ni ca l l a ngu a g e the neu t ra l
,
is
a xi s . I t v a ri es i n p os i ti on w i th th e n a tu re of th e su b s t a n ce
s t r a i n e d : b e i n g i n c om m on pi n e woo d a t a di s ta nc e of a bou t
,
-
,
fi ve eigh th s of th e d ep th f rom th e u pp er su r fa c e or th ree

-
,
eigh th s f rom th e u n d er s u r fa c e C l ea r ly if s uc h a pi ece Of

.
,
w ood i n stea d Of b ei ng su bj ect to a d ownwa r d f orc e i s sec u re d

, ,
a t i ts en d s a n d s u bj e ct to a n u p wa r d f or ce th e di s t r ib u ti on
,
of th e c om p res s i on s a n d t en s i on s will be r ev er s e d a n d th e ,
neu t ra l a xis will be n ea rest t o th e u ppe r su r f a c e Fig 2 8 2 . .
r ep r es en t s t h ese Op pos it e a t tit u d es of th e ba r a n d th e c h a ng ed
pos ition Of i ts neu t ra l a xi s : the a rrow i n di ca tin g th e di rec

ti on of th e force p r od u ci ng th e u p wa r d b end a nd th e fa i nt ,
d otted li n e a s h ow i ng the p revi ou s p os iti on of th e n eu t ra l a xis

, .
Bet ween th e two n eu t r a l a xes w ill be s ee n a cent ra l s p a ce ;

a n d i t i s obvi ou s th a t w h en th e ba r h a s i ts s t ra in f rom tim e
to ti m e rev ers ed th e r ep ea ted ch a n ges Of its m ol ecu l ar con

,
diti on m u s t a ffect the c en tra l sp a c e i n a wa y di ff erent f rom

th a t i n whi ch th ey a ffect th e two ou t er sp a ces Fig 28 3 i s a . .
di a g ra m c onv eyi ng som e id ea of th es e c ont ra sts i n m ol ec u l a r

co n diti on I f A B C D be the middl e p a rt of a ba r thu s
.
t rea ted whil e G H a nd K L a re the a l terna ti ng neu t ra l

,
a x es ; th en th e for c es t o which th e ba r i s i n ea c h c a s e s u bj ec t ,
m a y be r ea dily S h ow n S u pp os i n g the d efl ec ti ng for ce to

.
be a c ti ng i n t he di rec ti on o f the a rrow E th en the t en s i ons

,
t o whi c h the fib res b etween G a n d F a re e xp os ed w ill be ,
re p res ent ed by a s er i es of li n es i n c rea s i n g i n l ength a s th e
di st a n c e from G i nc rea ses ; s o th a t the t ri a ngl e G F M will ,
exp res s the a m ou n t a n d di s t rib u ti on Of a l l the m ol e c u l a r
t ens i ons Bu t the m ol ec u l a r c om p res si ons th rou gh ou t th e

.
s p a ce f rom G to E m u s t b a l a n ce th e m ol ec u l a r t ens i ons ;

,
a nd h enc e i f the t ri a ngl e G E N be m a d e eq u a l to the tri

,
TH E S H A P E S OF VE RTE BRA T E S K E L E T ONS .
21 1
an gl e G F M the p a ra llel li nes Of whi ch it is compos ed ( h ere

"
d ott ed for the s a ke of di s ti ncti on ) will exp res s the a m ou nt

a nd di s t r ib u ti on of the c omp res si on s b et ween E a nd G .
S imi l a rly, wh en d efl ecti ng forc e i s i n the di recti on of the

th e
a rrow F t h e co m p r es s i on s a nd t ens i on s will be qu a n tit a tiv ely
,
symb oli zed by the t r i a n gl es K F O a nd K E P , . A nd

thu s the sev era l S p a ces occ u pi ed by fu ll li nes a nd by d otted
li nes a nd by th e tw o t ogeth er wil l r ep res en t th e different
,
a cti on s t o whi c h diffe ren t p a r t s Of th e t r a n s v er s e s ec ti on a re
s u bj ect by a lt ern a ti ng t ran sv ers e st ra i ns H ere th en it i s

.
, ,
m a d e m a n i fest to the eye th a t th e cent ra l sp a ce b etween G

a nd K i s di f
,
f erently c onditi on ed f rom the S p a ces a b ov e a nd
b el ow it ; a nd th a t the di fference of c on diti on is sh a rply
m a rk ed off T he fib res f orm i ng the ou t er su rf a c e C D a re
.
,
s u bj ec t t o vi ol en t t en s i on s a n d vi ol en t c o mp res s i ons
. P ro
g ress i n g i nw a rd s the tensi ons a nd comp ress i ons d ec rea se
the t en si ons th e m ore ra pidly . A s we a pp roa ch th e p oi nt G ,
the t en s i on s t o w hi c h th e fib res a re a lt erna t ely s u bj ect b ea r ,
s m a ll er a n d s m a ll er ra ti os to th e c omp r ess i on s a nd dis a pp ea r

,
a t the p oi n t G T h ence to th e c ent re occu r c omp ressi on s

.
only of a lt er n a ti n g i nt en s iti es b ecomi n g a t the c ent re sm a ll

, ,
a nd e u a q l ; a nd from the cent re w e a dv a nc e th rou gh a rev ers e ,
se ri es of c h a n g es to th e oth e r s id e
, .
T h u s i t i s d em on stra bl e th a t a ny su b st a n ce i n wh i c h th e
p ower of r esi sti ng com p ress i on i s u neq u a l to th e p ower Of
r es i sti ng t en s i on c a nn ot be su bj ect to a l tern a t i n g t r a n s v ers e
,
s tra i n s w i th ou t h a vi n g a centr a l p orti on di f

,
ferenti a t e d i n its
c on di t i on s from th e ou t er p orti on s a n d c onsequ ently dif ,
ferenti a ted i n i ts s t ru ctu re T hi s c on c l u s i on m a y ea s ily be

.
v erifi ed by exp eri m ent I f s om ethi ng h a vi n g a cert a i n

.
tou gh ness bu t n ot di ffic u lt to b rea k a s a thick pi ece of s h ee t

,
l ea d be b en t from s id e to s id e till it i s b rok en the s u r f a c e Of

, ,
f ra ctu re w ill exhibit a n u nl i k eness of t extu re be tween th e

i nner a nd ou t er p a rts .
§ 255 . A nd n ow for th e
ppli c a ti on o f
a thi s s ee mi ngly
i rrel ev a nt t ru th T h ou gh it h a s no Obvi ou s c onnecti on with
.
t h e i nterp r eta ti on Of v erteb ra l s tru ctu re we sh a ll S oon see ,
th a t i t f u nd a m ent a lly c oncerns u s .
T he s i m pl es t typ e of v ert e b ra t e a nim a l the fish h a s a , ,
m od e of l oc om oti on whi ch i nvolv es a lt ern a ti n g t ra ns v erse

s tra i n s I t i s n ot i nd eed su bj ect ed to a lt ern a ti n g t ra ns v er s e
.
, ,
s tr a i ns by s om e ou ter a g en cy a s i n th e c a s e w e h a v e bee n
,
i nv es tig a ti ng : it su bj ect s i t self t o t h em B u t th ou gh th e .
s t r a i ns a r e h e re i n tern a lly p r od u c e d i nst ea d of ext ern a lly
p rod u ced th e c a se i s not th erefore rem ov e d i nt o a wh olly

,
”4 different ca t egory For su p .

~
p os i ng Fig 2 8 4 to rep resent .
the ou tli ne Of a fi sh wh en
b e n t on on e sid e ( the d ott ed li ne s rep res en ti n g i ts ou tli ne
w h en th e b en d i s r ev er s ed ) i t i s cl ea r th a t p a rt Of th e s u b
,
s ta n c e for mi ng the c on v ex h a l f m u s t be i n a st a te O f t ens i on .
T hi s s ta t e Of t en s i on impli es the exi stenc e i n th e oth er h a l f

o f s om e c ou nt er b a l a ncin g c omp res si on
-
A n d b et ween th e.
two th ere m u st be a n eu tra l a xi s The w a y i n which this

.
c on cl u s i on is r ec oncil a bl e with th e f a ct th a t th ere i s t ens i on

s om ewh ere i n th e c onca v e sid e of a fi sh si nc e the c ur v e i s ,
T HE S H A P E S OF V E RTE BRA TE S KE L E T ONS .
21 3
c a u sed by mu scu l a r c ont ra cti ons on th e conca v e s id e w ill be ,
m a d e cl ea r by the ru de ill u st ra ti on whi ch a bow su ppli es .
A bow m a y be b en t by a th ru s t a g a i n s t i ts middl e ( th e t wo
en d s b ei n g h e ld b a ck ) or i t m a y be b en t by con tr a c ti n g
,
a s tr i n g t h a t u n i tes i ts en d s ; bu t th e di st r ib u ti on s of m e
ch a mi ca l forces wi th i n the wood Of th e bow t hou gh not qu i te ,
a lik e i n th e two ca s es will be v e ry s imil a r

,
N ow whil e the .
mu scu l a r a cti on on the conc a v e s id e of a fi sh di ffers from th a t

r ep r es en te d by th e tigh tene d s tri n g of a bow th e di ffer en c e ,
i s n ot s u c h a s to d est roy th e a pplic a bili ty of the ill u st r a ti on

the p a ra ll el h old s S O fa r a s thi s th a t w ithi n th a t p or ti on of
,
’
th e fi s h s b ody w hi ch i s p a ss iv ely b en t by the c ont r a c ti n g
'
m u s cl es th ere m u s t be a s i n a st ru ng bow a p a rt i n com

, , ,
p res s i on a p a rt i n t en s i on a n d a n i nt erm edi a t e p a r t whi ch

, ,
i s neu t ra l .
A fter thu s s eei n g th a t ev en i n th e d ev el op ed fi sh w i th

its compl ex l ocomotive a pp a ra tu s thi s l a w of the t ra ns v ers e
,
st ra i n h old s i n a qu a lifi ed wa y we sh a ll u n d ersta n d how

,
mu ch m ore it mu s t h old i n a ny for m th a t m a y be su pp osed

to i niti a t e th e v ert eb ra t e typ e — a for m d ev oid of th a t
s egm enta ti on by whi ch the v ert eb ra t e typ e i s m ore or l ess
ch a ra cteri z ed We S h a ll see th a t a ssu m in g a ru dim ent a r y
.
a ni m a l,
s till si m pl er th a n t h e A m p hi ox us t o h a v e a f eebl e
,
p ower of m ovi ng it self th r ou gh th e wa t er by the u nd u l a ti on s

of i t s b ody or s o m e p a rt o f i t s b ody th er e will n ecess a r ily
, ,
c om e i nt o pl a y c ert a i n rea cti ons whi ch m u s t affect the m edi a n

p or ti on of the u ndu l a ti ng m a ss i n a w a y u n lik e th a t i n
whi ch th ey a ff ec t i t s l a t er a l p or ti on s A nd if th ere e xi s t s i n
.
thi s m edi a n p or ti on a ti s s u e w hich k eep s i t s pl a ce wi th a ny

c ons t a n c y we m a y exp ect th a t th e di fferen ti a l c onditi on s
,
p rod u c ed i n it by the t ra n s v ers e s tr a i n w ill i niti a t e a dif,
ferenti a t i on I t i s t ru e t h a t th e d i s t rib u ti on o f th e vi s cera

.
i n t he A m p hi oxu s Fig 1 9 1 a nd i n t he typ e fro m whi c h w e

,
.
,
m a y su pp os e it t o h a v e a r i s en i s s u ch a s t o i n t er fe re w i t h this
,
p r oces s I t is a l s o t ru e th a t the a cti on s a n d rea cti ons de

.
s crib ed w ou ld not of th em s elv es giv e t o th e m edi an p orti on

21 4 MORP H OL OGICA L D E VE L OP M E NT .
a cyli nd ric a l s h a p e lik e th a t of the c a rtil a gi nou s r od ru nni ng

,
a l ong th e b a ck o f t he A m p hi ox us Bu t wh a t we h a v e h ere
.
t o not e i n t he fi rs t pl a c e i s t h a t th es e h a bit u a l a lt erna t e

,
fi exi ons h a v e a t end ency t o m a rk ofi f rom th e ou t er p a rt s

a n u n lik e i nn er p a rt which m ay be s ei ze d h old of m a i n
, ,
t a i ned a n d f u rth er m odifi ed by na t u ra l s el ecti on s h ou ld

, , ,
a ny a dv a nt a g e th er eby res u lt A n d we h a v e t o n o t e i n t h e
.
s ec on d pl a c e
,
th a t a n a dv a nt a ge i s lik ely t o res u lt T he .
c on t ra cti ons c a nn ot be effectiv e i n p rod u ci ng u nd u l a ti ons ,

u nl e s s the g enera l sh a p e o f th e b ody is m a in t a i ned E xt erna l
.
m u sc u l ar fib res u nopp os ed by a n i nt erna l res ist a nt m a ss ,
w ou ld c a u s e c oll a p s e of the b ody T o m eet t h e requ i re

.
m ents th ere m u s t be a m ea n s of m a i nt a i ni ng l ongit u di na l

rigidity with ou t p r ev enti ng b en d s f rom sid e t o s id e ; a nd s u ch
a m ea n s i s p res en t e d by a s t ru ct u re i niti a t ed a s d e s c r ib ed .
I n b ri ef wh eth er we h a v e or h a v e not th e a ct u a l c a u se we
, ,
“
h a v e h ere a t a ny ra t e a t ru e c a u s e T h ou gh th ere a r e
di ffi cu lti es i n t ra ci n g ou t the p roces s i n a d efi ni te wa y it ,
m a y a t l ea s t be s a id th a t th e m ech a ni ca l g enes is o f thi s r u di
m ent a ry v ert eb ra t e a xis i s qu it e c onceiv a bl e A n d ev en t he

.
di ffi cu lti es m a y I thi nk be m ore fu lly m et th a n a t fi rs t

, ,
s ight s ee ms p os s ibl e .
Wh a t i s t o be s a id of the oth er l ea di ng t ra it whi ch the

simpl es t v ert eb ra t e a n im a l ha s i n comm on with a l l high er
v ert eb r a t e a nim a l s— th e s egm ent a ti on of i ts l a t era l mu s cu l a r
m a s s es ? I s this too expli ca bl e on the m ech a n ic a l hyp o
, ,
th esi s ? H a v e we i n the a lt ern a ti ng t ra ns v ers e st ra i ns a

, ,
c au s e for the f a ct th a t whil e the ru dim ent a ry v ert eb ra t e a xi s

T HE S H A P E S OF V E RT E BRA TE S K E L E T ONS .
21 5
i s w ith ou t a ny divi s i ons th ere a re d efi nit e divi s i ons of the

,
s u b s t a nc e form i n g t h e a n im a l s s id es ? A
’
I thi nk w e h a v e .
gl a n ce a t the di s t ribu t i on of f orces u nd er th e t ra n s v erse

s t ra i n,
a s r ep res ent e d i n t h e for eg oi ng di a g r a m s w ill s h ow ,
how m u c h m ore s ev ere i s th e s t r a i n on t h e ou t er p a rts th a n

o n t h e i nner p a rt s ; a n d h ow c on s equ ently a n y m o difi c a t i on s
, ,
o f s t r u ct ure ev ent u a lly n ec ess it a t ed w ill a ri s e p eriph era lly

,
b efore t h ey a ri s e c ent ra lly T he p ercep t i on of thi s m a y be

.
en f or ced by a s i m pl e exp er im ent T a k e a s ti c k o f s ea li ng

.
wa x a n d wa r m it s l ow ly a nd m o d era t ely be f ore th e fi re s o a s ,
t o giv e it a littl e fl exibili t y T h en b en d it gently u ntil it i s

.
cu rv ed in t o a s em i ci rcl e -
On th e con v ex s u r fa ce s m a ll
.
c ra ck s w ill be see n a nd on th e c onc a v e su r f a ce w ri nkl es ;

,
w hil e b et ween th e t w o t he s u b st a n ce rem a i n s u n di s t orte d :

I f th e b en d be rev ers ed a nd re rev ersed ti m e a fter t i m e -
, .
th es e cr a ck s a nd wri nkl es will b ecom e fi s su res whi c h g ra d u

a lly d ee p en .B u t n ow i f c h a n ges of thi s cl a ss ent a il e d by
, ,
a lt ern a ti n g t r a n s v ers e s t ra i ns c om m en c e s u p er fici a lly a s t h ey

, ,
m an i fes tly m u s t ; th ere a ri s e the fu rt h er qu es ti ons— Wh a t

will be the s p eci a l m odifi ca t i ons p ro d u ced u nd er th es e s p eci a l
c onditi ons ? a nd th rou gh w h a t s t a ges w ill th ese m odi fi ca
ti ons p rogr es s ? E v ery one h a s lit era lly a t h a n d a n ex a m pl e
o f t h e w a y i n w hi c h a fl exibl e e xt ern a l l a y er th a t i s n ow
ext en d ed a n d n ow c o m p resse d by th e b en di n g o f t h e m a s s it
,
c o v er s,
becom es c rea sed ; a n d a gl a nce a t the p a l m s a nd th e
fi ngers will s h ow th a t the crea ses a re nea r one a n oth er
wh er e th e s ki n i s thi n a n d fa r a p a rt w h ere th e s ki n i s thi c k
,
.
B etw een thi s f a m ili a r c a se a nd th e c a se o f the r hi n o cero s

hid e i n whi c h th ere a re bu t a few l a rge fold s v a r i ou s gra d a
, ,
ti on s m a y be t ra ced N ow the lik e mu s t h a p pen w i th the

.
i ncr ea s i n g l a y ers of cont r a ctil e fib res f or mi ng t he s id es of

the m u s c u l a r t u nic i n su ch a typ e a s th a t su pp osed Th e .
b en di n g s w ill p r odu ce i n th em s m a ll w ri nkl es w hil e th ey a re

thi n bu t m ore d ecid ed a n d c om p a r a tiv ely di s t a n t fis sures
,
a s th ey bec om e thick . Fig 2 8 9 w hich i s a h ori zont a l l ongi

.
,
tu di n a l s ec ti on sh ows h ow th es e thi c k en in g l a y er s w i ll
,
21 6 MORP HOL OGI CA L D E V E L OP ME NT .
a dj u s t th em s elv es on th e con v e x a nd th e c onca ve su r fa c es ,
s u pp os i n g th e fib re s o f w hi c h t h ey a re com p o s e d t o be ob
liq ue a s th ei r fu n cti on requ i res ; a nd it

,
i s n ot di ffi cu lt t o s e e th a t w h e n on c e
d efinit e divi s i on s h a v e b een est a bli s h ed,
th ey will a dv a n ce i nwa r d s a s the l a y ers

d ev el op ; a n d w ill s o p rod u ce a s eri es of
m l
,
m u s cu l a r b u n dl es H ere t h en we h a v e
.
s om ethi n g l i k e th e m yocom m a ta [ or m y ot om es a s n ow c a ll ed ]
whi ch a re t ra cea bl e i n the A m p hi oxus a nd a re cons picu ou s

,
i n a l l s u p eri or fi s h es .
§ 2 5 6 T hese a re highly s p ec ul a tiv e concepti on s I h a v e

. .
v ent u r ed t o p res ent th em with th e vi ew o f i m plyi ng th a t

th e hyp oth es i s of th e m ec h a n i ca l g en es i s o f v ert eb ra t e st r u e
t u re i s not wh olly a t f a u lt wh en a ppli ed t o t he m os t ru di
m ent a ry v ert eb ra t e a ni m a l L es t it s h ou ld be a ll eged th a t
.
t h e qu es ti on is b egged if w e set ou t with a typ e whi ch l i k e ,
the A m p hi ox as a l rea dy di s pl a y s segm ent a ti on th rou gh ou t

-
i ts m u s cu l a r sy s t em it s ee m ed need fu l t o i n dic a t e c onc ei v

,
a bl e m o d es i n w hi c h th er e m a y h a v e b een m ec h a n i ca lly r o
p
du ced th os e l ea di ng t ra it s th a t di s ti ngu i s h th e A m p hi ox us .
A l l I i nt en d t o s u gg es t i s th a t m ech a ni c a l a c ti ons h a v e b een

a t w or k a n d th a t p rob a bly th ey h a v e op era t e d i n th e m a nner
,
a ll e ge d : s o p r ep a r i n g th e w a y for n a t u r a l sel ecti on

.
B u t n ow l et u s ret u rn t o the regi on of es t a bli s h ed f a c t a n d

,
c on s id er w h eth er su ch a c ti on s a n d rea cti on s a s w e a ct u a lly
wit n es s a r e a d equ a t e c a u s es of th ose ob ser v e d di fferenti a ti on s

,
a n d i n t eg r a ti on s whi c h disti ng ui s h the m or e d ev el op e d v e rt e

-
b ra t e a ni m a l s L et u s see wh eth er the th eory o f m ech a n i ca l

.
g enes i s a ff or d s u s a d ed u ctiv e i nt erp ret a ti on of the i ndu ctiv e

g enera liz a ti ons .
B efore p r oc eed i n g w e m u s t n ot e a p roce ss o f f u ncti on a l

,
a d a pt a ti on whi ch h er e co op er a t es w i t h n a t u r a l sel ec t i on
-
.
I refer t o the u su a l f orm a ti on of d en s er ti s su es a t th o s e

p a rt s of a n orga nism which a re exp os ed t o the g rea t es t
TH E S H A P E S OF V E RTE BRA T E S K E L E T ONS .
st ra i n s— e ither c omp ress i on s or t en s i on s I n s t a nce s o f h a rd .

~
eni ng u n d e r com p r es s i on a r e m a d e fa m ili a r t o u s by t h e
s ki n. We h a v e th e g enera l c ont ra s t b et ween the s oft s ki n

c o v er i n g t h e b o dy a t l a r g e a n d t h e i n d u ra t e d s ki n cov eri ng
,
th e i nn er s u r fa ces of t he h a nd s a nd t he s ol es of t h e feet .
We h a v e th e fa c t th a t ev en wi t hi n th ese a rea s th e p a r ts
on w hi c h t h e p res s u re i s h a bit u a lly g rea t es t h a v e th e s ki n
a l wa y s t hi ck es t ; a n d th a t i n ea c h p er s on s p ec i a l p oi nt s
exp o se d t o s p eci a l p res su res b ec o m e s p eci a lly d en se— o ften
a s d ens e a s h or n F u r th er we h a ve th e c onv er s e fa c t th a t
.
,
the s ki n o f littl e u s ed h a n d s b ec om es a b n or m a lly thi n— ev en

-
l os i n g i n pl a ces th a t r ibbe d s t ru ct u re w hi c h di s ti ng ui s h es
, ,
s ki n s u bj ec t t o r ou gh u s a g e Of i nc r ea se d d en s ity di rec tly

.
f oll owi n g i ncrea se d t ens i on th e s k el et on s wh et h er of m en

, ,
o r a n im a l s fu rn i s h a b u nd a n t evid enc e
,
A n a t omi s t s ea s ily
.
di s c ri m i na t e b et ween th e b ones of a s t ron g m a n a n d th os e o f

a we a k m a n by t he gre a t er d ev el op m ent o f th ose ridg es a nd
,
c res t s t o whi ch t he m u scl es a re a tt a ch ed ; and na t u ra li s t s on ,
c om p a r i ng the rem a i ns of d om es ti ca t ed a ni m a l s with t h ose

of wild a ni m a l s o f t h e sa m e s p ec i es fi n d ki n d re d diff erence s
,
.
T he fi r s t o f th es e fa ct s s h ows u nmi s t a k a bly th e im m edi a t e

eff ec t o f fu n cti on on s t ru c t u r e a n d by obvi ou s a lli a n ce w i t h
,
it th e s econd m a y be h eld t o do th e s a m e : b o th i m plyi ng

th a t the d ep os it of d ense s u b s t a n ce ca p a bl e of grea t res i s t
a n ce,
c ons t a n tly t a k es pl a c e a t p o int s wh ere th e t ensi on is
e x ces s iv e .
T a ki ng i nt o a ccou n t th en thi s a d a ptiv e p r oces s con

, , ,
ti nu a l l y a id e d bv t he s u r viv a l of i n divid u a l s i n whi c h i t

h a s t a k en pl ac e m os t r a pidly w e m a y e xp ec t on t r a ci ng u p
, ,
th e ev ol u ti on o f th e v ert eb ra t e a xis t o fi nd th a t a s th e m u s
,
cu l a r p ower b ec om e s gre a t er t h e re a r i s e l a r g er a n d h a r d er
m a ss es o f ti s su e s ervi n g the mu s cl es a s p oi n ts d a ppu i ; a n d

’
th a t these a r i s e fi rs t i n th ose pl a ces wh ere th e s t ra i ns a r e

grea t est N ow thi s i s j u s t wh a t w e d o fi nd T he m yocom
. .
m a ta a re s o pl a ce d th a t th ei r a c ti on s a re lik ely t o a ffe c t fi rs t
th a t u pp er c oa t of the not och or d wh ere th ere a re f ou nd ,

21 8 MORP HOL OGI CA L D E VE L OP ME NT .
qu a d ra t e m a ss es of s om ewh a t d ens er ti s su e whi ch s eem ,

”
f a i ntly t o rep res en t neu r a l s pi nes ev en i n the A m p hi ox us
, .
I t i s by th e d ev el opm ent of th e n eu r a l s pi nes a nd a ft er th e m ,
o f t h e h aem a l spi nes

. th a t the s egment s of th e v erteb ra l
,
c ol u m n a r e fi rs t m a r k ed ou t ; a nd u n d er t h e i n crea s i n g s t r a i ns
o f m ore d ev el op ed m y ocom m a ta
-
it i s j u s t th es e p eriph era l
,
a pp en d a g e s of t h e v er t eb r a l s egm ent s th a t m u s t be m o s t
s u bj ec t t o t h e f or c es w hi c h c a u s e t h e for m a t i on o f d ens er
ti s su e . I t f oll ows f rom th e m ech a n ic a l hyp othes i s th a t a s

t he m u s cu l a r s egm ent a ti on m u s t b egi n ext ern a lly a nd p r o
g ress i nwa rds s o too m u s t the v ert eb ra l s egment a ti on
, , , .
B es id es thu s findi ng rea s on for t he f a c t th a t i n fish es w ith

wh olly c a rt il a gi nou s s k el et ons th e v ert eb ra l s egm ent s a re
,
i ndi ca ted by th ese p roces s es whil e yet th e n ot och or d is

,
u n s egm en t ed ; w e fi nd a lik e rea s on for t h e fa ct th a t th e
t ra n s i ti on f rom th e l ess d en se c a rtil a gi n ou s s k el et on t o the

-
m ore d ens e os seou s sk el et on p u r su es a p a ra ll el c ou rse I n

-
, .
th e e xi s t i n g L epi dos i r en whi ch by u n iti n g cert a i n pi s cin e a nd

,
a mphibi a n ch a ra c t er s b e t r a y s i t s c l o se a lli a nce w ith p r i m itiv e
typ es the a xi a l p a rt o f the v ert eb ra l col u mn i s u nos s ifi ed

, ,
whil e th ere is oss ifi ca ti on o f th e p er iph era l p a rt s S imil a rly .
w ith nu m er ou s g enera of fi s h es cl a s se d a s p a l azoz oi c T he .
fo s s il r em a i ns o f the m s h ow th a t whil e th e n eu ra l a nd h aem a l

s pi n e s c on s i s t ed o f b one th e cent ra l p a rt s o f th e v e rt eb rae
,
w ere n ot b on y I t m a y in s om e ca s es be n ot ed t oo b oth i n
.
, ,
e xt a nt a n d i n f os s il for m s th a t whil e t h e o s s ifi ca ti on i s com

,
l et e a t t h e ou t er e xt remiti es o f th e s pi n es i t i s i nc om pl e t e
p
a t t h ei r i nn er e xt rem iti es— th u s simil a rly implyi n g c ent ri
p et a l d ev el opm ent .
§25 7 . A ft er
th ese e xpl a n a ti ons th e p r oces s of ev ent u a l
s egm ent a ti on i n th e spina l a xis it s el f will be rea dily u nd er ,
s t ood. T he origi n a l c a rt il a gi nou s rod h a s t o m a i nt a in l ongi

t u di na l ri gidity whil e p ermitti n g l a t era l fl exi on A s fa s t a s .
it be c om es d efi nit ely m a rk e d ou t it will b egi n t o c onc ent ra t e

,
withi n it s elf a gr ea t p a rt of th ose p res su res a nd t ens i ons

T HE S H A P E S OF V E RTE BRA T E S K E L E T ONS .
c au e s d by t rans v ers e s t ra i ns A s a l rea dy s a id it m u s t be

.
,
a c t ed u p on m u c h i n th e s a m e m a nn er a s a bow th ou gh it is ,
be nt by f orces a cti ng i n a m o re i n di rec t wa y a nd like a bow ,
it mu s t a t eac h b en d h a v e t he s u b s t a nce of i t s convex sid e

, ,
ext en d e d a n d t h e su bst a nc e o f its c onc a v e s id e c o mp r es s ed .
S o l on g a s t he v ert eb ra t e a nim a l i s s m a ll or i nert s u ch a ,
c a rtil a gi nou s rod m a y h a v e s uffic i ent st r ength t o with s t a nd

th e mu s c u l a r s t ra i ns ; bu t oth er thi n g s equ a l t he ev ol u ti on
, ,
o f a n a ni m a l th a t i s l a rge o r a ctiv e o r b oth i m pli es m u s

, , ,
cu l a r st r a i n s w hi c h m u s t t en d t o c a u s e m o di fic a ti on i n s u ch a
c artil a gi nou s r od T he r es u lts of g rea t er b u lk a nd of grea t er

.
viv a ci t y m a y be b es t d ea lt wi t h s ep a ra t ely A s th e .
a nim a l in c r ea s es i n siz e t h e r od w ill g row b oth l on ger a n d

,
thi ck er On l ooki ng b a ck a t th e di a g ra m s of f or c es ca u s ed
.
by t ra n s v erse s t ra i n s it will be s een th a t a s the rod g row s

,
thi ck er i ts ou t er p a rts m u s t be e xp os ed t o m ore s ev ere ten

,
si on s a nd p ress u res if the d eg ree o f b en d is th e s a m e I t i s .
d ou btl ess t ru e th a t wh en the fi sh a dv a nci ng by l a t era l ,
u n d u l a ti on s b ecom es l ong er the cu rv a t u re a s s u m ed by the

, ,
b ody a t ea ch m ov em ent b ecom es l es s ; a n d th a t f rom this

c a u s e t he ou t er p a rt s of the n ot och ord a re oth er thi ngs ,
equ a l l es s s t ra i n ed — th e tw o c h a ng es th u s p a rti a lly neu t ra li

,
z i n g on e a n oth er B u t oth er thi ng s a re n ot equ a l

. F or .
Whil e s u pp os i n g th e s h a p e o f the b ody t o rem a i n c ons t a nt

, ,
th e f orc e e x ert ed i n m ovi ng t he b ody i nc rea s es a s th e c u b e s

of i t s dim en s i o n s th e s ecti ona l a rea o f t he n o t o ch or d on
, ,
w hi ch f a ll th e rea cti on s of thi s ex ert e d f or ce i ncr ea ses on ly ,
a s t he s qu a r es of th e dim en s i on s : wh en ce res u l t s a g rea t er
s t r es s u p on i t s s u b s t a nce T hi s h ow ev er w ill n ot be v ery

.
, ,
d eci ded wh ere th ere i s no c on s id era bl e a ctivity I t i s cl ea r .
th a t a u gm en ti ng b u lk t a k en a l one i nv o lv es bu t a m od er a te
, ,
res id u a r y i n c rea se o f s t ra i n on e a c h p o rti on o f t h e n ot och or d ;
a nd t hi s i s p r ob a bly t he rea s on wh it i s p os s ibl e for a l a rg e

y
s l u g i s h fi s h lik e t h e S t u rg e on t o ret a i n the n ot oc h ord a l
g ,
s t ru ct u r e . Bu t n ow p a ss i ng t o th e effect s of g re a t er
,
a c tivity a li k e dy n a mi c a l i nqu i ry a t on c e sh ows u s h ow r a pid

,
l y th e vi ol ence of the a cti ons a nd rea cti ons ri ses a s the m ove
m ent s b ec om e m ore viva ci ou s I n the fi rs t pl a ce the res i s t
.
,
a n ce o f a m edi u m s u c h a s w a t er i n c rea s es a s t h e s q u a r e o f
t he v el o c ity of th e b ody m ovi n g t h rou gh it ; s o t h a t t o m a i n

ta i n d ou bl e t he s p eed a fi sh h a s t o e xp end f ou r t i m es t h e
,
ener gy . B u t t h e fi sh h a s t o do m ore th a n t h i s— i t h a s t o
i ni tia te thi s s p ee d o r t o i m p res s on i t s m a s s t h e forc e i m pli e d
,
by thi s sp ee d N ow the vi s vi va of a m ovi ng b ody v a ri es a s

.
t he s qu a re of th e v el oc ity ; w h enc e it f oll ow s th a t t he en ergy

r equ i re d t o g ener a t e th a t vi s vi va i s m ea s u re d by th e s q u a r e
of th e v el oc ity it p ro d u c es .C ons equ ently did the fi sh pu t

,
it s elf i n m oti on i ns ta n ta n eou s l y the exp enditu re of energy i n

,
gen era t i ng i ts own via vi va a nd s i m u lt a neou s ly ov erc om i ng

t he res i s t a nce o f th e w a t er w ou ld v a ry a s th e f ou rth p ow er
,
o f th e v e l o c ity . B ut the fi sh ca nn ot pu t it self i n m oti on

’
i ns t a nt a neou s ly— i t m u s t do i t by i n crem ent s ; a n d thu s i t

resu lt s t h a t th e a m ou n t s o f t h e fo r c es e xp end e d t o giv e its e l f
di fi eren t v e l ociti es m u s t be rep res en t ed by s o m e s eri es o f

nu m b ers fa lli n g b et w een th e s qu a res a n d t he f ou rth p owers
o f th os e v el o citi es . Were the i nc rement s s l ow ly a ccu m u l a ted ,
the ra t i os of i ncrea s i n g eff ort w ou ld bu t littl e ex cee d th e ra ti os

o f t h e s qu a r es ; bu t w h oev er o b serv es th e s u dd en con v u l s iv e
,
a cti on w i th w hi c h a n a l a r m e d fi s h d a rt s ou t o f a s h a ll ow i nt o
d eep wa t er w ill see th a t th e v el ocity i s ra pidly genera t ed

, ,
a n d t h a t th er e for e t h e ra ti o s o f i nc r ea s i ng ef fort p r ob a bly

e x c ee d t h e r a ti os of t h e squ a res v ery c on s id era bly A t a ny
.
ra t e i t w ill be c l ea r th a t t h e e ffort s m a d e by fi s h es i n ru s hi n g
u p on p re y o r es c a pi n g en em i es ( a n d i t i s th es e ext r em e effo rt s
which h ere con c ern u s ) m u s t a s fi s h es b ec o m e m ore a ctiv e

, ,
ra pidly e x a lt th e s t r a i ns t o be b or ne by th e i r m o t or or g a n s ;
a n d th a t o f th es e s t ra i n s ,
th os e w hi ch fa ll u p on th e n ot o
c h o r d m u s t be e x a lt e d i n p r o p orti on t o th e res t T h u s t he
.
d ev el op m ent o f l ocom otiv e p ower which s u rv iv a l of the

,
fitt est m u s t t en d i n m os t c a s es t o f a v ou r i n v olv es su ch i n

,
c r e a s e o f s t r es s on t h e p rimitiv e c a rtil a gi n ou s r od a s wi ll
t end oth er thi n g s equ a l t o c a u s e i ts m odific a ti on

, ,
.
THE S HA P E S OF V E RT E BRA T E S K E L E T ONS .
221
Wh a t t i ts m odifi ca ti on be ? Con s id eri ng the com pli

'
m us
c a ti on o f t h e i n fl u en ce s a t w o rk c ons pi ri ng a s a b ove i ndi

, ,
c a t ed i n v a ri ou s w a y s a n d d eg r ee s w e c a nn o t exp ec t t o d o
, ,
m ore t h a n f orm a n id ea o f i ts a v era g e ch a ra ct er The n a t u r e .
o f t h e ch a ng es w hi c h t h e n ot och or d i s lik ely t o u n d erg o w h ere ,
g rea t er b u lk i s a ccomp a ni ed by higher a ctivity is ru d ely ,
i n dica t ed by Figs 2 9 1 29 2 a nd 2 9 3
.
, ,
T he s u cc es s iv ely
.
m m m r:
thi ck er li nes rep res ent th e s u ccessiv ely g rea t er s t ra i ns t o

w h ich th e ou t er l a yers of ti s su e a re exp ose d ; a nd the wid en
i ng i nt er s p a c es r ep r es ent th e g r ea t er e xt ens i on s w hi c h th ey
-
h a v e t o b ea r wh en th ey b ec om e c onvex or el s e the g rea t er

,
ga p s th a t m u s t be f orm ed i n th em .H a d th es e ou t er l a y er s
t o u n d er g o ext en s i on on ly a s on th e c onv e x s id e c ontin u e d
, ,
na t u ra l s el ecti on m ight resu lt i n th e f or m a ti on o f a t i s s u e
e l a s tic en ou gh t o a d m it o f th e req u i s it e s t re t c hi n g Bu t a t .
e a c h a l tern a t e b en d th es e ou t er l a y er s ,
b ec omi ng c on c a v e ,
a r e s u bj ec t t o i nc re a s e d c om p r es s i on— a c om p r es s i on w hi ch
they c a nn ot with s t a nd if th ey h a ve b ecom e s i m ply m ore

ext ens ibl e . T o w ith s t a n d thi s g rea t er c om p res s i on th ey m u s t
b ecom e h a r d er a s well a s m ore ext en s ibl e H ow a re th es e .
t w o requ i rem ent s t o be reconcil ed ? I f a s f a c t s w a rra nt

,
u s i n s u pp o s i n g a for m a ti on o f d en s er su b s t a n c e occu r s a t
,
t h os e p a rt s of th e n o t och or d wh ere t he s t r a i n i s g rea t e s t ;

it i s cl ea r th a t thi s f or m a ti on ca nnot s o g o on a s t o p r od u ce
a c o n ti nu ou s m a s s : th e p erp et u a l fl exi on s m u s t p r ev en t thi s .
I f m a t t er t h a t will n ot yi eld a t ea ch b en d i s d e p os it ed whil e

,
th e b endi ngs a re c on ti nu a lly t a ki n g pl a c e t he b en di ng s w ill

,
m a i n t a i n c er t a i n pl a c es o f di s c onti nu ity i n th e d ep os it
pl a c es a t whi ch th e wh ol e of th e st ret chi n g c on sequ ent on

ea c h b en d w ill be c on c en t r a t ed . A n d th u s the t en d en c y will
be t o for m segm ents of h a r d ti s s u e c a p a bl e o f g rea t resi s t a n c e
to c omp ressi on with i nt erv a l s fill ed by el a st ic t is su e c a p a bl e

,
o f g rea t res i s t a nc e t o e xt en si on— a v er t eb r a l c ol u m n .
A n d n ow ob s erv e h ow t he p rog res s of oss ific a t i on i s j u s t

s u c h a s c on forms t o this vi ew T h a t c en t r ip et a l d e v el op
.
m ent of s egm ent s whi ch h old s of th e v erteb ra t e a n i m a l a s a

w h ol e a s i f c a u s ed by t ra n s v ers e s t r a i n s it ou gh t t o d o a n d
, , , ,
w hi ch h old s of t he v ert eb ra l c ol u mn a s a w h ol e a s it ou ght ,
t o do h old s a l s o of th e c ent r a l a xi s
, On t he m ech a ni ca l
.
hyp oth es i s the ou t er s u r f a ce of th e n ot och ord s h ou ld be the

,
fi rs t p a rt t o u nd erg o i ndu r a ti on a n d th a t divi s i on i nt o s eg

,
m en t s w hi ch m u s t a cc om p a ny i n du ra ti on A n d a c c or di ngly .
,
i n a v er t eb ra l c ol u m n of whi ch th e a xi s i s b egi nni ng t o

oss i fy th e c ent ru m s c on s i s t of b ony r i ng s i ncl os i n g a still
,
c on ti nu ou s r od of c a r til a ge .
§25 8 . S u nd r y
th er genera l f a ct s di s cl os ed by the c om
o .
a r a t i ve m o r ph ol o gy of t h e Ver tebra ta s u pply f u th er co n

p r ,
fi rm a ti on L et u s t a k e fi rs t the s t ru ct u re o f th e s ku ll
. .
On c on s id eri ng th e a rr a ngem ent o f th e m u sc u l a r fl a k es or ,
m yoc om m a ta i n a ny or di n a ry fi sh w hi c h c om es t o t a bl e— a n
,
a r ra ng em en t a l rea dy sk et ch ed ou t i n t h e A m p hi oxu s— i t i s
n ot di f fi cu lt t o s ee th a t th a t p or ti on of th e b ody ou t of whi c h
th e h ea d of th e v ert eb ra t e a ni m a l b ecom es d ev el op ed i s a ,
p orti on w hi ch ca nnot s u bj ec t it s el f t o b en din gs i n the s a m e

d egree a s th e res t of th e b ody T he m u s c l es d ev el op ed th ere
.
m u s t be c om p a ra tiv el y s h ort a nd m u ch i n t erfered w ith by

,
th e p r e exi s ti ng or ifi c es
- H en ce th e c eph a lic p a rt will n ot
.
p a rt a k e i n a ny c on sid era bl e d eg ree o f the l a t era l u n dul a

ti on s ; a nd th ere will not t en d to a ri s e i n it a ny s u ch di s tinc t
s eg m en ta ti on a s a ri s es e l s ewh er e We h a v e h ere th en a n
.
, ,
e xp l a n a t i on o f th e fa c t th a t f ro m t h e b egi nn i n g th e d ev el op
,
m ent of th e h ea d foll ows a c ou r se u nlik e th a t of the spi na l

c ol u mn ; a n d o f the f a ct th a t th e s eg m ent a ti on s o fa r a s it ,
c a n be t r a ced i n the h ea d i s m os t rea dily t o be t r a c e d i n the

,
oc cipit a l r egi on a nd b ec om es l o s t i n th e r egi on o f t h e f a ce .
F or i f as we h a v e s een the s egm ent a ti on c ons equ ent on

, ,
TH E S H A P E S OF VE RT E BRA TE S K E L E T ONS .
m ec h a n i ca l a cti ons a nd rea cti ons m u s t p r og res s f rom w ith ou t

inwa rd s a ffecti ng l a s t o f a l l the a xi s ; a nd if a s we h a v e
, ,
s ee n
, th e regi on o f t h e h ea d i s s o ci rc u m s t a n c ed th a t t he
c a u ses of segm ent a ti on a ct bu t feebly even on i t s p er ip h ery ;
then th a t t erm i na l p orti on of the p rimitiv e n ot och or d w hi ch
i s i nc l u d e d i n t he h ea d h a vi ng t o u n d er g o n o l a t era l bend
,
i ng s m a y oss i fy w i th ou t divi s i on i nt o s egmen ts

,
.
Of othe r i nc id ent a l evid ences s u ppli e d by c omp a ra tiv e

m orph ol ogy l et m e next refer t o the s u p ernu mera ry b ones
, ,
which the th eor y o f Goeth e a n d Ok en a s el a b ora t ed by P rof .
Ow en h a s t o g et ri d of by g ra t u it ou s s u pp os iti ons
, ~
I n m a ny .
fi sh es for ex a mpl e th er e a re w h a t h a ve b een c a ll ed i nt er

, ,
neu ra l s pi n es a n d i n t er h aem a l s pi n es -
T h es e c a nno t by a ny
.
i ngenu ity be a ffili a t ed u p on th e a rch etyp a l v ert eb ra a nd ,
they a re th erefore a rbit ra rily r e j ect ed a s b on es b el ongi ng t o

the exo s k el et on ; th ou gh i n s h a p e a n d t ext u re th ey a re
-
s i m il a r t o t h e s pi nes b e t w een w hi c h th ey a re pl a ce d On th e .
hyp othes i s of ev olu ti on h owev er th es e a dditi on a l b ones a re

, ,
a ccou n t ed for a s a r i s i ng u n d er a c ti ons lik e th o s e th a t g a v e
o rigi n t o th e b ones a dj a cent t o th em A nd s i m il a rly with

.
s u ch b on es a s th os e c a lled s es a m oid ; t ogeth er with other s

t oo nu m er ou s to n a m e .
Of t he
foreg oi n g s y nth es is i s t o be t a k en
c ou r s e
si m ply a s a n a d u m b ra ti on of t he p rocess by which th e v ert e

b r a t e s t ru ct u r e m a y h a v e a r i s en th r ou gh the c on ti nu ed
a c ti on s o f k n ow n a g en c i es T h e m otiv e for a tt empti n g it h a s
.
b een tw o fold H a vi ng a s b efore s a id giv en r ea s on s for c on

-
.
, ,
c l u di n g th a t t h e s eg m ent s of a v ert eb ra t e a n i m a l a re n ot
h om ol og ou s i n the s a m e s en s e a s a re th os e of a n a nnu l os e
a ni m a l it seem ed n eed fu l t o do s om ethi ng t ow a rd s s h ow i n g
,
h ow t h ey a r e o th erw i s e t o be a ccou nt ed for ; a n d h a vi n g h ere ,
for ou r g enera l su bj ec t th e lik eness es a n d differen ces a m on g

,
the p a r t s o f o rg a ni s m s a s d et erm i ned by i n c id ent f orces it

, ,
s eem e d ou t o f t h e q u es ti on t o p a s s by th e p robl em p res ent ed
by th e v ert eb ra t e s k el et on .
L ea vi ng ou t a l l th a t i s hyp oth eti ca l the gen er a l a r gu m ent

,
m a y be b ri e fly p res en t e d th u s — T h e ev o l u ti on from t he
simpl es t k nown v ert eb r a t e a n im a l of a p owerfu l a nd a ctiv e
v erteb ra te a nim a l impli es the d ev el op m en t of a s tr onger
,
i nt erna l fu l cru m .T he i nt erna l f ul cru m c a nn ot be m a d e

s t r on g er w ith ou t b ec o mi n g m or e d en s e A nd i t c a nn ot be
.
c om e m or e d en s e whil e r et a i n i ng i t s l a t era l fl exibili t y w i t h ,
ou t b ec o mi n g divid e d i nt o s egm ent s Fu rth er i n c on form ity

.
,
with the g enera l p ri ncipl es thu s fa r t ra ced th ese segm ent s ,
m u s t be a lik e i n p rop orti on a s the forc es t o whi ch th ey a re

e xp osed a r e a lik e a n d u n lik e i n p r op orti on a s t h es e f orc es
,
a r e u n lik e ; a n d s o th ere n ec es s a rily r es u lt s th a t u n ity i n
v a ri ety by w hi c h th e v er t eb ra l c ol u m n i s fro m the be gi nni n g

c h a ra c t er i ze d . Onc e m ore w e see th a t th e expl a n a ti on ex
,
t en d s t o th ose i nnu m era bl e a n d m ore m a rk e d div e rgen c es

f rom h om og en eity whi ch v ert eb rae u nd erg o i n the v a ri ou s
,
high er a ni m a l s T hu s the p rod u cti on of v ert eb rae the pr o

.
, ,
d u cti on of lik enes s es a m ong v ert eb rae the p rod u cti on o f u n

,
lik enes ses a m ong v ert eb rae a nd th e p r od u cti on of u nlik e

,
n es s es a m on g v ert eb ra l colu m ns a re i nt erp r et a bl e a s p a rt s o f

,
o n e gen er a l p r oc es s a n d a s h a r m oni z i ng w ith o n e genera l

,
p ri ncipl e .
Wh eth er su ffi ci en t or i nsuffici ent the expl a na ti on h ere

,
”
giv en a s s ig ns ca u s es of k nown ki n d s p rod u ci ng eff ect s su c h

a s th ey a r e k now n t o p r od u ce I t d oes n ot a s a s olu ti on o f
.
,
o n e m y s t e ry ,
of f er a n oth er my s t ery of which no s ol u ti on i s
t o be a s k ed . I t d oes n ot a ll ege a P l a t on ic i Sea or fi ctiti ou s ’
,
en ti ty w hi ch expl a i ns th e v ert eb r a t e s k el et on by a b s orbi n g

,
i n t o it s el f a l l t he i nexpli ca bility On the cont rary it a s su m es

.
,
n othi n g b ey on d a g en ci es by which s t ru c t u res i n g enera l a re
m ou ld ed— a gen c i es by which th es e p a rt i cu l a r st ru ct u res a r e ,
i nd eed not ori ou s ly m odi fi a bl el A n a s cert a i ned c a u s e of cer

,
t a i n t ra it s i n v ert eb rae an d oth er b on es it ext end s t o a l l ,
ot h er t ra it s o f v er t eb rae ; a n d a t th e s a m e ti m e a s s i m il a t es
t he m orph ol ogi ca l ph en om en a th ey p res ent t o m u ch w id er

c l a ss es o f m or ph ol ogi c a l ph enom ena .
T HE S H A P E S OF VE RT E BRA TE S KE L E TONS .
— i n th e f oreg oi ng ch a pt er i s
[ N GTE T he theory f orth
s et ,
an e al b ora ti on o f on e s u gg es t ed at th e cl os e o f a c riti cis m
f the Ver tebra te

H om
’
o f P r o f Ow en s A r che typ e
. a nd ol ogi es o
S kel eton a l rea dy referre d

,
t o i n § 2 1 0 a s h a vi ng been pu b
l i s hed i n the Medi c o C hi r urgi ca l Revi ew for Oct ob er 1 8 5 8
-
,
.
I t i s n ow rep rod u ced i n A pp endix B S i nce t h e i s s u e of t hi s .
e l a b ora t ed exp os iti on i n N o 1 5 o f my s er i a l i n D ec e m b er

,
.
,
1 8 6 5 v erific a ti on s of it h a v e from tim e t o tim e b een p u bli s h ed

,
.
I n hi s w or k T h e P ri m a ry F a c tors of Orga ni c E vol u ti on ,
P r of C op e of P hil a d elphi a w r it es
.
Mr H erb ert S p en cer h a s en d ea v ou red t o a ccou n t for the

.
o r igi n o f t h e s egm en t a t i on o f m u s c l e s i n t o m y o t om es a nd ,
t h e divi s i on o f the s h ea th of th e n ot oc h or d i n t o v ert eb ra! ,
by su pp os i ng it to be du e t o the l a tera l s wi m m i n g m ov em en t s
o f t he fi s h es w hi ch fi rs t e xhibit th es e s t ru c t u res
, With thi s .
vi ew v a ri ou s l a t er a u th ors h a v e a greed a nd I h a v e offered ,
s o m e a dditi on a l evid ence of th e s ou nd ne s s o f this p os iti on
wi th res p ect t o the vert eb ra l a xi s of B a t ra chi a a n d the .

,
o r igi n o f limb a rti cu l a ti on s I t i s t ru e th a t th e or igi n o f

.
s egm en t a ti on i n th e v ert eb r a l co l u m n o f t h e t ru e fi s h es a n d
th e B a t r a chi a t u r ns ou t t o h a v e b ee n l es s s i m pl e i n i t s p ro
c es s th a n w a s s u gge s t e d by Mr S p enc er bu t his genera l
.
,
p ri ncipl e h olds good now th a t p a l eont ol ogy h a s cl ea red u p

,
the su bj ec t ( pp 3 67 .
A n a ll u s i on i n th e f or eg oi ng ext r a ct i s m a d e by P rof C op e .
t o cert a i n ob s erv a ti ons set for th i n h i s w or k en titl ed T he

O ri gi n of the F i ttes t On pp 3 05 —6 o f it w ill be f ou n d th e
. .
foll ow i ng s ent ences

N ow a l l th e P er m i a n l a n d a ni m a ls reptil es a n d b a t ra
,
-
,
chi a n s ,
ret a i n thi s n ot och or d with t h e e l em en ts o f os seou s
v ert eb rae i n a g rea t er or l es s d eg ree of c om pl et enes s T h ere

, .
a r e s om e i n S ou th A fric a I b eli ev e i n whi ch t h e o s s ifi ca ti on

, ,
h a s com e cl ea r th rou gh the n ot och or d ; bu t th ey a r e few .
T h ere i s s om ethi ng t o be s a id a s t o t he conditi on of

the c ol u m n f r om a m ech a n i ca l s t a ndp oi n t and it i s thi s ,
th a t the c h or d a exi s t s with i ts os s eou s el em ent s di s p os ed

,
61
a b ou t it ; a nd i n the P ermi a n b a t ra ch i a ns equ a lly rel a t ed to ,
s a l a m a n d er s a nd frog s th es e os s eou s el em en t s a r e a rr a n ged i n

,
th e s h ea th or s kin o f the ch or d a ; a n d th ey a re i n th e for m of

r eg u l a r c onc a v e se gm ent s v ery mu ch lik e s u ch segm ent s a s
,
ou ca n t a k e fr o m t h e ski n o f a n ora n g e — bu t p a rt s o f a
y
c yli nd er a n d h a vi ng g r ea t er o r l es s di m ens i on s a ccor di ng t o
,
t h e g r ou p or sp ec i es .N ow th e p oi n t of div ergenc e o f t h es e
,
s eg m ent s i s on t h e s id e of t h e c o l u m n T h e con t a c t s a re
.
pl a ced on the s id e of th e c ol u m n w h ere th e segm ent s s ep a r

a t e —the u pp er seg m ent s
h
r i s i n g a nd th e l ower s egm en t s
comi n g d ownw a r d .T o the u pp er s egm ent s a re a tt a ch ed th e
a r ch es a n d th e i r a rtic ul a ti ons a n d th e l ower s egm en t s a r e
,
lik e the s egm ent s of a cyli n d er I f you t a k e a fl exibl e

.
c yli n d er a n d c o v er it w ith a m ore or l ess in fl exibl e ski n o r

,
s h ea th,
a nd b en d t h a t c yli n d er s id ewi s e you o f c ou r s e will
,
fi n d th a t the w ri n kl e s or fr a ct u res of th a t p a rt of t he s u r fa ce
will t a ke pl a ce a l ong th e li ne o f the s h ort es t c u r v e whi ch ,
i s on t he s id e ; a nd a s a m a tt er of fa ct you h a v e b rea k s
, ,
o f v ery m u ch t h e ch ara ct er o f t h e s egm ent s of th e P erm i a n
B a t ra chi a . I n th e cyli n d er b en di ng b oth w a y s o f ,
c ou r s e the sh ort es t line of cu rv e i s right a t th e c ent re of th e

s id e o f th a t c yli n d er a nd th e l on ges t cu r v e i s o f c ou r s e a t
,
t h e s u mmit a n d b a s e a n d the s h ort es t cu r v e will be the p oi n t

,
of fr a ct u re . A n d th a t i s ex a ctly wh a t I p res u m e h a s
h a pp ened i n the ca s e o f the c ons t ru cti on of the s egmen ts of
t h e s h ea th of the v ert eb ra l c ol u mn by th e l a t era l m oti on of
,
th e a n i m a l i n s w im m i ng a n d w hi ch h a s been th e a c t u a l c a u s e
,
o f th e di s p o s iti o n o f t h e o s s e ou s m a t eri a l i n i t s f orm .
T h a t i s t h e s t a t e o f th e v ert eb r a l c ol u m n o f m a ny o f th e
Vert eb r a t a of the P erm i a n p eri od .
I n hi s es sa y on T he Mech a nic a l C a u s es of the D ev el op

m en t of th e H a r d P a rt s o f the Ma mm a li a p u bli s h ed i n the
”
,
A m eri c a n J ou rna l of M orp hol ogy ( Vol I I I ) P rof C op e h a s.

,
.
c a rri ed the i nt erp reta ti on fu rt h er by s h owi n g th a t i n ki nd red

,
wa ys the genesis of a rtic u l a ti on s a nd limb b ones m ay be ex -

T HE S HA P E S OF V E RT E BRA TE S KE L E T ONS .
22 7
pl a in ed On p 1 6 3 he enu nci a t es the genera l p ri ncipl e of

. .
his i nt erp ret a ti on a s f oll ow s

“
I t c a nnot h a v e b ee n otherwi s e th a n tha t s i nce the ,
m oti ons of a nim a l s c onti nu ed d u rin g the ev olu ti on o f th ei r

h a rd p a rt s thes e h a r d p a r ts g rew i n ex a c t a d a p t a t i on t o
,
t hese m ov em ents T hu s a t t he p oi n t s of grea t es t fl exu re

.
j oin t s w ou ld be f or m ed a nd b etween th es e j oi n ts th e d ep os it
,
”
w ou ld be c on ti nu ou s .
E vid ently if os seou s st ru ct u re s a re p rod u ced by d ep osit s

of c a lc a reou s m a tt er s i n p re—e xi s ti n g c a rtil a gi n ou s st ru ct u res ,
o r o t h er s t ru c t ur es o f fl e xibl e m a t er i a l s t h e d ep os it s m u s t be
,
s o c a r ri ed on th a t w hil e d en s e res i s t a n t m a s s es a r e p ro d u c e d
thes e m u s t a d m it of s u c h free m ov em ent s a s the c rea t ure s ’
life necess it a t es a nd m u s t s o f orm a d a pt ed j oi nt s

,
.
L et it b e u n d ers t oo d h owev er th a t th e hyp oth esi s set

, ,
f orth i n th e f oreg oi n g ch a pt er a nd ex tend ed by P rof C op e .

,
whi ch s er v es t o i nt er p r et a l a rge p a rt o f th e ph eno m ena

o f o s seou s s t ru c t u res i n t h e Ver tebra ta d oes not s er v e t o
,
i nt erp ret th em a l l Whil e t he f or m a ti on of h a r d p a rt s h a s

.
been i n l a rge m ea su re in iti a t ed a nd reg ul a t ed by t ens i ons a nd

p ressu res th ere a re h a r d p a rt s th e f orm a ti on of whi ch c a nnot
,
be thu s expl a i ned T he b ones o f th e s ku ll a re t he m os t

.
obvi ou s i n s t a nc es T h es e a re a pp a rent ly re fera bl e t o n o

.
ot h er c a u s e th a n t h e s u r vi v a l o f th e fi tt es t— th e s u rviv a l o f
i ndividu a l a n im a ls i n whi ch grea t er d ens ity of the b ra i n

c o v eri n g yi eld ed b ett er p r ot ec ti on a g a i n s t ext ern a l i n j u r i es .
Wi th ou t enu m era ti n g oth er i ns t a nc es whi ch might be given ,
it will su ffi ce t o rec ogni ze th e t ru th th a t na tu ra l sel ec ti on of

fa v ou ra bl e v a ri a ti ons a n d the i nh eri t a nce of fu ncti ona lly
p rod uced ch a n ges h a v e a l l a l ong c c op era t ed : ea ch of th em
-
i n s om e c a s es a ctin g a l one bu t i n oth er c a s es b oth a cti ng

,
t ogeth er ] .
C H A P T E R X VI .
TH E I
S H A P E S or A N M A L - CE L L S .
2 60 A M O N G
. i ls a s a m ong pl a nt s the l a w s o f m or
an m a ,
ph ol og i ca l di fferenti a ti on m u s t be c on f or m ed t o by th e m or
p h o l og i c a l u n it,
s a s w ell a s by t h e l a rg er p a rt s a n d by t h e
w h ol es f or m ed of th em I t rem a i ns h ere t o p oi nt out th a t

.
t he conf or m ity i s t ra c ea bl e w h ere th e conditi ons a re s impl e .
I n t he sh a p es a s su m e d by th os e r a pidly m u ltiplyi ng c ells

-
o u t o f whi ch ea c h a ni m a l i s d ev e l op ed th er e i s a cons pi cu ou s
,
s u b or di na ti on t o th e s u rrou n di n g a c ti on s .
Fig 29 4 rep resent s the c ell u l a r em b ry onic

.
m a s s th a t a ri s es by rep ea t ed s p ont a neou s

fi ss i on s I n it we s ee h ow th e c ells or igi n
.
,
a lly s ph eri c a l ,
a re ch a n ge d by p r es su re
a g a i ns t on e a n oth er a nd a g a i n s t t he limit
i ng m em b r a ne ; a n d h ow th ei r lik enes s es
a n d u nlik enes s es a re d et er m i ned by th e lik ene s s es a n d u n
lik enesses of th e forces t o whi ch th ey a r e exp os ed T hi s f ac t .
m a y be th ou ght s c a rc ely w orth p o i nti n g ou t B u t it i s

.
w ort h p oi nti ng ou t b eca u s e w h a t i s h ere s o obvi ou s a con

,
s eq u ence o f m e ch a n i c a l a c ti on s ,
i s i n oth er c a s es a c ons e
qu ence o f a cti on s c om p o s it e i n th ei r ki nd s a nd i nv olv ed i n
th ei r di s t rib u ti on Ju s t a s th e eq u a liti es a nd i nequ a liti es of
.
dim en s i ons a m ong a ggreg a t ed cell s a re h ere ca u s e d by th e

,
e q u a liti es a nd i neq u a liti es a m on g th e i r m u t u a l p res s u res i n
different di recti ons ; s o th ou gh l es s m a nifestly the equ a liti es

, ,
THE S HA PE S OF A NIMA L CE L L S .
22 9
a nd in eq u a liti es of di m ens i ons a m on g oth er a ggrega t ed c ell s ,
a re c a u se d by t h e eq u a liti es a n d i nequ a li ti es of t h e o sm otic

,
ch emi ca l therm a l a nd ot h er f orces b es ides the m ech a ni c a l

, , ,
t o w hi c h th ei r di fferen t p os iti on s s u bj ec t th em .
261 . T hi s we
h a ll rea dily see on ob s erv i ng the ordi na ry
s
s t ru ct u r es o f li m i t in g m e mb ra n es i nt erna l a nd ext erna l

, .
I n Fig 2 9 5 i s s h own a
.
,
m u ch m a gni fied s ecti on

-
of a p a pill a f rom th e
gu m . T he c ell s of w hi c h
it i s c om p osed ori gi n a t e
i n i ts d eep er p a rt ; a nd
a re a t fi r st a pp r o xim a t ely
S ph er ic a l T h os e of th em
.
whi ch a s th ey d evel op a re th ru s t ou t wa rd s by the new

, ,
cell s th a t c onti nu a lly t a k e th ei r pl a c es h a v e th ei r sh a pes

,
gra du a lly ch a nged A S th ey g row a n d s u cc es s iv ely a dv a nce

.
t o repl a ce the s u p erfi c i a l c ell s wh en th es e e xf oli a t e th ey

, ,
bec om e exp o sed to f orces w hi c h a re m ore and m ore different

i n th e di recti on o f th e s u r fa c e f rom wh a t th ey a re in l a t era l
di recti ons ; a nd th ei r dim ens i ons gr a d u a lly a s su m e c orre
s pondi ng di ffer en c es .
A noth er s p eci es o f limiti ng m em b r a ne c a ll ed cyli nd er ,
epith e li u m i s rep r es ent ed

,
i n Fig 2 9 6 . T h ou gh i t s
.
m ode of d evel op m ent i s

s u ch a s t o ren d er th e
s h a p es o f its c ell s q u it e
u nlik e th o s e of p a v em en t
epith eli u m a s th e a b o v e d esc ribe d ki n d i s s o m eti m es c a ll e d

,
-
,
i t s c ell s eq u a lly ex emplify th e s a m e g enera l t ru th For t he .
c hi ef c ont ra st which ea c h o f th em p r es en ts is the c ont ra s t ,
b etween i ts dim ens i on a t right a n gl es t o the s u r f a c e of the

m emb ra ne a nd i ts dim ens i on p a ra ll el t o th a t s u r fa ce
,
.
I t i s needl ess for ou r p res ent p u rp o s e t o ex a min e furth er

th e e vid enc e fu rn i s h ed by H i s t ol ogy ; n or i nd eed w ou ld

, ,
fu rth er e x a mi na ti on of this evid ence be likely t o yi eld

d efinit e res u lt s I n the c a s es given a b ov e we h a v e m a rk e d
.
di fferences a m ong t he i nc id ent f orc es ; a nd th ere f ore h a v e a

ch a nce of fi n din g a s we d o fi nd rel a ti ons b et ween thes e a n d
, ,
di fferences of f orm . B ut t he c ells comp os i ng m a s s es o f

t i s s u e a re s ev era l l y s u bj ec t t o forces which a r e i n d et erm i n a t e ;
a n d th erefore th e i n t er p ret a ti on o f th ei r S h a p e s i s i m p r a c
t i ca bl e. I t mu s t s u ffi c e t o ob s erv e th a t s o fa r a s the f a ct s g o
th ey a re c ong ru ou s with the hyp othesis .
C H A P T E R X VI I .
S U M M A RY OF M ORP H OL O GI C A L D E VE L OP ME NT .
262 . TH AT f ormu l a sh ou ld be c a p a bl e of exp ressi ng

a ny
a c omm on ch a ra ct er i n th e sh a p es of thi n g s s o u n lik e a s a
t ree a n d a c ow a fl ower a nd a centip ed e i s a rem a rk a bl e

, ,
f a ct ; a nd i s a f a ct whi ch a fford s s t ron g p ri m a fa ci e evid ence

o f t ru th. F or i n p rop or ti on t o the div er s ity a n d m u ltiplicity
o f t he c a s e s t o whi ch a ny s t a t e m en t a ppli es i s t h e p ro b a bility
,
tha t it s ets f ort h the es s enti a l rel a ti on s T h os e c onnexi on s

.
whi c h r em a i n c on st a n t u nd er a l l v a ri eti es of m a nif est a ti on ,
a re m os t lik ely t o be th e c a u s a l connexi ons .
S till high er will a pp ea r the lik elih oo d of a n a ll eged l a w of

o rg a n i c f orm p osses s i n g s o g rea t a c om p reh ens iv eness w h en ,
we r em em b er th a t on t h e hyp oth es i s o f E v ol u ti on th ere mu s t ,
exi s t be t ween a l l or g a n i s m s a n d th ei r envi r onm ent s c ert a i n ,
c ong ru iti es exp r es s ibl e i n t er m s o f th ei r a c ti on s a n d rea cti on s .
The f orces be in g on this hyp oth es is th e c a u ses of th e f or m s

, , ,
it i s i nfera bl e a p ri ori th a t the f orm s m u st a dmit of genera li

, ,
za ti on i n t erms of t he for c es ; a nd h en c e s u c h a g ener a li z a ti on

,
a rr iv e d a t a
p os teri ori g a i ns th e f u r th er p rob a bility du e t o
,
fu lfilm en t of a nti cip a ti on .
N ea rer yet t o c ert a i nty s eem s th e c on cl u s i on th u s rea ch ed ,
on fi n di ng th a t it d oes bu t a ss ert i n th ei r sp eci a l m a n if est a
ti ons the l a ws of E v ol u ti on i n gener a l— the l a ws of th a t

,
u niv er s a l r e di s t r ib u ti on of m a tt er a n d m oti on w hich h old

-
t h rou gh ou t th e t ot a lity of thi ngs ,

as w ell as in ea ch of its
p a rt s.
I t will be u s efu l t o gl a n ce b a ck o v er the v a ri ou s mi n or

i n ferences a rri ve d a t a nd c on t em pl a t e th em i n th ei r ens em bl e
,
f ro m th es e high er p oi nts of vi ew .
263 . p roces s of i nteg ra ti on whi ch ev ery pl a nt di s

T ha t
pl a y s du r i ng i ts life we f ou nd rea s on t o thi n k h a s g on e on
,
d u ri ng th e life of the v eget a l ki ngd om a s a wh ol e P rot o .
pl a s m i nt o cel l s cells i nt o f oli a foli a i nt o a x es a x es i nt o

, , ,
b ra n ch ed c ombi n a ti ons— su ch i n b ri ef a re the s t a g es p a ss ed

, ,
th rou gh by ev ery s h ru b ; a nd s u ch a pp ea r t o h a v e b een the

s t a g es th rou gh w hi ch pl a n t s of s u c ces s iv ely high er ki nd s -
h a v e been ev olv ed f rom l ower ki nd s E v en a m on g cert a i n .
g rou ps of pl a nt s now exi s ti ng we fi nd a ggreg a t es of the fi rst

,
or d er p a s s i ng th r ou gh v a ri ou s g r a d a ti on s i n t o a gg reg a t e s o f
t he s econ d or d er— h ere f orm i n g s m a ll i nc oh eren t i nd efi n it e , ,
a ss em bl a g es a n d th e r e formi n g
,
l a rge d efi nit e c ohe ren t , ,
f rond s . S imil a r t ra ns iti ons a re t ra c ea bl e th rou gh whi ch

th es e i nt egra t ed a gg rega t es of th e s ec on d or der p a s s i n t o
a gg reg a t e s o f th e t hi r d or d er : i n on e s p ec i es t h e u ni ons o f
p a ren t —fron d s w ith the f ron d s th a t bu d ou t from th em b ein g ,
t emp or a ry a nd i n a noth er s p eci es su ch u ni ons b ei n g l onger

,
c onti nu e d ; u ntil i n S p eci es s till high er by a g em m a ti on

, ,
which i s h a bit u a l a nd re gu l a r th ere i s p rod u ced a defi ni tel y

,
i nt eg ra t ed a ggrega t e of the thi r d ord er— an a xi s b ea ri ng

f ron ds or l ea v e s A nd ev en be t ween this typ e a nd a typ e
.
f u rth er c omp oun d ed a li nk occu rs i n the pl a nt s which c a s t

,
of f i n the S h a p e of b u lbil s s om e o f th e y o ung a x es th ey

, ,
p rod u ce . A s a m ong pl a nts s o a m ong a ni m a ls ,

A .
lik e s p on t a neou s fi s si on of c ell s end s h ere i n s ep a ra ti on th ere ,
i n p a rti a l a gg rega ti on whil e el s ew h ere by cl os er c ombi na

, ,
ti on of th e m u ltiplyi ng u nit s th ere a ri s es a c oh erent a n d

,
t ol era bly d efi nit e i ndivid u a l o f th e s ec on d ord er By t h e .
b u ddi ng of i ndividu a ls of the s ec ond ord er there a re i n s om e ,
c a s es p rodu c ed oth er sep a ra t e i ndivid u a ls lik e th em ; i n s om e

S U MMA R Y OF MORP H OL OGICA L D E V E L OP ME NT .
23 3
c a s es t emp ora ry a gg reg a t es of s u c h lik e i ndivid u a l s a n d i n

o th er c a s e s p er m a n en t a gg r eg a t es o f th em : c erta in of w hi c h
b ecom e s o d e fin it ely i n t egra t ed tha t the i ndividu a liti es of

th ei r c omp onent m embe rs a re a lm os t l ost i n a t erti a ry i n di
vi du a l i ty .
A l ong with t hi s p r ogress ive i nteg ra ti on th ere h a s g on e on

a p r ogres s iv e di fferen ti a ti on Veget a l u n it s of w h a t ev er
.
o r d er ,
origi na lly h o m og eneou s h a v e bec o m e het erog eneou s
,
whil e th ey h a v e b ec o m e u nit e d S ph er i c a l c ell s a ggrega ti n g

.
i nt o th rea d s i nt o l a m i nae i nt o m a s ses a nd i n t o s p eci a l t i s

, , ,
s u es,
l os e th ei r s ph erici ty ; a nd i n s tea d o f rem a i ni ng a l l
a lik e a s su m e i n n u m e r a bl e u n l i k en e s s es — from u n i formity
p a s s i n t o m u lti for mi ty F ron ds c o mbi n i ng t o form a x es
.
,
s ev era lly a cqu i r e d efi n it e differen c e s be t w een th e i r a t t a ch e d
en d s a n d th e i r free en ds ; whil e th ey a l so div erg e f r om o n e
a n ot h er i n th ei r s h a p es a t di ffer en t p a r t s o f th e a x es th e y
c o m p os e A n d a x es u niti ng i nt o a gg reg a t es of a s t ill high er

.
,
o r d er b ec o m e c ont ra s t e d i n th ei r S iz es
, ,
c u r v a t u res
,
a nd t h e
a rr a n g em ent s o f th ei r a pp en d a g es . S i m i l a rly a m ong

a ni m a l s T h os e com p on en t s of th em whi c h with a cert a i n
.
,
li cen s e we cl a s s a s m orph ol ogi ca l un it s whil e l os i ng th ei r

, ,
mi nor i ndivid u a liti e s i n th e m a j or i ndivid u a li t i es for m ed of

t h em ,
g row d efin i tely un lik e a s th ey g row d efi nit ely com
bi ned . A nd w h ere the a ggreg a t es so p ro d u ce d be c om e by ,
c oa l es c en ce s e gm en t s of a ggr eg a t es of a s till high er ord er

, ,
they t oo diverge from one a n oth er i n th ei r sh a p es

, , .
The m orph o l ogi ca l differenti a ti on whi ch th u s g oes h a n d

i n ha n d with m orph ol ogic a l i nt egra ti on i s cl ea r ly wh a t the
,
p erp etu a l l v c ompli c a tin g c onditi on s w ou ld l ea d u s to a n ti c i

-
p a t e E v ery a ddi ti on of a new u nit t o a n a gg reg a t e o f s u ch

.
u nit s m u s t a ffect th e c i r cu m s t a n ces o f th e o th er u ni t s i n a l l

,
v a ri eti es of w a y s a nd d eg rees a ccordi ng t o th ei r rel a tiv e

,
p os iti ons— m u st a lt er th e di s t rib u ti on of m ech a n i ca l s t ra i ns

th rou gh ou t th e m a ss mu s t m odi fy the p roces s of nu t riti on
, ,
mu st a ffec t the rel a ti on s of neighb ou ri ng p a rt s t o su rr ou nd

i ng di ffu s e d a cti ons ; th a t i s m u s t i n iti a t e a c h a ng ed i nci
,
dence off orces t endi ng ev er to p rodu ce ch a nged st ru ct u ra l

a rr a ng em ent s .
§ 2 64 . T hi s
b roa d s t a t em ent of the c orres p on d en ce be
t ween the g enera l f a ct s of Mor ph ol ogic a l D ev el opm ent a n d
t h e p ri nc ipl es of E v ol u ti on a t l a rge m a y be re d u c ed t o ,
s t a t em ent s of a m u c h m ore s p ecifi c ki n d T he ph enom ena .
o f sy m m et ry a n d u ns ymm et ry a n d a s y m m et ry w hi c h we ,
h a ve t r a c ed ou t a m on g org a n ic form s a re d em ons t ra bly i n ,
h a rm ony w ith th os e l a ws of the re di s t rib u ti on of m a tt er a nd

-
m ot i on t o w hi c h E v ol u ti on c on forms B esid es the my ri a d .
f old illu s t r a ti on s of th e i ns t a bility of the h om ogeneou s ,
af ford ed by th ese a ggr ega t es of u n its o f ea c h or d er which , ,
a t fi rs t a lik e l a p se gr a du a lly i nt o un lik enes s ; a n d b es id es

,
th e m yr i a d fold ill u s t r a ti on s o f th e mu ltipli ca ti on o f effect s

-
,
which th ese ev er c om plic a ti ng di fferenti a ti ons exhibit t o u s ;

-
we h a v e a l s o my ri a d f old ill u s t ra ti on s o f th e d efi ni te eq u a li
-
ti es a n d i nequ a liti es of s t ru ct u res p rodu ced by d efi nit e eq u a li

,
ti es a n d i nequ a liti es of f orces .
T he p rop os iti on a r riv e d a t wh en d ea li ng with th e c a u s es

o f E v ol u ti on th a t i n th e a cti ons a nd rea cti ons o f force a nd
,
m a tt er a n u nlik enes s i n eith er of th e f a ct or s neces sit a t es a n

,
u nlik en es s i n th e effect s ; a n d th a t i n th e a b s en ce o f u nlik e
n es s i n eith er o f th e f a c t or s th e ef f ec t s mu st be a like ( Fi rs t
P ri nci p l es ,
i s a p rop os iti on which impli es a l l th es e
p a rticu l a r lik eness es a nd u nlik enes s es of p a rt s whi ch we
h a v e b een t ra ci ng F or h a ve we not e v eryw h ere s een th a t
.
t h e s t r onges t con t ra st s a re b et w een th e p a rt s th a t a re m os t

con t ra st ed i n th ei r c onditi on s ; whil e th e m o s t s imil a r p a rts
a re th o s e m os t s i m il a rly c on diti on ed ?
-
I n ev ery pl a nt the
l ea din g difference i s b et ween th e a tt a ch ed en d a nd th e f ree
e nd ; i n ev ery b ra n c h it i s th e s a m e ; i n ev ery l ea f it i s
the s a m e A n d i n ev ery pl a nt th e l ea di ng lik eness es a re

.
th ose b et ween the two S id es of the b ra nch the tw o S id es of ,
the l ea f a n d th e two s id es of th e fl ower wh ere th es e p a rt s

, ,
a re t w o sid ed i n th ei r c on diti on s ; or b et ween a l l sid es of the

-
S U MMA RY OF MORP H OL OGI CA L D E VE L OP ME NT .
23 5
b ran ch a l l s id es of the l ea f a nd a l l S id es of th e fl ower wh ere

, , ,
thes e p a rt s a re si m il a rly c ondi ti oned on a l l S id es S o t oo i s it.

, ,
with a ni m a l s whi ch m ov e a b ou t T h e m os t m a rk e d con t ra s t s

.
th ey p resent a re th ose be tween the p a rt i n a dv a nce a nd the

p a rt b ehi nd a n d bet ween the u pp er p a rt a nd the u n d er p a rt ;
,
w h il e th ere i s c ompl et e corres p ond ence b et ween th e tw o s id es .
E xt erna lly the lik enes se s a n d di fferen c es a m ong limb s a nd ,
i nt erna lly th e lik enes s es a nd differenc es a m ong v ert eb ra a r e ,
exp ress ibl e i n t er m s of this sa m e l a w .
A nd h ere i nd eed w e m a y s ee c l ea rly th a t th es e t ru ths a re

, ,
c oroll a ri es f rom th a t u ltim a t e t ru th t o which a l l phenom ena

o f E v ol u ti on a r e refer a bl e I t is a n i nevit a bl e d e d u cti on
.
f rom the p ers i s t ence of f orce th a t org a n ic f orm s whi ch h a ve

,
been p rog ress iv ely ev olv ed mu s t p res ent j u st th os e f un d a

,
m ent a l t ra i t s of form which we fi nd th em p res ent I t ca nnot .
bu t be th a t d u ri n g t he i nt erc ou rs e b et ween a n org a ni s m a nd

i ts en vi ronm ent equ a l f orc es a ctin g u nd er equ a l c on diti on s
,
mu st p rodu ce eq u a l effect s ; for to s a y otherw i s e i s by i m ,
p lic a ti on t o s a y th a t s ome f orce c a n p rod u ce m ore or l es s

,
th a n i ts equ iv a l ent effec t whi c h i s t o d eny the p ers i s t enc e of

,
f orce H enc e th os e p a rt s of a n org a ni s m w hich a re by i ts

.
,
ha bits of li fe exp ose d to lik e a m ou nts a nd lik e com bi na

,
ti ons of a cti on s a nd rea cti ons m u s t d evel op a lik e ; whil e

,
u nlik enes s es of d ev el op m en t m u s t a s u na v oid a bly f oll ow
u n lik ene s s es a m on g th ese a g enc i es A n d thi s b ei ng so a l l

.
,
the s p eci a liti es of symm et ry a n d u n s ymm et ry a n d a symm et ry

which we h a v e t ra ced a re necess a ry c onsequ enc es
,
.
P A RT V .
P HYS I OL OGI C A L D E VE L OP ME NT .
C H AP T E R I .
TH E O
P R BL E M S OF P H Y S IO L O GY .
TH Eq ti b e t r ea t e d u n d er t he a b ov e titl e
§265 . u e s o n s t o
a re w id ely di fferen t fr o m th ose w hi c h it or di na r ily exp r es ses .
We h a ve n o a lt erna tiv e h owev er bu t t o u se P hys i ol o gy i n

, ,
a s en s e c o ext en s iv e w ith
-
t h a t i n which w e h a v e u s ed
Morph ol ogy We mu s t h ere con s id er the f a c t s of fu n cti on
.
i n a m a nner p a ra ll el t o t h a t i n whi c h we h a v e i n th e f ore ,
g oi ng P a rt c ons id ered the f a c t s of s tru ct u re A S hith ert o

,
.
, ,
we h a v e c onc erne d ou rs elv es with t h ose m os t genera l ph en o

m en a of orga n i c for m which h oldi ng i rr es p ec tiv e of c l a ss
,
a n d o r d er a nd s u b ki n gd o m
-
ill u s t ra t e the p roces se s of
,
i nt egra ti on a nd differen ti a ti on ch a ra ct eri zing E v olu ti on a t

l a r ge ; s o n ow w e h a v e t o c oncern o u rse lv es with t he evi
, ,
d eu ces o f th os e differenti a ti ons a nd i n t egr a ti ons of orga nic

fun c ti on s whi c h h a v e simu lt a neou s ly a ri sen a nd whi ch ,
s i m il a r ly t ra n s c en d th e limit s of z oo l ogi c a l a n d b ot a nic a l
divi s i on s H ow h et erogeneiti es of a c ti on h a v e p rogres s ed

.
a l on g w ith h et er o gene i t i es o f s t ru ctu r e — th a t i s th e i nq u i ry
b ef ore u s ; a nd obvi ou s ly i n p u rs u i ng it a l l th e sp eci a liti es

, ,
with whi c h P hy s i ol ogy u s u a lly d ea l s c an s erv e u s only a s

m a t eri a l s .
B efore ent er i n g on th e s t u dy o f Mor ph o l o gi c a l D ev el op

m ent, it wa s p oi nt ed ou t th a t w hil e fa c t s of s t ru ct u re m a y
be em pi ri c a lly gen er a li z ed a p a rt f rom fa c t s o f fu ncti on th ey ,
c a nnot be ra ti on a lly i nt erp ret ed a p a r t ; a nd th rou gh ou t the

239
240 P H Y S IOL OGICA L D E V E L OP ME NT .
foreg oi ng p a g es thi s t ru th h a s b een m a d e a bu n d a ntly m a ni

f es t H ere we a re obliged to recogn i ze th e i nter d ep end en ce
.
-
still m ore di s ti nc tly ; for t he ph enom ena of fu ncti on ca nn ot

e v en be con c e iv e d w ith ou t di r ect a n d p er p et u a l cons c i ou s n es s
o f th e ph eno m ena o f s t r u c t u r e .T h ou gh t he su bj ect m a t t er -
of P hy s i ol o gy i s a s b r oa dly di s ti ngu i s h e d from t h e s u bj ect
m a t t er of Morph ol ogy a s m oti on i s fro m m a tt er ; yet j u s t a s ,
t h e l a w s of m o ti on ca nn o t be k n own a p a rt from s om e m a tt er
m ov ed s o th ere c a n be no k nowl edge of f u ncti on w i th ou t a
,
kn owl edge of s om e s t ru ct u re a s p erform i n g f uncti on .
Mu ch m or e th a n t hi s i s obvi ou s T h e s tu dy o f fu ncti ons

.
,
c on s id ered from ou r p res en t p oi n t of vi ew a s a ri s i ng by

E v ol u ti on m u s t be c a rr i e d on m a i n l y by th e s t u dy of th e
,
c orrel a tive s t ru ct u res D ou b t l e s s by e xp erim ent i ng on th e

.
,
org a n i s m s w hi ch a r e g row i n g a n d m ovi n g a r ou n d u s w e m a y ,
a s c er t a i n th e c onn e xi on s exi s ti n g a m on g c er t a i n o f th e i r
a cti ons w hil e w e h a v e lit t l e o r n o k n ow l edg e o f t h e s p ec i a l

,
p a rt s concerned i n th os e a ct i on s I n a livi ng a n im a l t h a t
.
c a n be conv en i ently k ept u n d er o b serv a ti on w e m a y l ea rn

,
th e wa y i n w hich con s pi cu ou s fu n cti on s v a ry t og eth er— h ow

’
t he ra t e of a m a n s p u l se i nc rea s es w ith th e a m ou n t o f
m u s cu l a r ex erti on h e i s u nd er g oi ng ; or h ow a h orse s ’
ra pidity o f b rea thi n g i s i n p a r t d ep en d ent on h i s s p eed .
Bu t th ou gh ob s erv a ti ons o f thi s ord e r a r e i ndi s p ens a bl e

t h ou gh by a ccu m u l a ti on a n d com p a r i s on o f su ch o b s erv a ti on s
we l ea rn whi ch p a r t s p er form which f u n cti on s—4 th ou gh s u c h
o b ser v a ti o n s p ros ecu t ed s o a s t o di s cl os e th e a cti ons of a l l
,
p a rts u nd er a l l ci rc u m s t a nces c on s tit u t e wh en p rop erly

, ,
g enera li zed a nd c o ordi na ted wh a t i s c om m on ly u nd ers t ood

-
,
a s P hy s i ol o gy ; yet su c h o b s erv a ti on s h elp u s bu t a littl e
w a y t owa rd s l ea rni ng h ow fu ncti on s ca m e t o be es t a bli s h ed

a n d S p eci a li z ed We h a v e next to no p ower of t ra ci ng u p
.
th e gen es i s of a functi on c ons id ered p u rely a s a f u nc ti on— no

o pp ort u n ity o f ob s e rvi ng t h e p r o gres s iv ely i n c rea s i n g qu a n
-
ti ti es of a giv en a cti on th a t h a v e a ri s en i n a ny or d er of
o r g a ni s m s . I n nea rly a l l ca s es we a re a bl e only to S h ow
THE P R OBL E MS OF P H Y S I OL OGY .
24 1
th e g rea ter gr owth of th e p a rt w hi ch we h a v e fou nd p erform s

t he a c t i on a n d t o in fer th a t g rea t er a cti on of th e p a rt h a s
,
a c c o mp a n i ed g rea t er g row th of it T he t ra ci ng ou t o f
.
P hy s i ol ogi ca l D ev el op m en t th en b ec om es s u b s t a nt i a lly a
, ,
t ra ci n g ou t o f t h e d e v el opm en t o f the org a n s by w hi c h t he

f u ncti on s a re k n own t o be di s ch a rged— th e di fferent i a ti on
a nd i n t eg ra ti on of t he f u n cti o n s b ei n g p r es u m e d t o h a v e
p rog res sed h a nd i n h a nd wi th the di fferenti a ti on a nd i nt eg ra

ti on o f th e org a ns Betw een the i nq u i ry p u rsu ed i n P a rt I V
.
,
a n d t h e i n qu i ry t o be p u rs u e d i n thi s P a r t t h e c on t ra s t is ,
th a t i n the fi rs t pl a ce fa c t s of s t ru ct u r e a re now t o be u s ed
, ,
t o i nt er p ret fa ct s of fu nc ti on i n s t ea d o f c onv ersely ; a nd i n

, ,
t he s econ d pl a ce the f a c t s of st ru c t u re t o be s o u s ed a re not

,
th o se of cons pi cu ou s S h a p e s o m u ch a s th os e of mi nu t e t ext u re
a nd ch em i c a l c omp os iti on .
§ 2 6 6 T he p robl em s of P hy s i ol ogy i n the wid e s ens e

.
,
a b ov e d es cribe d a re lik e th e p r obl e m s o f Morph ol o gy t o be

, , ,
c ons idere d a s p robl em s t o whi ch a n s wers m u s t be giv en i n

t erm s of i nc id ent forces On th e hyp oth es i s of E v ol u ti on
.
th es e s p eci a li za ti ons o f ti s su es a n d a cc o mp a nyi ng c onc en t ra

ti ons of f u ncti ons m u s t lik e the S p ec i a li za ti on s o f s h a p e i n
, ,
a n o r g a ni s m a n d i ts c o m p on en t divi s i o ns be du e t o the a c ,
ti on s a n d rea cti ons whi ch i ts i nt ercou rse with th e envi ronm ent
i nv olv es ; a nd the t a s k b ef ore u s i s to expl a i n h ow th ey a re
w rou ght — h ow th ey a re t o be c om p reh end ed a s res u lts o f
s u c h a c ti on s a n d r ea cti on s .
Or t o d efi n e th es e p robl ems s till m ore sp ec ific a lly — T h os e

,
ext r em ely u n s t a bl e Su b s t a n c es c om p os i n g th e p r o t o pl a s m
o f whi c h o rg a ni s ms a re m a i nly b u ilt h a v e t o be t ra c ed

,
th r ou gh the v a ri ou s m odifi ca ti ons i n th ei r p r op erti es a nd

p ower s th a t a re en t a il ed on th em by c h a n ges o f rel a ti on t o
,
a genc i e s o f a l l ki nd s T h o se orga n i c c oll oid s whi ch p a s s fro m

.
liqu id t o s olid a n d f rom s ol u bl e t o i ns ol u bl e on the slight es t

m o l ecu l d t b
a r i s u r a n ce— th os e a lbu m enoid m a tt ers which a s ,
we s ee i n cl o tt ed bl o o d or the coa gu l a bl e lymph p ou red

62
24 2 P H Y SI OL OGICA L D E VE L OP ME NT .
ou t on b ra d ed s u rfa ces a nd c a u s i ng a dh es i on b et ween

a
i nfl a m ed m em b ra nes a s s u m e new form s wi th t he g rea tes t

,
r ea di n es s— a r e t o h a v e th e i r m et a m or ph os es st u di ed i n c o n
nexi o n wi t h th e i n fl u en c es a t w o rk T h ose c om p ou nd s whi ch

.
,
a s w e s ee i n t h e qu i c kly a c qu i r e d b r own ne s s o f a bit t en a ppl e

-
o r i n th e d a r k s t a i ns p r o d u ce d by t h e mi l ky j u i ce o f a D a n d e
li on im m edi a t ely b e gi n t o a l ter w h en t he s u rrou ndi n g a ct i ons

,
a lt er a r e t o be ev ery wh er e c o ns id er e d a s u n d e rg o i ng m o di fi
“
c a ti ons by m odifi ed con diti on s Org a ni c b odi es cons i st in g

.
,
o f s u b s t a nc es th a t a s I h ere p u rp osely r emi nd t h e rea d er

, ,
a r e p r one b ey on d a l l ot h ers t o c h a n g e w h en t h e i n c id en t
f orc es a re ch a ng ed w e mu s t con t em pl a t e a s i n a l l th ei r p a rt s
,
differently ch a n ged i n res p on se t o th e di fferen t ch a n ges of

t he i n c id ent forces A nd th en we h a v e t o reg a r d th e con
.
c om it a nt di fferenti a ti on s of th ei r rea ct i ons a s b ei ng c oncom i

t a nt differenti a ti ons of th ei r fu n cti on s .
H er e a s b efore we m u s t t a k e i nt o a ccou n t t w o cl a sses

, ,
of fa ct or s We h a ve t o b ea r i n mi nd the i nh erit e d res u l t s o f

.
a cti on s t o w hich a n tece d ent org a n i s m s w ere e xp os ed a nd t o ,
j oi n with th es e the res u lts of p r es ent a cti ons E a ch org a ni s m

.
i s t o be c o ns id ered a s p res enti ng a m ovi ng equ ilib ri u m of

fu n c ti ons a nd a c o r rel a tiv e a rr a n gem en t o f s t ru ct u res p ro
, ,
du ced by t h e a gg reg a t e of a c t i on s a n d r ea c t i on s th a t h a v e
t a k en pl a ce bet ween a l l a nc es t ra l orga ni s m s a n d th ei r cmvi
r on m ent s Th e t end ency i n e a ch org a ni s m t o rep ea t thi s
.
a dj u s t ed a r ra n g em ent o f fu n c ti ons a n d s t ru c t u res m u s t be ,
reg a rd ed a s fro m tim e t o ti m e i nt e r fere d w ith by a c t i on s t o
w hi ch i t s i nh eri t e d eq u i l ib r i u m i s n ot a dj u s t ed— a c t i on s t o
w hi ch th eref or e i ts eq u ilib r i u m h a s t o be re a dj u s t ed
, ,
-
A nd .
i n s t u dyi ng phy s i ol ogi ca l d ev el o p m en t w e h a v e i n a l l ca s es

t o c ont empl a te the p ro g res s i ng c o m p r omi se b et w een th e ol d
a nd th e n ew en di n g i n a r es t or ed b a l a nc e o r a d a p t a ti on
,
.
Mani fes tly ou r d a t a a re s o s ca nty th a t n othi ng m o re

th a n v ery g enera l a nd a pp ro xi m a t e i nt erp ret a ti ons of thi s
ki nd a r e p os s ibl e I f t h e h yp oth es i s o f E v ol u ti o n fur
.
ni s h es u s with a ru d e c onc epti on of the wa y i n which the

T HE P R OBL E MS OF P H YSI OL OG Y .
243
m ore c on s pi cu ou s a nd imp ort a nt di fferenti a ti on s of f u ncti on s

h a ve a ri sen it i s a s mu ch a s ca n be exp ect ed
,
.
§
267 . It w ill be
b e st fo r b r evity a nd cl ea rnes s
,
t o d ea l
,
w ith th es e phy s i ol ogi c a l p robl em s a s we d ea lt wi th th e

m orph ol ogi ca l ones— t o ca rry on the i n d u ctiv e st a t ement a nd
t he d ed u ctiv e i nt erp ret a ti on h a n d i n—h a nd : s o di s p os i ng o f
-
ea c h g enera l t ru t h b e fore p a ss i n g t o t he n ext T rea ti ng

.
s ep a r a t ely v eget a l org a n i s m s a n d a n im a l o r g a n i s m s w e w ill

,
i n ea c h ki ngd o m c ons id er — fi r st th e physi ol ogic a l di fferen

,
t i a t i ons a nd a c c om p a nyi ng ch a ng es of s t ru c t u re which a r i s e

be t ween ou t er ti s su es a n d i nner ti s s u es ; next th ose w hi c h
,
a r i se be t w een differen t p a r t s o f th e ou t er ti s su es ; a n d fi na lly , ,
th os e whi ch a r i s e b et ween differen t p a rt s of th e i nner ti s su es .
Wh a t littl e h a s t o be s a id c oncerni ng phy s i ol ogic a l i nt egra

ti on mu s t c om e l a s t For th ou gh i n t ra ci ng u p Mor ph o
.
,
l ogic a l E v ol u ti on we h a v e t o s t u dy th ose p roces ses of i nt e

,
g ra ti on by whi ch or g a n ic a gg reg a t e s a r e f orm ed b ef ore ,
s t u dyi n g th e di f f erenti a ti ons th a t a ri se a m ong th ei r p a rt s ;

we m u s t con t ra ri w i se i n t ra c i ng u p P hy s i ol ogi ca l E v ol u ti on
, , ,
s t u dy t h e g en es i s o f t h e di fferen t f u ncti ons b efor e we s t u dy
the i n t erd e p en d en c e th a t ev ent u a lly a ri s es a m ong th em a nd

c ons titu t es phy s i ol ogic a l u nity .
CH A P TE R I I .
I
D FFE RE N T A T I IO N S B E T WE E N TH E O U TE R AND INNE R
T SSI UE S OF P L A N T S .
2 68 . impl est pl a nt p res ent s a c ont ra s t b etween its

TH E s
p eriph era l su b s t a nce a nd its c ent ra l s u b s t a nce I n ea ch p ro .
t ophyte be it a s ph eri c a l c ell or a b ra nch e d t u b e or s u c h

, ,
a m or e S p ec i a li z ed for m a s a D es mid
-
a m a r k e d u n lik en es s
,
e xi s t s b et ween th e limiti n g l a y er a n d th a t whi c h it limits .
T h ese v eget a l a ggrega t es of th e fi rs t ord er m a y differ w id ely

fr om one a n o th er i n the n a t u res o f th ei r ou ter c oa t s a nd i n
the n a t u res of th ei r cont ents A s i n the P a lm ell a f orm o f
.
-
one o f t h e l ow er A l g ae th ere m a y e xi s t a c l o thi ng of j elly

,
or a s i n D i a t om
,
t h e w a ll s m a y t a k e the f orm of s ilici ou s
,
v a lv es v a ri ou s ly s cu lpt u red The c on t a i ned m a tt er m a y be

.
p a rtly or wh olly h ere g reen th ere red a nd i n oth er c a s es

, ,
b rown B u t a mid a l l th es e div ers iti es th ere i s thi s one

.
u n i for mity— a s t ron g di s ti ncti on b et ween t h e p a rt s i n c ont a c t
w ith th e en vi ron m ent a nd th e p a rts n ot i n c on t a c t with th e

envi ronm ent .
Wh en w e rem emb er th a t this t ra it i s one whi ch th es e

S impl e livi ng b odi es h a v e i n co mm on w ith b odi e s th a t a re
not livi ng — wh en w e r em emb er th a t ea c h i n org a n ic m a s s
ev ent u a lly h a s i t s ou t er p a rt m ore or l es s di f f erenti a t ed from

its i nner p a rt h ere by oxid a ti on th ere by d ryi n g a nd el se
, , ,
wh ere by th e a cti ons of l ight of m oi s t u re of f ros t ; we c a n

, ,
s c a r c ely res i s t th e concl u s i on th a t i n th e one c a s e a s i n th e

,
oth er th e c o nt ra st is due t o th e u nlik e a cti on s t o whi ch the

,
2 44
TH E OU TE R A ND I N NE R T ISS UE S OF P L A NTS .
24 5
p a rt s a re s u bj ect Giv en a n origi na lly h om ogeneou s p orti on

.
-
o f p rot o pl a s m a nd it f o ll ow s fr o m t h e g ene r a l l a ws of E vol u

,
ti on ( F i rs t P r i n ci p l es 1 49 fi rs t th a t it mu s t l os e
, ,
i t s h o m ogen eity a n d sec ond th a t t h e l ea di ng di s s imil a riti es

, , ,
m u s t a r i s e b etween the p a rt s m os t di s s i m il a r ly c on di ti on ed
-
th a t i s b e tw een t h e ou t s id e a nd th e i n s id e
,
T he ex t eri or .
m u s t b ea r a m ou nt s a nd ki nd s o f f or c e u nlik e the a m ou nts

a n d ki n d s w hi c h t h e i n t er i or b ea r s ; a n d f ro m t h e p ers i s t ence
o f f or ce it f o ll ow s i nevit a bly th a t u n lik e eff ects m u st be
w rou gh t on th em— th ey mu s t be di fl erenti a ted .
Wh a t i s the li m i t t owa r d s which the d i ff erenti a ti on t ends ?

We h a v e s een th a t t h e re—di s t ribu ti on of m a tt er a n d m oti on
wh ence u nd er cert a i n c on diti on s
,
ev o l u ti o n r esu lt s ,
ca n ,
n ev er cea s e u n til eq u ilib r i u m i s rea c h ed — p r o xim a t ely a
m ovi n g eq u ilib ri u m a nd fi na lly a com pl et e eq u ilib ri u m ( F i rs t

,
P ri nci p l es ,
1 70 H enc e th e di fferen ti a ti on m u s t
,
g o on u n til it es t a bli s h es su c h di fferences i n th e p a rt s a s

s h a ll b a l a n ce th e differen ces i n th e for ce s a c ti n g on t h e m .
Wh en d ea li ng wi th equ ilib ra ti on i n gen era l we s a w th a t t his ,
p r oc es s i s w h a t i s ca ll ed a d a pt a ti on ( F i rs t P ri n ci p l es 1 73 ) ,
a nd,
i n thi s w ork w e s a w th a t by it th e t ot a l ity of fu n c
,
ti ons of a n org a n i s m is b r ou ght i nt o corres p ond en ce with the

t ot a lity of a cti ons a ffecti n g it 1 59 Ma n i festly i n
thi s c a se a s i n a l l o th ers eith er d ea t h or a dj u s tm ent mu s t
, ,
e v en t u a lly r e s u lt A f orce f a lli ng o n one of th es e min u t e

.
a ggre g a t es o f p r o t opl a s m m u s t exp en d i t self i n w or ki n g its

,
e qu iv a l ent o f c h a n g e I f thi s f orce i s su c h th a t i n expen d

.
i ng i t s el f it di s t u rb s b ey on d rectifi ca t i on t h e b a l a nce of t he
o r g a ni c p rocess es t h en th e a ggr eg a t e i s di s i n t eg r a t ed or de
,
c om p os e d .B u t i f it d oes n ot o v erth row th a t m o vin g equ i

lib ri u m const i tu ti ng th e li fe of th e a ggr eg a t e th en th e ,
a gg reg a t e conti nu es i n th a t m od i fi e d for m p r o d u ced by th e
e xp en dit u r e o f t h e for ce T h u s by di rec t equ ilib ra ti on con

.
, ,
t i nu a l l y f u rth ered by i ndi re ct equ il i b ra ti on th ere m u s t a ri se ,
t hi s di s t i ncti o n b et w een t he ou ter p a rt a d a pt e d t o m eet ou t er

f orc es and the i nner p a rt a d a pt ed to m eet i nner f or c es A nd
, .
24 6 P H YSIOL OGI CA L D E VE L OP ME NT .
t h ei r p ectiv e a cti ons a s thu s m eetin g ou t er a nd i nner

r es ,
f orc es mu s t be wh a t we c a ll th ei r res p ectiv e f u ncti ons

,
.
§ 269 . A gg reg a t es d of
o r d
th e
er exhibi t
s econp a r a ll e l
t ra it s a d m i t ti n g of p a ra llel i nt erp ret a ti ons

,
I nteg ra t ed .
m a s s es of c ell s or u nit s h om ol og ou s wi th p rot ophyt es ,
h a bit u a lly S h ow u s c ont ra s t s b et ween the ch a ra ct ers of the

s u p erfici a l ti s s u es a nd t he c ent ra l ti s su es S u ch a m ong th es e.
a gg reg a t es o f th e s ec on d o r d er a s h a v e th ei r com p on en t u n it s
a r ra n g ed i nt o t h rea ds or l a mi na s i ngl e or d ou bl e c a nn ot o f
, , ,
c ou rs e fu rni s h c on t ra s t s of this ki nd ; for a l l thei r u ni t s a re

,
a s m u c h e xt ern a l a s i n t ern a l We mu s t t urn t o the m ore or

.
l es s m a s s ive f or m s .
Of th ese a m ong F u ngi th e c om m on P uff b a ll is a g oo d

, ,
-
e x a m pl e — g oo d b ec a u s e it p resen t s thi s f u n d a m en t a l di f feren

t i a t i on bu t littl e c om pli ca t ed by oth ers I n it we h a v e a .
c ort i ca l l a yer o f i nt erwov en hyph a obvi ou s ly u nlik e the

m a ss of sp ores whi c h i t i n cl oses S o fa r a s the u nlik eness .
b et ween ext er na l a n d i n t ern a l p a rt s i s c on c erned we s ee h ere ,
a rel a ti on a n a l o g ou s t o th a t exi s ti n g i n th e S impl e c ell ; a n d
we s ee i n it a S i m il a r m ea n i n g : th ere i s a phy s i ol ogic a l

di ff erenti a ti on c orres p ondi ng t o the di fferenc e i n the i ncid enc e
o f f orc es .
U nd er v a r i ou s form s th e A l gae S h ow j u s t th e s a m e rel a

ti on . Wh ere a s i n C odi u m B u rs a we h a ve the r a m i fi ed
, ,
t u bu l a r b ra nch es of th e th a ll u s a ggreg a t ed i nt o a h oll ow

gl obu l a r m a ss th e ou t er a n d inner s u r fa c es a re c on t ra s t ed
,
b ot h i n c ol ou r a nd s t ru ct u re th ou gh the t u b u l es c om p os i n g
,
t h e two s u rfa c es a re conti n u ou s w i th one a n other I n Ri va .
l a ri a a g a i n w e s ee th e lik e b oth i n th e r a di a l a rr a n ge m en t
, , ,
o f the i m b edd e d th rea d s a n d i n th e di ffer ence o f c ol ou r
b et ween the e xt eri or of t he i m b eddi ng j elly a nd i ts i n teri or .
T he m ore d ev el op ed A l g ae of a l l ki nd s r ep ea t th e a ntith es i s
-
.
I n b ra nch ed s t em s w h en th ey c ons i s t o f m or e th a n S i ngl e

,
r ow s o f c ell s th e ou t er c ell s b ec o m e u nlik e th e i nner a s S h own

, ,
i n Fig 3 5 . S uch typ es a s Chrysym eni a r os ea S h ow u s thi s

.
THE OU TE R AN D I NNE R T ISS U E S OF P L A NTS .
2 47
un lik enes s v ery c ons pi cu ou s ly A nd it h old s even wi th ri b

.
bon s h a p ed f ron d s
-
Wh erev er one of th es e i s c omp os ed of
.
th ree f ou r o r m ore l ayers a s i n L a m i na ri a a nd P u nc ta ri a

, , , ,
the c ells o f th e ext erna l l a ye rs a re s t rongly di s ti ngu i s h e d

f rom th os e of t he i nterna l l a y ers b oth by th ei r c omp a ra tiv e
,
sm a lln es s a nd by t h ei r d eep c ol ou r .
§ 2 7 0 T he high er pl a nt s v a ri ou s ly di s pl a y th e like
.
fun d a m ent a l di s ti n cti on be tw een ou t er a n d i nner ti s su es .
E a c h l ea f thi n a s it i s ex emplifi es t hi s diff erenti a ti on of t he

, ,
p a rt s i m m edi a t ely i n c ont a ct wi th t h e envi ronm ent from the

p a rt s n ot i n i m m edi a te c on t a ct w i t h th e envi ronm ent I ts .
epid er m a l cell s for m i ng a p r ot ec ti n g e n v el op e div erg e phy s i

, ,
c a lly a nd ch e m i c a lly f r o m th e m es o phyll c ell s whi ch ca rry ,
on t h e m ore a c tiv e fu n c t i ons A nd th e cont ra s t m a y be

.
ob s er v ed t o es t a bli s h i ts el f i n t h e c ou r se o f d ev e l o p m en t At .
fi rs t the c omp on ent ce ll s of the l ea f a re a l l a lik e ; a n d thi s

unlik enes s b etween th e c ell s o f th e ou t er a n d i nner l a y ers ,
a ri s es s i m u lt a n eou s ly w i t h t he r i se o f di fferenc es i n th ei r c on
di ti ons — d i ff erences t h a t h a v e a ct ed on a l l a nc es t ra l l ea v es
a s th ey a ct on th e i n divid u a l l ea f .
A n u nlik ene s s m ore m a r k ed i n ki n d bu t S i m il a r i n m ea n

i ng e xi s t s b etw een th e b a rk of ev er y b ra nch a nd t h e ti s s u es
,
i t cl oth es . T he ph a nog a m i c a xi s es p ec i a lly w h en i t u n d er

,
g oes w h a t i s kn own a s s econ d a ry thi ck eni n g i s com m onl y ,
c h a r a ct er i z ed by a n ou t er z one o f cell s ( t h e c o rk l a y er ) di f fer

i ng fro m th e i nner l a y ers i n c h a r a c t er a n d f u nct i on a s it ,
di ffers f rom th em i n p os iti on S u bj ect a s thi s ou t er l a y er i s

.
t o the u n m i tig a t ed a ct i ons of f orc es a rou n d — t o a b ra s i on s t o ,
ext rem es o f h e a t a n d c old t o ev a p or a t i on a n d s oa ki n g w i t h

,
w a t er— i t s u n i t s h a v e t o be b rou gh t i n t o eq u ilib r i u m w i th

th ese m ore vi ol en t a cti o n s a n d h a v e a cqu i red m ol ecu l a r
,
c on s ti t u t i o ns m or e s t a bl e t h a n t h o s e o f th e i n t e r i or cell s .
T h a t i s t o s a y th e forc es w hi ch di ffer en ti a t e th e c orti c a l p a rt

,
from th e res t a re th e for ces w hich i t h a s t o r es i s t a nd fro m ,
whi ch it p a s siv ely p rot ect s th e p a rt s withi n . H ow

24 8 P H Y SI OL OGI C A L D E V E L OP ME NT .
cl ea rly this h eterogenei ty o f s t ru ctu re a nd functi on i s c ou s e

q uent u p on i n t ercou rs e wi t h the envi ronm ent ev ery t ree
‘
a n d s h ru b s h ow s T he y ou n g S h o ot s a lik e of a nnu a l s a nd
.
,
p erenni a l s a re qu it e green a nd s oft a t thei r ext remi t i es

,
.
A m on g pl a n ts of s h or t liv es t h er e i s u su a lly bu t a s light

,
d ev el opm ent of b a rk or none a t a l l : s u ch t ra c es o f it a s the

su r f a c e o f th e a xis a cqu i r es b ei n g seen on ly a t i t s l ower m os t
or old es t p or ti on I n l ong liv ed pl a nt s h owev er thi s for m a

.
-
, ,
ti on of a t ou gh op a qu e coa ti ng t a k es pl a ce m ore ra pidly ; a nd

S h ow s u s di s ti nc tly th e c onnexi on b et ween t he d eg r ee o f
di fferenti a ti on a nd the l ength of exp os u re For i n a g row i n g .

,
twig w e s ee th a t the b a r k i n vi s ibl e a t the bu d thi ck en s by

, , ,
i ns ens ibl e gr a d a ti ons a s we go d ownwa rd s t o the j u ncti on o f

t he t wig w i th th e b r a nc h ; a nd w e c om e t o s till t hi c k er p a r t s
of it a s we d es cend a l on g t h e b r a nc h t ow a rd s t he m a i n s t e m .
Moreov er on ex a m i n i ng m a i n s t em s we fi nd th a t whil e i n
,
s om e t rees t h e b a r k c ra ck e d by e xp a n s i on o f th e w o od d ro p s
, ,
of f i n fl a k es l ea vi ng exp os ed p a t ch es o f the i nner ti s s u e whi c h

,
p res ently b ec om e g reen a nd fi na lly d evel op new b a rk ; i n

o th er t r ees t h e ex foli a t e d fl a k es c on ti nu e a dh erent a nd i n th e ,
c ou rs e of y ea rs form a ru gged fi ss u red c oa t : s o p rod u ci ng

a still m ore ma rk ed cont ra s t b et ween ou t s id e a n d i n
s id e . Of cou rs e th e es t a bli s h m en t of t hi s h et er o
g en ei ty i s f u rth e red by n a t u ra l s el ec t i on w hi ch w h ere a , ,
p rot ectiv e c overi n g i s nee d ed giv es a n a dv a nt a ge to th os e

,
i ndivid u a l s i n w hi c h it h a s b ecom e s t r onges t Bu t th a t thi s .
div ergence of s t ru ct u re c om m ences a s a di rec t a d a p ta ti on i s ,
c l ea rly S h own by ot h er f a cts th a n th e f or eg o i n g T h er e i s .
th e f a c t th a t m a ny o f th e pl a n t s whi ch i n ou r ga rd en s
d ev el op b a r k with c on s id era bl e ra pidity do n ot d ev el op it ,
with the s a m e ra pidity i n a g reenh ou s e A n d th e re i s th e

.
fa c t th a t pl a n ts whi ch i n s om e cli m a t es h a v e th ei r s t em s
, ,
c overed only by thi n s em i t ra n s p a rent l a yers a cqu i re thick

-
,
op a q u e l a y ers wh en t a k en t o oth e r c li m a t es .
J us t no ti ng for the sa ke o f com pl et enes s th a t i n the

, ,
r o ot s o f the high er pl a n t s th ere a r i se s a cont ra s t b et ween

THE OU T E R A ND INNE R T ISS U E S OF P L A N TS .
24 9
t
ou er a nd i nner p a rt s p a ra ll e l t o th e one w e h a v e tra ced i n
,
th ei r b ra nch es l et m e d ra w a tt enti on t o a not h er di fferenti a

,
ti on o f the s a m e u lti m a t e na t u re which t he high er pl a nt s ,
e xhibit t o u s— a di fferenti a ti on w hi c h fa m ili a r th ou gh it i s , ,
g a i ns a new m ea n i ng by a s soc i a ti on w ith th os e na m ed a b ov e ,
a nd m a k es th ei r m ea n i n g s t ill m or e m a n if es t E a c h g rea t .
pl a nt s h ow s it When by the b u ddi ng o f a x es ou t of a x es

.
, ,
t h ere i s p rod u ced one o f th o s e highly c om p ou n d ed P ha n og a m s -
w hi c h we c a ll a t ree t he cent ra l p a rt o f th e a gg reg a t e be

,
c om es fu nc t i on a lly a n d s t ru c t u r a lly u n lik e t h e p eriph era l
p a rt. On l ooki n g i n t o a l a rg e t ree or ev en a s m a ll one ,
wh i c h h a s thick fo li a g e lik e t he L a u r el we s ee t h a t t h e i n
, ,
t erna l b ra nch es a re a lm os t or qu it e b a re o f l ea v es w hil e th e ,
l ea f c l a d b ra nc h es f orm a n ex terna l s t ra t u m ; a nd a l l ou r
-
e xp eri enc e u n it e s i n p r o vi n g t h a t t hi s con t ra s t a r i s es by
d egr ees a s f a s t a s the gr owth of the t ree ent a il s a c ont ra s t be

,
t wee n the c onditi ons t o whi c h i nner a nd ou t er b ra nch es a re

ex p osed N ow wh en i n th ese m os t c om p os it e a gg reg a t es
.
,
-
,
w e s ee a differen ti a ti on be t w een pe r iph era l a n d c en t ra l p a rt s

d em on s t ra bly ca u s ed by a di fference i n the r el a ti on s of th es e
p a rts to envi roni ng forces w e get s u pp or t for the conc lu s i on
j ,
oth er wi se rea ch e d th a t th ere i s a p a r a ll el ca u s e for th e p a r a l

,
l el d i fl erenti a ti ons exhibit ed by a l l a gg reg a t es o f l ower ord er s

— b ra nch es l ea v es c ell s .
, ,
§ 271 . B e f ore
l ea vi n g thi s m os t g en er a l phy s i ol ogi ca l
di fferenti a ti on it m a y be w ell to s a y s om ethi ng r esp ecti ng
,
cert a i n se c on d a ry u n lik enes ses whi c h u su a lly a r i se be t w ee n
i nt eri or a n d ext eri or F or th e c on t r a s t i s not a s m ight be

.
,
su pp os e d f r o m t h e for eg oi n g d es cripti on s a s i m pl e c ont r a s t : ,
it i s a c om p ou n d c ont ra s t T he ou t er s t ru ct u r e it s elf i s
.
u su a lly di vi s ibl e i n t o c o nc ent r i c s t ru c t u r es T hi s i s e qu a lly .
t ru e of a p r ot oph yt e a n d of a pha nog a m i c a xi s Bet w een .
th e c ent re o f a n i n d ep end ent v eget a l cell a n d i t s s u r fa ce ,
th ere a re a t l ea s t tw o l a y ers ; a nd th e b a r k coa ti n g th e s u b

s t a n ce o f a s h o o t be s id es b ei n g it self c omp ou nd i ncl u d es
, ,
25 0 P H Y SIOL OGI CA L D E VE L OP ME NT .
a n ot h er ti ssu e lyi n g b et ween it a nd the w ood Wh a t is the .
phy s ic a l in t erp ret a ti on of th es e f a ct s ?

Wh en a m a s s of s om ethi ng we di s t ingu i s h a s i nert m a tt er
i s exp os ed t o ext erna l a g enc i es c a p a bl e o f w orki ng ch a n ges i n
i t— wh en it i s ch em ic a lly a ct ed u p on or wh en b ei n g dry it
, , ,
i s a ll owed t o s oa k or w h en b ei ng w et i t i s a ll owed t o dry

, , ,
the c h a nges s et u p p rog res s i n a n equ a bl e w a y f rom the

s u r f a c e t owa rd s th e cen t re A t a ny ti m e d urin g the p roces s
.
s pp s i ng n o o th er a cti on s u p erv en es ) h e m o difi c a t i on

( u o t
w rou ght fi rs t c ompl et ed a t th e ou t s id e eith er g ra du a lly
, ,
dimi ni s h es a s we a pp roa ch th e cent re or c ea s es s u dd en ly a t

,
a cert a i n di s t a n ce from the cent re Bu t n ow s u pp ose th a t

.
t h e m a s s i n st ea d of b ei ng i nert i s th e s ea t of a ctiv e ch a nges

, ,
— S u pp os e th a t it i s a p o rti on of c o mpl ex c oll oid a l s ubst a nc e

,
p erm ea bl e by ligh t a n d by fl u id s ca p a bl e of a ffecti ng i ts

u n s t a bl e m ol ec u l es— su pp o s e t h a t i t s in t eri or i s a s ou r c e o f
f orces c onti nu a lly lib era ted a n d di ff u s i ng th em s elv es ou t

wa rds . I s it n ot lik ely t h a t w hil e a t the cent re th e a cti on
o f th e i nt ern a lly lib er a te d f or c es w ill d o mi n a t e a n d whil e a t
-
,
t he s u rfa ce th e a ct i on of th e envi ron i n g forc es w ill d omi na te ,
th er e w ill be b et ween t he t w o a certa i n pl a ce a t whi c h th ei r

a c ti on s b a l a n ce ? Ma y we n ot exp ec t t h a t thi s will be th e
pl a ce wh ere the m os t u ns ta bl e m a tt er exi s t s— the pl a ce ou t
s id e o f w hi ch t h e m a t t er b ecom es rel a t iv ely s t a bl e in th e
f a ce of ex tern a l forces a nd i ns id e of w hi ch th e m a tter be

,
c om es rel a tiv ely s t a bl e i n the f a c e of i nt ern a l forces ? A nd

m u s t w e n ot conc l u d e th a t th ou gh p a rt of th e a dj u s tm ent i s
d u e t o i ndi rec t eq u ilib ra ti on th e i n iti a ti on of i t i s du e t o
,
di rect equ ilib ra ti on ?

Bu t w e a re h ere c hi e fly c oncerned w ith th e m or e gen er a l
i nt erp ret a ti on w hi ch i s i nd ep en d en t o f a ny s u ch s p ecu l a t i on
,
a s t h e for e g oi n g . T h ese c ontra s t ed ti ss u es a n d th e c on tr a s t ed

f un cti on s th ey s ev era lly p er form a re be y on d q u es ti on s u b
, ,
o r di n a ted t o th e rel a t i on s o f ou t s id e a nd i n s id e A n d th e
.
e vid en c e m a k es it t ol era bly cl ea r th a t t h e u nlik e a c ti on s o r
f orces i n v olv ed by therel a ti ons o f ou t s id e a nd i n s id e d et er ,
mi ne th es e c ont ra s ts — p a rtly di rectly a nd p a rtly i ndi rectly .

I
D F F E RE N T A T I IO N S AM ONG TH E O UTE R I
T SS UE S or P L A NTS .
2 72 . m oti onl ess p rot ococcoid form s of l ower A l gae

TH E ,
whi c h d o not p erm a nently exp os e a ny p a rt s of th ei r s u rf a ces

t o a c ti on s u n lik e th os e w hich ot h er p a rt s a re exp os ed t o h a v e ,
no p a rt s of th ei r s u rf a ces u n lik e th e res t i n f u ncti on a nd

com p os iti on T hi s is wh a t th e hyp o th es i s p rep a r es u s for
. .
I f physi ol ogic a l differenti a ti ons a re d et er mi ned by differen ces

i n t he i nc id en c e o f f orc es th en th ere will be n o s u ch differ
,
enti a ti on s w h ere th ere a re n o s u c h diff er enc es C on t ra ri wi s e

.
,
it is to be exp ec t ed th a t the m os t c onspic u ou s u nlik enes s of

f u ncti on a nd m i nu t e s t ru ct u re will a ri s e b et ween th e m os t
di s s imil a rly c i rcu m st a nced p a rt s of the s u r f a c e We fi nd .
th a t th ey d o T he u pp er end a n d the l ower end or m ore

.
, ,
s t r ictly s p ea ki n g t h e free en d a n d th e a tt a c h e d end h a bi t u

, ,
a lly p r esen t th e s t ron ges t phy s i ol o gi c a l co n t r a s t s .
E v en a ggrega t es of th e fi r s t or d er ill u s t ra t e this t ru th .
S u c h s o ca ll ed u nic ell u l a r pl a n t s a s th os e d eli nea t ed i n

-
Figs 4 5 a n d 6 S h ow u s on c om p a r i n g the content s Of

.
, , , ,
t h ei r fix ed en d s a n d t h ei r l oos e en d s th a t di ffer ent p r ocesses

,
a re g oi ng o n i n th em a nd th a t di fferen t
, fu nc ti on s a r e
b ein g p erf orm ed by th ei r limiti n g m em b ra n es Ca u l er pa .
p r ol i fer a whi ch
“
, con s i s t s o f a littl e cr ee pi n g s t em wi th
r oot s b el ow a n d l ea v es a b ov e or igi n a ti n g i n th e g ro wth
“
,
o f a b o dy whi c h m a
y be rega r d ed a s a n i n divid u a l cell ,
s u ppli es a still b ett er ex a mpl e

-
. A m on g a g gre
25 1
25 2 P H YS IOL OGICA L D E VE L OP ME NT .
g a t es of the s ec on d ord er a lik e c onn exi on i s displ a yed i n

m or e v a ri ou s m od es bu t with equ a l cons i s t ency A s b ef ore .
,
t h e P u ff b a ll s erv e d t o e x em plify th e p r im a r y phy s i ol ogic a l

-
di fferenti a ti on o f ou t er p a rt s fro m i nn er p a rt s ; s o h ere it , ,
s u ppli es a s i m pl e ill u s t ra ti on of t h e w a y i n whi c h th e

di fferenti a t ed ou t er p a rt i s re di ff eren ti a t ed i n c orres pon
-
,
d enc e wi th th e chi ef c ont ra s t i n i ts rel a t i ons t o th e envi ron

m ent T he only m a rk ed u nlik enes s whi ch th e c ortic a l l a yer
.
o f t he P u ff b a ll p r e s en t s
-
,
i s th a t b et ween th e p orti on n ext
t h e g rou nd a nd th e o pp os it e p or ti on The b ett er d ev el op e d
.
-
F u ngi exhibit a m ore d ec id e d h et erogeneity of p a ra ll el ki nd .
S u c h i n c ru s ti ng A l g ee a s Ra l fs i a verru c osa fu r ni s h a ki n
d red c on t ra s t ; a nd i n the high er A l g ae it i s u ni formly
r ep ea t ed . P ha n oga m s di s pl a y thi s phys i ol ogi
c a l differ en ti a ti on v ery c on s pi cu ou s ly T h a t ea rth a n d a i r
.
a r e u n lik e p o rti on s o f t h e envi r on m ent s u bj ecti n g r oo t s a n d

,
l ea v es t o u nlik e phy s i ca l forces w hi c h ent a il on th em unlik e

,
r ea c ti on s a n d th a t th e u n lik e f u ncti on s a n d s t ru c tu res o f

,
t h ei r res p ec tiv e s u rf a c es a re fi t t ed t o th ese u nlik e ph y s i c a l

f orc es a re fa mili a r fa ct s whi ch it w ou ld be needl es s h ere
,
t o n a m e w ere it n ot th a t th ey m u s t be c ou n t e d a s c omi ng
,
w ithi n a wi der g rou p o f fa c t s .
I s thi s u nlik en es s b et ween th e ou t er ti s su es of th e a tt a ch e d

en d s a n d th o se o f t h e f ree en d s i n pl a nt s d et erm i ned by
,
t h ei r c onv ers e w ith th e u nlik e p a rt s o f th e en vi ronm ent ?

T h a t th ey r esu lt f ro m a n equ ilib r a ti on p a rtly a ri s i n g i n
th e i ndivid u a l a n d p a rtly a r i s i n g by th e su rviv a l of i ndi vi
d u a l s i n whi ch it ha s b een c a rr i ed fu rth es t is i nfera bl e ,
( i p ri ori ; a n d thi s a p ri ori a r g um ent m a y be a deq u a t ely
enfor c e d by a rgu m ent s o f th e i n d u ctiv e ord er A few .
t ypi c a l ones m u s t h ere s u ffi c e . T he g em m u l es

o f th e M a r cha n ti a a r e l ittl e di s c s h a p ed m a s s es o f cell s
-
c omp osed of tw o or m ore l a yers T h ei r S id es b ei ng a lik e

.
,
th ere i s n ot h i ng t o d et ermin e whi ch s id e fa ll s l owerm os t

wh en on e o f th em i s d et a ch ed Whi ch ev er s id e fa ll s l ower
.
m os t h owever p res ently b egi ns t o s end ou t rootl et s whil e

, , ,
T HE OU TE R T ISS U E S OF P L A NTS .
25 3
th e u pp erm os t s id e b egi n s t o a s s u m e th os e ch a ra ct er s whi ch

di s ti ngu i s h the fa ce of the f rond When thi s di fferen ti a ti on
.
h a s c omm ence d t h e t end ency t o i ts c ompl et e es t a bli s hm en t

,
b ec om es m ore a n d m ore d ecid ed ; a s i s p rov ed by th e f a ct

th a t if the p os iti ons of the s u rf a c es be a lt ered th e gem mu l e ,
b ends i ts elf s o a s to re a dj u s t th em : the c h a nge t owa rds

-
equ ilib r iu m w ith envi r on i ng f orces h a vin g b een once set u p ,
there is a cq u i red a s it were a n i nc rea s i ng m oment u m whi ch

, ,
res i s t s a ny co u nt er—ch a ng e Bu t th e e vid enc e S h ow s th a t

.
a t t h e ou t s et the rel a ti on s t o ea rth a nd a i r a l one d et er

,
mi ne the di fferenti a ti on of th e u nd er s u rfa ce from the

u pp er . T he e xp eri ences of t he ga r d ener m u lti ,
plyi ng h i s pl a nt s by cu tti n g s a n d l a yers c onstit u t e a n oth er,
cl a ss of evid ences not t o be om itt ed : th ey a re c om m onpla c e

bu t i ns t ru ctiv e e x a m pl es of phy s i ol ogi ca l differen ti a ti on .
Whil e c i rcu m s t a nce d a s it u su a lly i s the m eri s t e m a ti c ti ssu e

,
o f ea ch b ra nc h i n a P ha n og a m c on ti nu es t o p erfor m its
Ordi n a r y fu nc ti on— r eg u l a r ly p ro d u c i n g on its ou t er S id e the
c or ti ca l s u b s t a n ces a nd on i ts i nn er s id e th e v a scu l a r a nd
,
woody ti s s u es B u t ch a nge th e c on di ti ons t o th ose whi c h

.
t he u nd erg rou n d p a rt of th e pl a n t i s exp ose d t o a nd th ere ,
be gi ns a noth er di fferenti a ti on res u lti n g i n u n d erg rou n d S t ru e

t u res Cont a c t w ith wa t er oft en su ffi ces a l one t o p r od u ce
.
this res u lt a s i n the b ra nch es of s om e t ree s w hen th ey d roo p

,
i nt o a p ool or a s o cca s i ona lly with a cu tti ng pl a ced i n a

,
b ottl e of wa t er ; a nd wh en the light i s ex c lu d ed by i m

b eddi ng the end o f the cu t t i n g or th e middl e o f the s till
,
a tt a c h ed b ra nch i n th e ea rth thi s p rod u c ti on o f ti s s u es

, ,
a d a pt ed t o t h e fu n c ti on of a b s orbi n g m o i s t u re a nd mi nera l
c on s tit u e nt s p r ocee d s s till m ore rea dily With su c h c a se s

.
m a y be gr ou p ed th o s e i n which thi s d ev el o pm en t of u n d er
g rou nd orga n s by an a b ov e g rou n d ti s su e is not excep

-
,
t i on a l bu t h a bit u a l Creepi ng pl a nt s fu rni s h g oo d ill u s

.
t r a t i on s . F rom th e s h oot s of the Grou nd I vy rootl et s a re -

,
p t ou t i nt o the s oil i n a m a nner di fferi ng bu t littl e from

u
th a t i n which th ey a re pu t ou t by a n i m b edd ed l a yer ; s a ve

2 54 P H YS IOL OGI CA L D E VE L OP ME NT .
th a t the p roces s f oll ow s n a t u ra lly i n d u ced c onditi on s i ns t ea d -
o f f o ll ow i ng a rt i fi c i a l l y i nd u ced c onditi ons -

B u t i n t he .
com m on I vy w hi c h i ns t ea d o f ru nn i ng a l on g the s u rfa ce

,
o f th e ea r th r un s u p i n c li n ed or v ert i c a l s u rfa c es we s ee t h e
, ,
p roces s i nteres ti ngly m odified with ou t b ei ng es sen ti a lly

ch a nged T he rootl et s h ere di fferen ti a t ed by th ei r con
.
,
di ti on s i nt o org a n s of a tt a ch m en t m u c h m or e th a n org a ns of
a b s o r pti on s till d ev el op on t h a t S id e o f th e S h oot ne xt t h e
,
s u pp orti ng s u r fa c e a n d d o n ot d ev el op w h er e t h e S h oo t
, ,
g rowing a w a y fr om t he t ree or w a ll i s s u rr ou nded equ a lly ,
o n a l l s id es by light a n d a i r : th u s S h ow i ng u n d eni a bly , ,
th a t the p r odu cti on of the root l ets i s d eterm i ned by th e

di fferenti a l i ncid ence of for ces T h ou gh s u r viv a l o f t h e
.
fi t t es t d ou btl es s fu rth ered thi s t ra ns iti on yet it cl ea rly did

n ot i n iti a t e it . T h a t g r eenn es s which m a y be
o b serv e d i n th es e I vy b ra n ch r oo tl e t s whil e th ey a re qu it e
-
y oun g s oft a nd u n s h a d ed i nt rod u ces u s t o f a ct s whi ch a re

, , ,
th e c on v ers e of th e foreg oi n g fa ct s ; a nd p r ov es th a t th e p a rts

ordi na rily imb e dd ed i n th e s oil a n d a d a pt ed t o its a c ti on s ,
a cqu i re ,
o ft en i n v ery m a rk e d d eg rees the s u p erfici a l ,
st ru ctu res of the a e ri a l p a rt s wh en th ey a re e xp os ed t o light ,
a nd a i r T hi s m a y be w it n es sed i n Ma i ze which w h en
.
, ,
l u xu ri a nt sen d s ou t f rom i t s nod es nea r the gr ou n d c l u s t ers

, ,
o f roo ts th a t a r e thick s u c cu l ent a nd of th e s a m e c ol ou r a s

, ,
th e l ea v es E x a mpl es m ore fa m ili a r t o u s i n E ngl a nd occ u r

.
i n ev ery fi eld o f t u rnip s On noti ng h ow green i s the u n

.
c ov ered p a rt of a t u rnip root a nd h ow m an ifes tly th e -

,
a r ea o v er which th e gr eennes s e xt en d s v a ri es wi th th e a rea
e xp o se d t o light a s well a s wi th th e d eg ree of t h e exp os u re

, ,
it w ill be seen th a t b ey ond qu es ti on root tiss u e a s su m e s ,

-
t o a c ons id era bl e ext en t th e a pp ea r a nc es a n d f u n cti on o f

l ea f tis s u e w h en su bj e ct t o th e s a m e agenc i es
-
,
L et u s n ot .
fo rge t t oo th a t wh ere exp os ed root s d o not a pp roa ch i n

, ,
s u p erfici a l c h a r a ct er t owa r d s l ea v es th ey a pp r oa ch i n ,
s u p erfi c i a l c h a r a ct er t ow a rds s t em s : b ec omi n g cl oth e d

with a thi ck fi s su red b a rk lik e th a t of th e t r unk a nd
, ,
T HE OU T E R T IS SU E S OF P L A NT S .
255
b ra nch es . B u t the m os t c oncl u s iv e evid enc e i s

fu rni s h e d by th e a c t u a l s u b s tit u ti ons of s u r fa c e s t r u c t u res -
a nd fu nc t i ons t h a t oc cu r i n a er i a l o rg a n s whi c h h a v e t a k en
,
t o g ro wi n g p er m a nen tly u nd er g rou nd a n d i n u n d er g r ou nd ,

-
o rg a n s which h a v e t a k en t o gr owin g p er m a nen tly i n th e
a ir . On th e one h a nd th er e i s th e rh iz om e e x em plifi ed by
,
Gi nger— a s te m w hi c h i ns t ea d of sh ooti n g u p v ertic a lly

, ,
r un s h or iz on t a lly b el ow t he s u r fa ce o f t he s o il a n d a s s u m e s ,
the c h a r a c t er o f a root a lik e i n co l ou r t ext u re a n d

, , ,
p ro d u c ti on of rootl et s ; a nd th ere i s th a t ki nd of s w oll en

u n d er g r ou n d a xi s b e a r i n g a xill a r y b u d s w hi c h t h e P o t a t o
-
, ,
ex em p l i fi es— a s t ru ctu re w hi c h th ou gh h om ol ogic a lly a n ,
a xi s ,
s i m u l a t es a t u be r ou s roo t i n su rfa ce ch a ra ct er a nd -
,
w h en e xp os e d t o th e a i r m a n i fes ts n o g rea t er rea di nes s t o

,
d evel o p c hl orophyll th a n a t u b erou s r o ot d oes On the other .
h a nd there a re the a eri a l r oot s o f cert a i n Orchid s whi ch

, ,
h a bit u a lly g reen a t th ei r tip s con ti nu e gr een th rou gh ou t ,
th ei r w h ol e l ength s wh en k ept m oi s t ; w hich h a v e b ec om e

l ea f lik e not only by thi s d evel op m ent of c hl orophyll bu t
-
,
a l so by the a cqu i re m en t o f s t o m a t a ; a nd w hi c h do n ot b u r y
th em selv es i n the s oil wh en th ey h a v e the opp ort u nity *

.
T hu s we h a v e a eri a l org a n s s o c ompl etely ch a nged t o fit

u n d er gr ou n d a c ti on s t h a t th ey will n ot res u m e a eri a l f u nc
-
,
ti ons ; a nd u nd er gr ou nd org a ns s o com pl et ely ch a n ged t o

-
fit a e ri a l a cti ons th a t they will n ot r es u m e u nd er g rou n d

,
-
fu nc ti ons .
T h a t th e phy s i ol o gi c a l differ en ti a ti on b et ween th e p a rt Of

’
a pl a n t s s u r f a c e whi c h i s e xp os e d t o light a nd a i r a n d the
p a r t which i s e xp os ed t o d a rkn es s a nd m oi s t u re a nd s olid

A cri ti ca l co m m e nt m a de on thi s s en t ence r un s as fol l ow s The
a eri a l roots of m os t ep i ph yt i c orchids con ta i n ch l o roph yl l i n t h ei r co rt e x
th rou ghout t hei r l eng th, b ut th e cort e x be i ng v
co ered by a
‘
v el a m en ’
of a i r
conta i ni ng ce l l s whi ch b re ak up a nd re fle cti nci dent l i gh t , t h e gre en col ou r
i s not v i s i bl e th rou gh thi s opa q ue coa t . Wh en m oi s te ned th e c el l s of th e
v el a m en ta k e up w a te r a nd th e gre en co l ou r im m ed i a t el y s h ow s th ro ugh .
S uch root s do not h owe v er po s s e s s s tom a t a Th e . roo t s of c erta i n s p e ci es of
A ngr wc um ,
h ow ev er, conta in t h e wh ol e of th e a s si m i l a ti n g ti s s ue of th e
”
pl a nt .
25 6 P H Y S IOL OGICA L D E VE L OP ME NT .
m a tt er i s p rim a rily due t o th e u nlik e a ct i on s of thes e u nlike

,
p a rts of th e envi ronm ent i s th en c l ea r ly i m pli ed by ob s erve d

, , ,
f a c t s— m ore c l ea r ly i nd eed th a n wa s t o be exp ect ed C on

, ,
.
s i deri ng h ow s t ron g m u s t be th e i nh er i t e d t en d en c y of a pl a n t
t o a ssu m e t h os e S p eci a l ch a ra ct ers phy s i ol ogic a l a s well a s

,
m orph ol ogi ca l whi c h h a v e r es u lt ed from a n enor m ou s a ccu

,
m u l a t i on o f a n t ece d en t a cti on s it m a y be ev en th ou ght

,
s u rp ri s i ng th a t t hi s t end ency c a n be cou nt era ct e d t o s o g rea t

a n e xt en t by ch a ng ed con diti on s S u ch a d egree of m odi fi
.
a bili ty b ec om es c om p reh ens ibl e only w h en w e r em emb er

’
h ow littl e a pl a nt s f u n c ti ons a re i nt eg ra t e d a nd h ow m u ch , ,
th eref ore th e fun cti ons g oi n g on i n ea ch p a rt m a y be a ltered

,
w i t h ou t h a vi ng t o ov ercom e the m o m ent u m of the fu ncti on s

th rou gh ou t the wh ol e pl a nt B u t thi s m odi fi a bi l i ty be i ng a s
.
g rea t a s it i s we ca n h a v e no di fficu lty i n u nd ers t a n di ng

,
h ow by the cu m u l a tiv e a i d of n a t u ra l s el ecti on thi s p ri m a ry

, ,
di fferenti a ti on of the s u rf a ce in pl a nt s ha s b ec om e wh a t we
s ee it .
2 73 . We will l ea v e n ow th es e c ont ra s t s be t ween t he f ree

s u rfa ces of pl a n t s a nd thei r a tt a ch ed or i m b e dd e d su r fa ces ,
a nd t u rn ou r a tt en ti on t o th e se c on d a ry c on t ra s t s exi s ti ng
b et ween di fferen t p a rt s o f th ei r free s u r f a ces Were a f u ll .
s t a t em en t of th e e vid ence p ra cti ca bl e it w ou ld be p rop er ,
h ere t o d well on th a t whi ch i s f u rn i s h e d by th e i n feri or

cl a ss es I t might be p o i nt ed ou t i n d et a il th a t wh ere a s
.
,
a m on g th e A l g ee th e f ree s u r f a ces a re n ot dis s imil a rly c on

,
d i ti oned th ere is no s y s t em a tic differenti a ti on of th em — th a t

,
the f r ond of a n Ul va th e ribb on s h a p ed divi s i ons o f a

,
-
L a m i na ri a a nd the dich ot om ou s e xp a n s i on s of the F u ci

,
which cl oth e the r ock s b etween tid e m a rk s a re a lik e on b oth-

,
s id es ; bec a u s e s wa ve d a b ou t i n a l l di recti on s a s th ey a re by
,
th e w a v es a n d ti des th ei r s id es a re equ a l ly a ffect ed

, C on .
vers el y f r om th e F u n i m ight be d r a wn a b u n d a nt p r oo f th a t
, g
e v en a m ong T h a ll ophyt es u nl ik enes s es a ri s e b et ween di fferent
,
p a rts of the f ree su rfa c es wh en th ei r ci rc u mst a nc es a re unlik e .

TH E OU T E R T ISS UE S OF P L A N T S .
25 7
In h l a t era lly gr owi ng ki nds a s a re s h own i n Fig ; 1 9 6 6

su c -
,
t h e h oney co mbed u n d er s u rfa ce a nd th e s m ooth l ea t h ery

-
u pp er s u r f a ce h a v e th ei r c on t r a s ts r el a t e d t o con t r a st ed c on
,
di ti ons ; a nd i n th e a dj a c ently fi g ured A g a r ic s a nd oth er -

,
s t a lk e d gen er a the pil eu s e xhibit s a p a r a ll el di fferenc e ex

, ,
pli ca bl e i n a p a ra ll el wa y Bu t p a s s i n g over Crypt og a m s it

.
m u s t s u ffic e if w e ex a mi ne m ore a t l en gth th es e t ra it s a s th ey
a r e di s pl a y e d by P h a n og a m s L et u s fi rs t not e t he di s
.
si m il a r iti es bet ween the ou t er ti s s u es of st em s a nd l ea v es .
T h a t th es e di s s imi l a riti es a ros e by d egree s a s fa s t a s th e ,
u nit s o f w hi ch t h e pha n og a m i c a xis i s com p ose d b ec a m e
i n t egr a t ed i s a c onc l u s i on i n h a rm ony w ith the t ru t h t h a t i n

,
ev er y s h oot o f e v er y pl a n t , th ey a re a t fi r s t slight a n d b ec om e
g ra d u a lly m a rk ed A l rea dy i n b ri efly t ra ci ng the c ontra s t s

.
,
be t ween th e ou t er a nd i nner ti s su e s o f pl a n t s s ome fa ct s ,
h a ve been na m ed s h owi ng by impli ca ti on h ow the c es s a ti on

, ,
o f th e l ea f f u ncti on i n a x e s i s du e t o th a t c h a n g e of c on di
-
ti ons ent a il ed by the di s ch a rge of other fu ncti on s H ere .
w e h a v e t o c ons id er m ore cl os ely f a c ts of this cl a s s t ogeth er ,
with oth ers im m edi a t ely t o the p oi nt On p u lli ng .
Of f f rom a s t e m of gr a ss th e su cc es s i v e s h ea ths of its l ea v es

‘
the m or e i nc l os ed p a rt s of w hi c h a re of a f a i nt er g reen th a n
-
the ou t er p a rt s it wi ll be f ou n d th a t th e t u b u l a r a xi s ev en
,
tu a l l y rea ch e d i s of a still fa i nt er g reen ; bu t w h en th e a xi s

ev ent u a lly s h oots u p i nt o a fl ower i n g st em i t s exp o s ed p a r t ,
a cqu i r es a s b right a g reen a s th e l ea v es I n oth er M ono .
c otyl ed on s th e l ea f s h ea th s of whi c h a r e s u cces s iv ely b u rs t

,
-
a n d ex fo li a t e d by th e s welli ng a xi s it m a
y be ob s erv ed th a t
,
wh ere th e d ea d s h ea th s d o n ot m u ch ob s t ru c t t he light a n d
a i r th e su rfa c e o f th e a xi s u n d ern ea th i s fu ll o f c hl or ophyll
, ,
D endr o bi u m is a n ex a m pl e B u t wh en th e d ea d s h ea th s
.
a ccu m u l a t e i n t o a n op a qu e en v el op e t h e c hl or ophyll i s a b ,
s en t a n d a l s o w e m a
, y i nfer th e fu n ct i on whi ch i t s p resen ce
, ,
h a bit u a lly i m pli es C a rryi n g with u s thi s e vid ence we s hall

.
,
rec o gn i ze a lik e rel a ti on i n D i co tyl ed ons Whil e i ts ou t er .
l a y er rem a in s t olera bly tra nsp a rent a n ex ogenou s st em or ,
63
”
25 8 P H Y S IOL OGICA L D E V E L OP ME NT .
b ra n ch c onti nu es t o S h ow by th e form a ti on o f chl oroph y ll

, ,
th a t it s h a res i n the d u ti es of th e l ea ves ; bu t i n p rop ort i on

a s a b a r k w hi c h th e ligh t ca n n ot p ene t ra t e i s p ro d u c ed by
t h e a dh ere nt fl a k es of d ea d s ki n or by th e a c tu a l d ep os i t of
,
a p rot ec t iv e s u b s t a n ce the differen t i a t i on o f d u t i e s b ecom es

,
m ore d ecid ed . C a ct u s es a nd E u ph orbi a s s u ppl y

u s w i t h c on v ers e fa c t s h a vi n g t h e s a m e i m pli c a ti on T he .
s u ccu l ent a x es s o s t ra n g ely com bi ned i n th es e pl a nt s m a i n ,
t a i n for a l ong ti m e the t ra ns l u c ency of thei r ou t er l a yers

a nd th ei r g reenness a n d t h ey s o effi c i ently p erform t h e O f fic es
o f l ea v es t h a t l e a v e s a re n ot p r od u c ed I n s om e ca ses a x es
.
,
th a t a re not s u c cu l ent p a r ticip a t e l a rgely i n the l ea f fu ncti on -

,
or e nti rely u s u rp i t— s t ill ,

h owev er by f u lfilli ng the s a m e
,
e ss ent i a l con diti ons . Occa si ona lly a s i n S ta ti ce bra s s i ccefol i a

, ,
s t em s b eco m e f r i n ged ; a nd th e f ri nges th ey be a r a ssu m e ,
a l ong with th e thi nness o f l ea v e s t h ei r d a rk er green a nd

,
g enera l a s p ect . I n th e genu s Ru scus th e fl a tt ened a xis ,
s imu l a t es s o cl os ely t he l ea f s t ru c t u r e th a t w ere it n ot fo r th e

-
,
fl ower b orne on i ts m i d r ib or edge i ts a xi a l na t u re wou ld

, ,
h a rdly be s u sp ect ed A n d l et u s n ot omit t o n ot e th a t wh ere

.
a x es u s u rp th e ch a r a c t ers o f l ea v es i n th ei r a ttit u d e s a s w ell

,
a s i n th ei r s h a p es a n d t hi ckn es ses th er e a re c ont r a s t s b etween

,
th ei r u n d er a nd u pp er s u rfa ces a nsweri n g t o th e cont ra s t s

,
b et ween the rel a ti ons of th es e s u rf a ces t o the light Of this .
Ru s cus a n dr ogynu s fu r nish es a s t riki ng ex a mpl e I n i t th e .
difference whi c h th e u na id ed eye p erceiv es i s m u ch l es s con

S p i cu ou s t h a n th a t di scl osed by th e mic r o s cop e ; for I fi nd
th a t whil e th e fa c e of th e p seu d o l ea f ha s n o s t om a t a the b a ck

-
,
i s a b u n d a n tly s u ppli ed w ith th em One m ore ill u s t ra ti on

.
mu s t be a dd ed E qu a lly for th e m orph ol ogi c a l a nd phy s i o

.
l ogi ca l t ru th s which i t en forces the M ii hl en beckia p l a tycl a da

,
i s one of th e m os t i n s t ru c tiv e of pl a n t s I n it the s i m u l a ti on

.
o f f or m s a nd u su r p a ti on o f f u nc ti on s a r e c a rr i ed ou t in a ,
m u c h m ore m a rv ell ou s w a y th a n a m ong th e Ca c ta cece .
I m a gin e a gr owth resembli ng i n ou tli ne a v ery l ong will ow

l ea f bu t with ou t a mid rib a n d h a vi ng i t s two s u rfa ce s a l i k e
, ,
.
T HE OU T E R T ISS U E S OF P L A NTS .
259
I m a gi ne th a t thi s thi n g reen s em i t ra nsp a ren t st ru e

a cro s s , ,
-
t u re th ere a re f rom ten t o thi rty divi s i ons w h ich p rov e t o be

, ,
t he su c c es s iv e n od es of a n a xi s I m a gi ne th a t a l ong the
.
e dges o f t hi s l ea f s h a p e d a gg re g a t e of i n t ern o d es th ere a ri s e

-
,
a xill a ry b u d s s o m e o f whi c h un fo ld i nt o fl ow ers a n d o th ers

, ,
o f whi c h S h oot u p v ertic a lly i nt o g r owt h s lik e the one w hi c h
b ea rs th em I m a gi ne a w h ol e pl a nt thu s s eem i ngly c omp os ed

.
o f j o i nt e d will ow l ea v es g r o-
wi g
n f ro m o n e n r
’
a oth e s edg es ,
a n d s o m e con c epti on will be f orm e d o f the Mil hl en bec ki a .
T h e t wo fa c t s whi c h h a v e m ea ni ng for u s h ere a ra — fi rs t th a t ,
t h e p er form a nc e o f l ea f fu ncti ons by t h es e a x es g oes a l on g

-
w ith the a s su mp t i on o f a l ea f lik e t ra ns l u cenc y a nd s ec on d

-
, ,
th a t th ese fl a t t ened a x es r eta i ni n g th ei r u p right a ttit u des

, ,
a n d th er efo re k eepi n g th ei r tw o f a c es s imil a rly c on diti oned ,
h a ve th es e t w o f a c es a lik e in c ol ou r a nd t e xtu re .
T h a t phy s i ol ogi ca l di fferent i a ti on o f t h e s u r f a c e which

a r i ses i n P h a n og a m s b et ween a xi a l o rg a n s a nd f oli a r or g a n s ,
i s thu s t ra cea bl e w i t h t ol era bl e cl ea rnes s t o th os e di ff eren ce s

b et ween th ei r con diti ons w hi ch i ntegra ti on h a s en t a il ed
p a rt ly i n th e wa y a b ov e d es c rib ed a nd p a rtly in other wa ys
s t ill t o be n a m ed By i t s rel a t iv e p os iti on a s b ei n g sh a d ed
.
,
by th e l ea ves the a xi s i s l es s f a v ou ra bly c i r c u m s t a n ced for

,
-
p er formi ng th ose a s s i m il a tiv e a cti ons effect e d by the a i d of

light F u rther th a t rel a tiv ely s m a ll ra ti o of s u rf a ce to m a s s
.
,
-
i n the a xi s which i s neces s it a t ed by i ts f u ncti ons a s a s u pp ort

,
a n d a c h a nnel for c i rcu l a ti on p rev ent s it f rom t a ki ng i n

, ,
w i th th e s a m e f a c ili t y a s th e l ea v es th os e s u r rou nd i n g g a ses

,
fro m whi c h m a tt er i s t o be a s s imil a t ed B oth th es e S p eci a l.
ca u s es h owev er i n c omm on with th a t p revi ou s ly a s s igned

, , ,
fa ll withi n th e genera l c a u s e A n d i n t h e f a c t th a t wh ere

.
th e di ffer en ti a l c on diti on s do n ot exi s t the phy s i ol o gi ca l ,
di fferen ti a t i on d oes not a r i s e or i s oblit era t ed we h a v e cl ea r

, ,
p r oof th a t it i s det erm i ned by u nlik eness es i n the rel a ti ons

of th e p a rt s t o th e en vi r on m en t .
§ 2 74 . F rom thi s m os t g enera l c ont ra s t b etween i l

a er a
2 60 P H Y SIOL OGI C A L D E V E L OP ME NT .
f c ti s su es— th os e o f a x es a n d th os e of foli a— we p a s s
s u r a e-
n ow t o t h e m ore s p eci a l c on t ra s t s o f lik e ki n d e xi s ti ng i n
foli a t h em se lv es L ea v es p res ent u s with su p er fi c i a l d ifferen

.
t i a t i on s of st ru ct u re a n d fun ct i on ; a nd we h a v e t o c on s id er
t h e rel a ti ons b et ween th es e a n d th e en vi r oni n g f orc es .
Ov er the w h ol e s u r fa c e of ev ery pha nog a m i c l ea f a s ov er ,
th e fron d s of the P teri dop hyta there ext en d s a si m pl e or

,
c om p ou n d epid er m a l l a y er f orm ed of cells th a t a re c l osely

,
u n i ted a t th ei r e dge s a n d d ev o id ( i n t h e Fl ower in g P l a nt s )
o f t h a t g ra nu l a r col ou r i ng m a tt er ( chl or ophyll ) con t a i ne d i n
th e l a y ers of cell s t h ey i nc l os e : t he res u l t b ei ng t h a t t he

m em b ra n e form e d o f th em is c om p a ra t iv ely t ra ns p a rent .
On the s u b m er ge d l ea v es of a qu a tic P h a n og a m s t hi s ou t er ,
l a y er is thi n d eli ca t e a nd p erm ea bl e by w a t er ; bu t on

, ,
l ea v es exp os ed t o the a i r a nd es p eci a lly on th ei r u pp er s u r

,
f a ces i s c om p a r a tively s t rong d en s e oft en s m ooth a n d

, , ,
i m p erm ea bl e by wa t er : b ei ng thu s fi tt ed t o p rev ent the

r a pid es ca pe o f th e c on t a i n ed j u i ce s by ev a p ora ti on S .
l a rl y whil e the l ea v e s of t erres t ri a l pl a nt s whi c h liv e i n tem

,
p era t e clim a t es u s u a ll y h a v e c omp a ra tiv ely thi n c oa ts thu s

,
c om p os ed i n c lim a t e s th a t a re b oth h ot a nd dry l ea v es a re

, ,
c om m on ly cl o th ed wi th a v ery thi c k c u ti cl e N or i s thi s a l l

. .
T he ou t s id e of a n a eri a l l ea f di ffer s from th a t o f a s u bm erg ed

l ea f by c on t a i ni ng a d ep os it of wa xy s u b s t a nce Wheth er .
thi s be ex u d ed by th e exp osed s u rfa ces of the cell s a s s om e ,
c ont en d o r w h et h er it i s d e p os it e d wi t hi n t he c ell s a s th ou ght

, ,
by oth ers m a t ters not i n s o fa r a s the genera l res u lt i s con

,
c erned I n e ith er c a s e a w a t erp r oof c oa ti ng i s f orm ed a t the

.
o u t er m os t s id es o f th ese ou t erm os t c ell s ; a nd i n m a ny c a ses
p r od u ces t h a t p oli s h by whi ch th e u pp er s u rfa ce o f the l ea f is

m o r e or l es s di s ti ngu i s h ed f r o m t he u n d er s u r f a c e T hi s
.
e xt er na l p elli c l e p r es en t s u s with a n o th er c ont r a st of a lli e d
m ea n in g On the u pp er s u r fa c es of l ea v es s u bj ect t o th e
.
di rec t a c ti on of th e s u n s ra y s th ere a re eith er few or n one

’
,
o f th os e mi nu t e o p en i n g s or s t om a t a
,
th r ou gh wh ich g a s es
,
c a n ent er or es c a p e ; bu t on the u n d er su r f a c es th es e st om a t a
T HE O UT E R T IS S U E S OF P L A NT S .
261
a re abu nd a nt : a di s t ribu ti on whi c h while perm itti ng f ree

,
a b s or pti on of th e nee d fu l c a rb on ic a c id p u ts a c h eck o n t h e

,
e xit o f wa t ery v a p ou r . T wo genera l ex cepti ons t o thi s a r

r a nge m en t m a y be no t ed .L ea v es th a t fl oa t on th e w a t er
h a v e a l l th ei r s t om a t a on thei r u pp er s id es a nd l ea v es th a t
,
a r e s u b m erg ed h a v e no s t om a t a — m odi fi ca ti on s o bvi ou s ly a p
p rop ri a t e t o th e c on diti on s . W h a t i s t o be s a id
res pec t i n g th e g enes i s o f t h es e differe nti a ti ons ? F or the

l a s t there s ee m s no di rect c a u s e : i ts ca u s e mu s t be i n di rect .
T he u nlik e a cti ons t o w hich t he u pp er a n d u nd er s u rfa c es o f

l ea v es a re s u bj ect h a ve no a pp a ren t t end ency to p rodri ce
,
u n lik e nes s in t h e nu m b er of th ei r b rea thi ng h ol es H ere .
the na t u ra l s el ect i on o f sp ont a n eou s v a ri a ti on s fu rn i s h es the

o nly fea s ibl e expl a na ti on . F or th e fi rs t h owev er th ere i s a
, ,
'
p os sibl e c a u s e i n the i m m edi a t e a cti ons of i nc id en t f o rces ,
whi ch s u rviv a l of the fitt es t conti nu a lly f u rth ers .
T he fl ui d exh a li ng th rou gh the w a ll s of th e cell s next the

air ,
will be lik ely t o l ea ve be hi nd s u s p end ed s u b s t a nces on
thei r ou t er s u r f a c es On rem emberi ng th e p ellicl e w hi c h i s a pt
.
t o f orm on thi c k s ol u ti on s or e mu l s i on s a s th ey dry a n d h ow ,
t hi s p elli c l e a s it g row s re t a r d s th e f u r th er d ryi ng it w ill be ,
p erceiv ed th a t th e d ep os it of w a xy m a tt er next t o the ou t er

s u rfa ces o f t he cu ti c u l a r ce ll s i n l eav es i s n ot i m p r ob a bly
,
i niti a t ed by the ev a p ora ti on whi ch it ev ent u a lly ch ecks

‘
.
.
S h ou ld it be s o th ere resu lt s a v ery s i m pl e ca s e o f equ ilib ra

,
ti on Wh ere the l oss of w a t er i s t oo gr ea t thi s w a xy p ellicl e

.
,
l eft behi nd by the esca pi ng w a t er will p rot ect m os t th os e i n

di vi du al s of th e s p eci es i n whi ch it i s thi c k es t or d enses t ; a nd
by inh er it a nce a n d c onti nu a l s u rviv a l of th e fitt es t th ere w ill ,
be es t a bli s h ed i n the s pe c i es th a t thick ness of th e l a y er w hi c h

b ri ngs the ev a p ora ti on to a b a l a nc e wi th the s u pply of wa t er .
A noth er s u p er fi c i a l differenti a ti on s till m ore f a m ili a r h a s

, ,
t o be n ot ed ,
E v ery chi ld s oon l ea rn s t o di s ti ngu i s h by it s
c ol ou r the u pper sid e o f a l ea f f ro m its u nder S id e if the l ea f ,
i s one th a t h a s grow n i n s u ch w a y a s t o es t a bli s h the rel a

ti ons of u pp er a nd und er T he u pp er s u rf aces of l ea ves a re
.
2 62 ,
P H YSIOL OGICA L D E V E L OP ME NT .
ha bit u a lly of a deep er g reen th a n the u nd er Mic rosc opic . .
e x a mi na ti on s h ows th a t t hi s d ee p er g reen resu l t s f r om the

cl os er cl u st eri ng of th os e p a renchy m a cell s fu ll o f chl orophyll -
th a t a re i n s om e way concern ed wi th the a s s imil a tiv e a c ti ons ;

‘
whil e b enea th th em a re m ore nu m er ou s i nt erc ell u l a r p a ssa ges

c om mu nic a ti ng wi t h th os e Op eni ng s or s t om a t a th rou gh
which is a b s or b ed t he n eedf u l ai r N ow wh en it i s rem em .
bered th a t the f orm a ti on of chl orophyll i s cl ea rly t ra cea bl e t o

the a cti on of light— wh en it i s rem em b ered th a t l ea v es a re p a l e
where th ey a re mu ch sh a d ed a n d col ou rl es s wh en d evel op ed
i n the d a rk a s i n the h ea rt of a Ca bb a ge— wh en it i s rem em
,
b ered th a t s u c cu l ent a x es a nd p eti ol es lik e th ose o f S ea k a l e ,

-
a nd Cel ery rem a i n w hit e w hil e th e light i s k ept fro m th em

,
a nd b ec om e g reen w h en e xp ose d ; it c a n n ot be qu e s ti one d
tha t t hi s g rea t er p rod u cti on o f chl orophyll next t o the u pp er

s u r fa c e of a l ea f i s di rectly c ons equ ent on the g rea t er
,
a m ou n t o f light rec eiv ed H ere a s i n s o m a n y o th er c a s es

.
, ,
we m u s t reg a rd the diff erenti a ti on a s i n p a r t du e t o di rect

equi lib r a ti on a n d i n p a rt t o i ndi r ect eq u ilib ra ti on F a m ili a r .
fa cts c omp el u s t o c on cl u d e th a t from the b egi nni ng ea ch ,
i ndivid ua l f oli a r org a n h a s u nd ergone a cert a i n imm edi a t e

a d a pt a ti on o f i ts s u r f a c e s t o th e i n c id en c e o f light
; th a t
wh en th ere a ros e a m od e o f gr owth whi ch e xp osed the l ea ves
o f s u cc ess iv e g enera ti on s i n simil a r wa y s thi s imm edi a t ely ,
p rod u c ed a d a pt a ti on ever t end in g to be t ra ns m itt ed w a s

, ,
f u rth ered bv the su rviva l of i ndivid u a l s i nh eritin g it i n the

g rea t es t d egr ee ; a nd th a t s o th ere wa s g ra d u a lly es t a bli s h ed
th a t di fference b etween the tw o s u r fa c es w hi ch ea ch l ea f di s
pl a y s b e f ore i t u nf olds t o the light bu t whi ch b ecom es m ore ,
m a rk ed wh en it h as u n f old ed *
.
The c urrent doc tri ne th at chl oroph yl l

i s the s pe ci a l s ub s t a nce c oncerned
in v egetal as s i m i l a ti on, e i th er a s a n agent o r as a n i nc identa l p rod uc t , m us t
be t a ken wi th con s idera bl e q ual i fi cat ion . Be s i des t he fa ct th a t a m ong the

A lg w th ere a re m any re d a nd b rown kinds whi ch th ri v e ; a nd b esi de s th e
fact th at am ong th e l ower A rch e goni a te s t h ere a re s pe ci es whi ch a re p urp l e or
choc ol ate-col oure d ; the re i s th e fa ct th at P ha nog am s a re not all g reen . We
ha v e the v
Copper-Beech, w e h a e th e bl ack-p urpl e Col eu s Verschq fieI/u ,
'
a nd
flou i sh
-
we h a v e the re d va ri ety Of Ca bbage, whi ch s ee m s to r as wel l as th e

TH E O U T E R TI SS U E S OF P L A NT S .
2 63
F ro m the di na ry ca s es l et u s now p a s s t o the exc epti ona l

or
c a ses We w ill l ook fi rs t a t th ose i n wh i ch the t wo fa ce s of

.
t he l ea v es di ffer bu t littl e or not a t a l l — th ei r ci rc u m s t a nc es

,
bei ng s imil a r o equ a l L ea ves th a t g row i n a pp ro xim a t ely

r
.
u p ri ght a t t i t u d es a n d a tt it u d es whi c h d o n ot m a i n t a i n th e
,
rel a tiv e p o s iti ons o f t h e t w o s u r f a ces with c on s t a n cy m a y be ,
e xpe c t ed t o di s pl a y a n u n u su a l lik enes s b et w een th e tw o
s u r f a ces ; a nd a m on g t h em we s ee it T he Gra s s es m a y be .
na m ed a s a grou p ex empli fyi n g thi s rel a ti on ; a n d if i n s t ea d ,
of com p a r in g th e m a s a gr ou p with oth er g r ou p s w e c om p a r e ,
th os e d wa rf ki nd s of th em which s p rea d ou t th ei r l ea v es
h oriz ont a lly with the l a rge a s pi ri ng ki nd s s u ch a s A ru nd o
, , ,
w e t ra c e a lik e a n tith es i s : i n th e on e the cont ra s t o f u pp er

a nd un d er i s v ery o bvi ou s w hil e i n t h e o th er it i s s ca rcely t o
,
be d et ect ed L ea v es of v a ri ou s o th er Monoc otyl ed ons th a t

.
gr ow i n a s i m il a r w a y S imil a r ly S h ow u s a nea r a pp roa ch t o

,
u ni f ormity o f t he t w o s u r f a c es ; a s i ns t a nc e t h e g enu s Ol i vi a
a nd th e thi nn er l e a v e d ki nd s o f Yu cca
-
Wh ere the con .
t ra s t of Upper a nd u nd er i s grea tly di m i nished by the a s su m p

ti on of a roun d ed or cyli n d ri ca l f or m i ns t ea d of a fl a tt ened ,
form the s a m e thi n g h a pp en s

,
The g enu s K l ei n ia fu rn i s h es
.
illu st ra ti on s I t m a y be rem a rk ed t oo th a t ev en w ithi n

.
, ,
the limit s o f thi s g enu s th ere a re i n s t ru ctiv e v a ri a t i on s ; for

whil e i n K l ai ni a ficoi des the l ea v es s h a p ed lik e pea p ods ,
-
,
a re b roa d es t i n a v ertic a l di rec ti on a nd h a v e th ei r l a t era l

,
su rfa ces a lik e i n c on diti ons a n d st ru ct u r e i n ot h er s p e ci es ,
the l ea v es b r oa d er h ori z ont a lly th a n v ertic a lly exhibit

, ,
u n lik eness b et w ee n th e u pp er a n d u n d er sid es E qu a ll y .
t o th e p oi nt i s th e e vi d enc e fu r ni s h ed by v ertic a lly g rowi n g -
l ea v es th a t a re cyli nd ric a l a s th os e of S a n s evi era cyl i ndri ca

, ,
or a s th os e o f th e Ru s h —t r ib e : th e S imil a rly pl a c ed s u r f a c e -
h a s a l l a rou n d a s i m il a r c h a ra ct er Of ki nd red m ea n i n g
.
,
oth er v a ri e ti e s Chl orophyl l th en m u s t be re ga rded s i m pl y a s th e m os t

.
, ,
g enera l of th e c ol ouri ng m a tt ers fou nd i n th ose p a rts o f pl a n ts i n wh i ch

a s si m il a ti on i s b e i ng fect ed
ef bv th e a g ency of l i gh t Though i t i s al w a ys
.
p re s ent a l ong wi th the re d a nd b rown pi gm ent s , ye t th ere i s m u ch e v i dence to

s how t ha t th es e a re th e act ua a s s im i l a t i v elpi gm ents .
P H Y S I OL OGICA L D E VE L OP ME NT
'
2 64 .
a nd s till m ore c on cl u s iv e a re the c a s es i n which the u nd er

,
s id e o f th e l ea f b ei n g m ore e xp os e d t o light th a n th e u pp er
,
S id e ,
u s u rp s t h e ch a ra ct er a nd fu n cti on Of th e u pp er s id e .
I f a c om m on Fl a g be p u ll e d t o pi eces it will be s ee n t h a t ,
w h a t a ns wers t o t he f a ce i n ot h er l ea v es f orms m erely the ,
i n s id e of t he s h ea th i nclu din g the y ou nger l ea v es a nd i s ,
o bli t er a t e d high e r u
p T he tw o su rf a ces of th e bl a d e a ns wer
.
t o th e t wo u n d er h a lv es o f a l ea f th a t h a s b een a s it w ere , ,
f old ed t ogeth er l engthwa y s w i th the two h a lv es of its u pp er

,
s u rfa ce i n cont a ct A nd h ere i n d ef a u lt of a n u pp er s u rf a c e

.
, ,
t he u n d er s u rf a c e a c qu i res i t s ch a ra ct er a n d di s c h a rges its

f u nc ti on A lik e s u b s tit u ti on occu rs i n A ri s tea c orym bos a ;
.
a nd th ere a re s om e o f t he Orc hid s a s L ockha r tia whi c h dis ,

.
,
pl a y i t i n a v ery obvi ou s w a y .
When j oi ned with the foreg o ing evid ence th e evid ence ,
whi ch a n oth er ki nd of s ub s tit u ti on su ppli es i s of g rea t

w eight . I r efer t o th a t whi ch occu r s i n the A u s t ra li a n
A ca c i a s a l rea dy i ns t a nc e d a s th row i n g light on m orph o
,
l ogi ca l ch a ng es I n th es e pl a nt s the l ea v es p rop erly S O c a ll ed

.
a re u nd ev el op ed a n d th e f ootst a lk s fl a tt ene d ou t i nt o f o li a
, ,
ceou s s h a p es a c q u i r e v ei n s a nd mid r ib s a nd s o fa r s im u l a t e
, ,
l ea v es a s ordi na rily t o be t a k en for th em : a f a c t i n it s elf of

mu ch phy s i ol ogi ca l s ignific a nc e Bu t t h a t whi ch it c oncern s
.
u s e s p ec i a lly t o no t e i s t h e a b se nce o f di s ti ncti on bet ween

,
t he t w o f a ces of t h es e phyll od es a s th ey a re na m ed a nd , ,
th e c a u se of i ts a b sen c e T h es e t ra ns f orm ed p eti ol es do not

.
fl a tt en th em s elv es ou t h ori zont a lly s o a s to a cq u i re u n d er ,
a n d u pp e r sid es a s m os t t ru e l ea v es do ; bu t th ey fl a t t en
,
th em selv es ou t v erti ca lly : the res u lt bei ng th a t th ei r two

sid es a re si m il a rly ci rcu mst a nc ed with res p ect to light a nd
o th er a genc i es ; a nd th ere i s c ons eq u ently nothi n g t o c a u s e
th ei r differenti a ti on A nd th en we find a n a na l og ou s c a s e
.
where differenti a l c onditi ons a ri s e and wh ere s om e di ff er ,
enti a ti on re s u lts I n Oxa l i s bupl euri fol i a Fig 6 6 th ere is a

.
,
.
,
simil a r fl a tt eni ng ou t of the p eti ol e i nt o a p s eu d o l ea f ; but -
in it the fl a tt ening t a k es pl a c e i n the s a m e pl a ne a s the l ea f,

T HE OU T E R T ISS UE S OF P L A NTS .
2 65
so a s to p rod u ce d er a n d a n u pp er s u r f a ce ; a n d h ere
an u n
th e tw o s u rfa c es of t h e p s eu d o l ea f a r e s ligh t ly unl i k e — i n

-
c ont ou r if in not hi ng el se .
2 75 . We now c om e to ch phys i o l ogic a l d ifferenti a ti ons

su
a m ong the ou t er ti s s ue s of pl a n t s a s a r e di s pl a y ed in t h e
,
c ont ra s t s b et wee n f oli a r org a ns on the s a m e a xi s or on ,
di fferent a x es — c ontra s ts b etw een t he s ee d l ea v es a n d t he -
l ea v es s u b s equ ently f orm ed b et wee n s u bm erge d a n d a eri a l

,
le a v es i n c ert a i n a qu a ti c pl a n ts b et w ee n l ea v es a nd b ra c t s
, ,
a n d b et wee n b r a ct s a n d s ep a l s T o d ea l ev en b r i efly w ith

.
th ese impli es i n form a ti on whi c h ev en a p r ofes s ed b ota ni s t

w ou ld h a v e t o i nc rea s e by s p ec i a l i nqu i ri es b e f ore a t t em pti ng ,
i nt erp ret a ti ons H ere it mu st s u ffi ce t o s a y s om ethi n g

.
res pe ctin g th ose m a rk e d u n lik enes ses e xi s ti n g b e t w ee n t he
ti s s u es of th e m ore c h a ra ct eri s ti c p a r t s of fl owers a n d the ,
tis su es of the h om ol og ou s f oli a r orga ns .
I t wa s p oi nt ed ou t i n 1 9 6 th a t the t ermi n a l f o li a o f a
,
p h a n o a m i c a xi s h a v e s u n d ry ch a r a ct e rs i n c om m on w ith su c h
g
f rond s a s t h os e ou t of whi ch w e c onc l u d ed th a t the ph a n o
g a m ic a xi s ha s a ri sen by i nt eg ra ti on— c om m on ch a ra ct ers of
a ki n d t o be e xp ec t e d I n th ei r s impl e cell u l a r c om p os iti on
.
,
c om p a ra tiv e wa nt of chl orophyll a n d d efi c i ency of v a scu l a r

,
s t ru c t u re s the u n d ev el op ed en d s o f l ea f S h ee t s a nd th e de
,
-
vel oped en d s o f fl ower s h oot s a pp r oa c h t o t h e f ro n d s o f the

-
,
s impl er A rch eg on i a t es We a l s o not e d b et ween th em a noth er

.
resem bl a nc e I t is s a id of th e J u nger m a nni a cece th a t

.
,
“
t h ou gh u nd er cert a i n ci rc u m s t a nces of a p u re g reen th ev ,
a re i ncli n ed t o be s h a d e d w ith red p u r l e c h oc ol a t e o r ot h er

p , , ,
ti nt s ; a nd a n s w eri n g t o thi s we h a v e th e fa c t s th a t su c h
c ol ou rs c omm on ly oc cu r i n the t er m i n a l f o li a o f a ph a n o
ga m ic a xis wh en a rres t of i ts d ev el opm ent l ea d s t o the

,
f orm a ti on of a fl ow er a nd th a t v ery f requ ently th ey a re

,
vi s ibl e a t the ends of l ea f a x es I n the un f oldi ng p a rt s of

-
.
sh oot s m ore or l es s of red or c opp er c ol ou r or ch oc ol a t e

, ,
-
,
c ol ou r, m a y g enera lly be s een : oft en, i n d eed , it ch a ra c

2 66 P H Y SIOL OGICA L D E V E L OP ME NT .
teriz es th e l ea ves for s om e ti m e a ft er they a re u nf old ed .
O cc a s i ona lly the t ra ce s of i t a re p erm a nent ; a nd a s i n ,
t he s c a rl et t erm i na l l ea v es o f P oi ns et ti a p u l c herri m a w e see ,
th a t it m a y b ecom e a nd c onti nu e ex t rem ely c ons pi cu ou s

, , .
T h e qu es ti on then n e w t o be a s k ed i s— h a s thi s c ol ou ri ng
, ,
by which the im m a t u re p a rt of the pha nog a m i c a xi s i s cha

r a c teri ze d ,
a nythi ng t o do w ith t he c o l ou r i n g o f fl ow ers ?
H a s thi s di fl erence b et ween u n d ev el op e d f oli a a nd foli a t h a t

a re fu rth er d ev el op e d been i ncrea s ed by na tu ra l s el ecti on
,
where a n a dva nt a ge a cc ru e d f r om it u ntil it h a s end ed in

,
the s t ron g c ont ra s t we n ow s ee ? I thi nk we m a y not i rra

ti on a l l y i nfer th a t thi s h a s ha pp ened .
F a ct s v ery nu m erou s a nd v a ri ed u nit ed t o wa rr a nt u s i n

, ,
c oncl u ding th a t g a m ogenes i s comm ences wh ere th e f orc es

whi ch c ond u c e t o growth a re nea rly eq u ilib ra t ed by the forces
whi ch res i s t growth 7 8 ) a nd t he i ndu ct i on th a t i n pl a nt s ,
f ertili zed g erm s a re p rod u ced a t pl a ces w here th ere i s a n

a pp r oa ch t owa r d s this b a l a n ce w e fou n d t o be i n h a r m on y
,
with the dedu cti on th a t an a dv a nt a ge t o the sp eci es mu st be

ga i ned by s endi ng off migra ti ng p rogeny f rom p oi nts where
nu t r iti on i s f a ili n g
. Oth er thi ngs equ a l f a il u re of nu t riti on
,
m a y be exp e ct e d i n p a rt s w hich h a v e th e m os t rem ot e or m os t
i ndi rec t a cces s t o th e m a t eri a l s fu rni sh ed by the r oot s

m a t eri a l s whi ch ha v e to be ca rr i ed g rea t di s t a nc es by a v ery
i m p er fect a pp a ra tu s . The e nds of l a t era l a x es a re th eref ore
the p rob a bl e p oi n t s of fructi fica ti on i n a ggrega t es of the
,
thi rd ord er th a t h a v e t a k en t o g rowi ng v erti ca lly But i f .
th es e p oint s a t whi c h nu t riti on i s f a ili ng a re a l s o the p oi nt s

,
a t whi c h th e c ol ou rs i nh erit ed f ro m l ower typ es a re lik ely t o
rec u r i n m ore m a r k e d d eg ree s th a n el sew h ere ; th en we m a y
i nfer th a t the organ s of fru cti fica ti on w ill not u nfrequently

c o exi s t with s u ch c ol ou r s a t t h e end s o f s u ch a x es
-
H ow .
m a y the res u lti ng c ont ra s t b et ween the old er f ronds a n d th e
f rond s next the ger m p rod u ci ng org ans be i nc rea s ed ? I f

-
u nin terfered with it w ou ld be lik ely t o dimi nish T h es e

.
t ra its i nh erit ed f rom rem ot e a nc est ry might be exp ec t ed

TH E OU T E R TIS S U E S OF P L A NTS .
l wly t o f a d e a w a y H ow th en i s the i n t ens ific a ti on of

s o .
, ,
the m t o be expl a i ned ?

I f a cont ra s t o f the ki nd d es c rib ed fa v ou r s the p rop a ga ti on
o f a ra c e i n which it exi s t s it will be m a i n t a i ned a nd
,
i nc rea s ed ; a nd i f we t a k e i nto a cc ou nt a n a gency of which

Mr D a rwi n h a s s h own the grea t i m p ort a n ce— the a gen cy of
.
i n s ect s— we s h a ll h a v e littl e d iffi cu lty i n u n d ers t a n di ng h ow

s u c h a c on t ra s t m a y fa cilit a t e p r op a g a ti on We ca nnot of .
,
cou r s e here a s su m e th e a gency of i n s ect s s o s p ec i a lized in

,
th ei r ha bit s a s B ee s a nd Bu tter fli es ; for th ei r s peci a lized

h a bit s i m ply the pre—e xi s t ence of th e c ontra s t to be expl a i ned .
Bu t th ere is a n i n sect a gen c y of a m ore gener a l ki nd w hi c h

-
m a y be fa i r ly cou nt ed u p o n a s com in g i n t o a cti on Va ri ou s .
s m a ll Fli es a nd Beetl es wa nd er over the su r fa c es o f pl a nt s i n

s ea rch of f oo d I t i s a l egi t im a t e a s s u m pti on th a t they will
.
f req u ent m ost th os e p a rt s i n whi c h th ey fi nd m os t f ood or ,
f ood m os t t o thei r l iki n g— e s p ec i a lly if a t the s a m e ti m e

t h ey g a i n the a dv a nt a ge of con cea l m en t N ow the end s of
.
a x e s fo rm ed of y ou n g s o ft a nd cl ose ly p a c k e d f o li a a re th e
, , ,
-
,
p a rt s which m or e th a n a ny others offer th es e s ev era l a dva n

t a ges T h ey a ffor d s h elt er from en emi es ; th ey f requ en tly
.
c ont a i n ex u d ed j u ic es ; a nd wh en they do not th ei r ti s s u es ,
a r e s o t en d er a s t o be ea s ily pi er ce d i n s ea r c h o f t h e s a p .
I f th en f ro m the fi rs t a s a t p res ent th ese end s of a x es

, , , ,
h a v e been f a v ou rit e h a u nt s of s m a ll i n s ect s ; a n d i f where ,
th e c l osely c l u s t er ed f oli a con t a i ned th e g en er a tiv e org a n s

-
,
th e i n s ec t s f requ en ti n g th em oc c a s i ona lly ca rri e d a dh eren t

fr u cti fyi n g cell s fr om one pl a nt t o a noth er a nd s o a id ed ,
fertili z a t i on ; it wou ld f oll ow th a t a nything whi c h m a d e

s u ch t erm i n a l cl u s t er s m or e a tt r a c tiv e t o s u c h i n sect s or ,
m ore c ons pi cu ou s t o th em or b oth wou ld fu rth er the mu lti

, ,
pli ca ti on of th e ra ce a nd w ou ld s o be conti nu a lly i n crea sed

,
by the ext r a mu l t ipli ca ti on o f i ndi vi dpa l s i n wh i ch it w a s

grea t es t H ere we fi nd th e clu e T his c ont r a s t of c ol ou r
. .
b et ween th e f oli a next to the f ru ctifyi n g p a rts a n d a l l oth er

foli a mu s t c on s t a ntly h a v e f a cilit a t ed i ns ec t a gen cy ; s u p
,
-
2 68 P H YS I OL OGI CAL D E VE L OP ME NT .
p os ing th e i ns ec ts t o h a v e h a d th e p ower of di s t i ngu i s hi ng

b etween col ou rs T h a t Bees a n d Bu tterfli es h a v e thi s p ower
.
is m a nifes t .T h ey m a y be wa t ch ed flyi n g fro m fl ow er t o

fl ower di sreg a rdi ng a l l oth er p a rt s of th e pl a nt s
,
A nd i f .
th e l es s s p eci a li zed i n s ect s p os s es sed s om e d eg ree o f su c h

-
dis crimi na ti on th en the i niti a l c ont ra s t s of c ol ou r a b ov e

,
d es crib ed wou ld be m a i nt a i ned a n d i n crea s ed : L et s u ch a ‘
conn exi on be on c e e s t a bli s h ed a nd it m u s t t en d t o b ec om e

,
m ore d ecid ed I n s ec ts m os t a bl e t o di s c ern th e p a rt s o f

.
pl a n t s whi ch a ff or d wh a t th ey s eek w ill be th os e m os t lik ely ,
t o s u rviv e a n d l ea v e off s p ri ng P l a nt s p res enti ng m os t o f

.
t h e d es i re d food a n d s h owi n g m os t c l ea r ly wh ere it li es w ill

, ,
h a v e thei r fertili za ti on a nd mu l tiplic a t i on fu rthere d i n the

grea t est d egr ee A nd s o th e m u t u a l a d a pt a ti on wi ll b ec om e
.
e v er c l o s er ; whil e it is rend ere d a t th e s a m e ti m e m ore
v a r i ed by the S p eci a l requ i rem en t s o f th e i n sec t s a nd of

t h e pl a n ts i n ea ch l oc a lity u n d er ea ch ch a ng e o f c on ,
d i ti on s . Of c ou r s e the genes i s of th e s wee t ,
s ecre ti ons a nd th e od ou r s o f fl ower s h a s a p a r a ll el i nter re

p ,
t a ti on
. T h e s imu lt a neou s p rod u cti on of h on ey or s om e ,
ki n d red su b s t a nc e i s i m pli ed a b ov e ; S i n ce u nl es s a b a it
, ,
co exi s t e d w ith t h e col ou r

-
th e c ol ou r w ou ld n ot a tt ra ct
,
i n sect s a nd w ou ld not be m a i nt a i ned a n d i nt ens ifi ed by

,
n a t u ra l s el ec ti on Gu m s a nd res i ns a n d b a l s a m s a re f a mili a r
.
, , ,
p rod u ct s of pl a nts : a pp a rently i n m a ny ca s es ex cret ed a s , ,
u s el ess m a tt ers f rom v a r i ou s p a rt s o f th e i r s u rf a c es T h ese .
s u b s t a nces a dmitti ng of wid e v a ri a ti on s i n q u a lity a s th ey

, ,
d o a fl ord Opp ort u niti es for th e a cti on of na t u ra l s el ecti on

,
wh er ev er a ny of th em a tt r a ctiv e t o i ns ec t s h a pp en t o be
, ,
p rodu ced nea r the org a n s of fru ctifica t i on A nd thi s a cti on .
O f n a t u ra l s el ec ti on on c e s et u p m a y l ea d t o t h e es t a bli s h ,
m ent of a l oca l exc reti on t o the p rod ucti on of a n exc reti on

,
m ore a nd m ore a tt ra ctiv e a nd to the di s p os a l of the org a n

,
c ont a i ni ng it i n s u ch a wa y a s m os t t o f a cilit a t e the ca rry ’
i ng a w a y o f p oll en S imil a rly a nd s im u lt a neou s ly wi th

.
od ou rs . Od ou rs lik e c ol ou rs d ra w i ns ects t o fl ow ers

, , A ft er .
T HE OU T E R T IS S U E S OF P L A NT S .
269
o b s ervi ng h ow B ees c om e s wa r mi n g i nt o a h ou se wh ere

h oney i s l a rg ely exp os ed or h ow Wa s p s fi n d th ei r w a y i n t o
,
a s h o p c o nt a i n i ng m u c h rip e fru it it c a nnot be qu es ti oned ,
th a t in s ects a re t o a c ons id era bl e ex tent gu id e d by s cent .
Bei ng th u s s en s itiv e t o th e a rom a ti c s u b st a n ces whi c h fl ow ers

exh a l e th ey m a y w h en th e fl owers a r e i n l a r g e m a s ses
, ,
be ,
a tt r a c t e d by t h em f r om di s t a n ces a t w hi c h t h e fl ower s th em
s elv es a r e i n vi s ibl e
.
A nd m a n i fes tly the fl owers whi c h s o
.
,
a tt ra ct t h e m fr om th e g rea t es t di s t a nc es i ncr ea s i ng t hereby ,
t h ei r c h a nces of effi ci en t fertili za t i on w ill be m os t li k ely t o ,
p er p etu a t e th em s elve s T h a t i s t o s a y s u rviv a l of th e fit t es t

.
,
m u s t t en d t o p r o d u ce p er fu m e s th a t a r e b o th m or e p ow erfu l
a n d m o re a tt r a c tiv e .
T h es e phy s i ol o gi ca l di fferen t i a ti on s th en w hi ch m a r k Off , ,
the foli a r o rg a n s c on s t it u ti n g fl ow er s fro m oth er fo li a r or g a n s ,
a r e th e co n s eq u en ce s o f i n di re c t e qu ilib r a t i on T h ey a re not .
du e t o th e i m m edi a t e a ct i on s o f u n lik e i ncid en t for ces on

t he p a rt s of the i n divid u a l pl a n t ; bu t th ey a re du e t o t he
a cti on s o f s u ch u n lik e i n c id ent f or c es on th e a gg reg a t e of
i ndivi du a l s gen era ti on a fter g en era ti on

,
*
.
2 76 . Th e ity of i nt erp ret a ti on whi ch we h ere fi nd for

un
ph en om ena of s u ch v a r i ou s or d ers c ou ld h a r dly be fou n d ,
Th i s p l a ce a s a ny for not i ng th e fa ct, th a t th e gre a t er p a r t

s e em s a s fit a
o f wh a t we c al l bea u t
y i n the orga ni c w orl d, i s i n s om e w a y d e pe ndent on
x
th e s e ua l r el a t i on I t i s no t onl y so w ith the c ol ou rs a nd odou rs of owe rs
. fl .
It i s s o, t oo , wi th t h e b ri l li an t p l um a ge of bi rd s ; a nd i t i s p ro b a bl e t hat the
c ol ou rs o f th e m ore co ns i cuou s
p i ns ect s a re i n p a rt s i m il a rl y de te rm i ne d The .
re m a rk a b l e c i rcu m s t a nce i s , th a t t h es e ch a r ac t e ri s t i c s , wh i ch h a ve ori gi na ted
by fu rt heri ng th e p rodu ct i on of th e be st o ffs pr i ng, wh i l e t h e y a re na tura ll y
t h os e w hi ch rende r t h e o rga n is m s pos s e s s i ng th em a t tra ct i v e to one anoth er,
d i re ctl v or i ndi re ct l y, sh o ul d a l so b e t h os e whi ch a re g en era l l y a tt r a c ti e

so v
t o u s — th os e w i th ou t wh i ch t h e fi el ds a nd w oo ds w oul d l o s e h a l f th e i r ch a rm .
I t i s i n tere s t i ng , too , t o ob s er ve ho w th e con c e pt i o n of hum an b e a u ty i s i n a
con s idera b l e degree th u s or i g i na t ed . A nd t h e t ri t e v

ob ser a t i on that th e
e l e m ent o f be a u ty w hi ch g row s out of th e s e xua l rel a ti on i s so p re dom i na nt
i n a s th e ti c p rod uc t s— i n m u si c, i n th e d ram a , i n fi cti on, i n poe t ry— ga i ns a
ne w m e an i ng wh en we s ee how dee
p down i n o rga ni c na t ure thi s connex ion
e xtends .
w ere th e ph enom ena o th erwi s e c a u sed T h a t th e st ronger .
a nd th e f ee bl er cont r a s ts a m ong t he di f ferent p a rt s of th e

ou t er ti s s u es i n pl a n t s s h ou ld S O c ons t a n tly occu r a l ong with
,
s tr onger a n d f eebl er c ont r a s t s a m on g th e i ncid ent forc es i s ,
i n it s el f weighty evid en c e th a t u nlik e ou t er a c ti on s h a v e

ca u se d un lik e i nn er a cti ons a nd c orres p ondi ngly u nlike
,
-
s t ru ct u res ; e ith er by c h a n gi n g th e fu n cti ona l eq u ilib ri u m i n
t h e i ndivid u a l or by ch a n gi n g it i n th e ra c e or by b oth
, ,
.
E v en i n th e a b s en ce of m or e di rec t p roof th ere w ou ld be ,
grea t s ign ifi ca nce i n the m a rk ed diff e renc es th a t h a bit u a lly

e xi s t b et wee n th e e xp ose d a n d i m b edd ed p a rt s o f pl a nt s be ,
t ween t h e s tem s a nd th e l ea v es a n d b et w een t h e u pp er a n d

,
u nd er s u r f a c es of th e l ea v es T h e s ign ific a nce of t h es e differ

.
e n ces i s i nc re a s ed w h en w e di s c o v er t h a t th ey v a ry i n d eg r ee
a s th e di ffer en c es i n t he con diti on s v a ry i n d eg r ee S till .
g rea t er b ec om es the force o f th e evi d en ce on fi ndi ng th a t

th es e s t rongly c ont ra s t ed p a rt s m a y wh en pl a c ed i n one
-
,
a n oth er s c o
’
n diti on s a n d k ept i n th em f ro m g ener a ti on t o
,
’
g enera ti on p erm a nen tly a ss u m e one a n oth er s fun cti ons a n d
, , ,
’
i n a g rea t d eg ree on e a n oth er s s t ru ct u res
,
E v en m ore c on .
e l u s iv e yet i s t h e a r gu m en t r end er ed by the di s c ov ery th a t

, ,
wh ere t h es e s u b s tit u ti ons of f un c ti on a n d st ru ct u re t a k e pl a c e ,
t he s u p eri n d u ced m odific a ti ons differ i n diff er ent ci r cu m

st a nces ; j u s t a s th e origi na l m odifi ca ti ons do T he f a ct th a t .
a fl a t tened s t e m s im u l a ti ng a v ertic a lly g rowi ng l ea f h a s i ts-
two s u rfa c es a lik e w hil e w h en it s imu l a t es a h or i zont a lly

,
gr ow i ng l ea f i ts u pp er a nd u n d er s u rfa c es diff er i s a fa c t ,
which s t a ndi n g a l one might p rov e li t tl e bu t p r ov es m u c h

, , ,
wh en j oi ned with a l l th e oth er evid enc e A n d its p r ofou n d

.
m ea n i n g b ecom es th e m ore obvi ou s on di s c overi ng th a t th e

s a m e thi ng h a pp ens with p eti ol es w h en th ey u s u r p l ea f
fu nc ti on s .
Fi na lly w h en we r em emb er h ow r a pidly a na l og ou s m odi

,
fi ca t i on s o f f u ncti on a n d s t ru c t u re a r i s e i n t he su perfi ci a l
t i s su es of i ndi vid u a l pl a nt s th e gen era l i n f erence ca n s c a rc ely
,
be res i s t ed Wh en we m eet with S O st riki ng a c a s e a s th a t

.
THE OU T E R T ISS U E S OF P L A NTS .
of the B eg oni a -
l ea f a f ra gm ent of w hi ch s t u ck i n th e gr ou n d
,
p rod u ces r oot s f rom i ts u nder s u rfa ce a n d l ea v es fro m i ts

u pp er s u r fa ce— w h en w e s ee th a t th ou gh ,
i n thi s c a s e th e
,
typi ca l st ru c t u re of the pl a n t p resently b egi ns t o c ont ro l t he

org a n i z i n g p r oces s yet the i n iti a l di f
,
feren ti a ti on s a r e s et u p
by the di fferenti a l a ct i ons of th e envi ronm en t ; the p res u m p
ti on b ec om es ext remely s t rong th a t th e h et erogeneiti es o f
s u rf a ce which we h a v e c ons id ered resu l t a s a ll eged di rectly
, , ,
o r i ndi rectly f r om h et erog eneit i es i n th e i n cid ent f orc es .

CH A P T E R I V
I
D FFE RE N T A T I IO N S AM ON G TH E IN NE R I UE S
T SS OF
*
P L ANTS .
2 77 p a s s i ng from pl a nts f or m ed of threa d s or thi n

. IN
l a m i na t o pl a nt s h a vi ng s om e m a s s iv enes s we fi nd th a t a fter
, ,
t he ext erna l a n d i nt ern a l p a r t s h a v e b ec om e di s ti n gu i sh e d

f ro m one a n oth er th ere a ri se di s ti n cti ons a m ong th e i nt erna l
,
p a rt s th em s elves a s w ell a s a m ong th e ext erna l p a rt s

,
th em selves : the p rim a rily di fferenti a t e d p a rt s a re b oth r e-
di fferenti a t ed .
F rom typ es of v ery l ow org a ni s a ti on illu s t ra ti ons of this

m a y be d ra wn I n th e thi nner ki n d s o f L a m i na ri a th ere
.
exi s ts bu t th e s i ngl e c on t ra st b etw een t h e ou t er l a y er o f c ell s
a n d a n i nner l a y er ; bu t i n l a rg er s p eci es o f t h e s a m e g enu s ,
as L di gi ta ta th ere a r e th ree u nl ik e l a y ers on ea ch s id e of a

.
,
c ent ra l l a yer di fferi ng f rom th em— a u gm ent a ti on of b ulk is

a c c om p a n i e d by mu ltipli ca ti on of c on cent ric i nt ern a l s t rue
t u res h a vi ng th ei r u nlik enes s es obvi ou s ly rel a t ed t o u nlik e

,
nes s es i n th ei r c on diti on s I n F u rcel l a ri a a n d v a ri ou s A l gae

.
o f s i m il a r ly s w oll en f or m s th e lik e rel a ti on m a b e t r a c ed

y ,
.
Ju s t i ndi ca ti n g th e genera lity of thi s c ontra s t bu t not ,
Stu d e n t s of v ege t al
ph ys i ol ogy, fa m il i ar wi th th e cont rov e rs i es re specti ng
s undry p oi nts de a l t w i th i n th i s ch a pte r, w i l l p rob a bl y be s urp ri s e d to fi nd
t ak en fo r gra nted i n i t, p ropos i tions whi ch th ey h a v e h a bi t ua l l y re g a rded a s

o p en to dou bt . Hence i t s eem s needful to s ay th a t the concl u s i ons h ere se t
forth , h a v e re s ul te d from i nv e s t ig a t i ons un de rt a k en for th e p urpos e of form

i ng v
opi n i on s on s e e ra l uns e ttl e d q ue s ti ons wh ic h I h a d t o t rea t, bu t whi ch I
c oul d fi nd i n book s no a de q u a te d a t a for t rea ting . The det a i l s of t h es e
i nv es ti ga ti ons , and t h e enti re a rg um ent of whi ch t hi s ch a pte r i s pa rtl y an
a bs t ra c t, will be found i n A pp endi x C.
2 72
TH E I NNE R T ISS U E S OF P L A NT S .
2 73
t
a t em pti ng t o s eek i n thes e l ow er typ es for a ny m ore sp eci fic
i nterp reta ti on o f it l et u s p a ss t o the high er typ es T he
, .
a r g u m ent will be a mply en forced by t h e e vi d ence obt a i ne d
from t h e m . We w ill l ook fi rs t a t the c on di t i ons whi c h th ey

h a v e to fu lfil ; a nd th en a t the w a ys i n whi ch the functi ons
a n d s t ru c t u r es a d a pti n g th e m t o th es e conditi on s a r i s e .
§ 2 78 . At e r r e s t r i a l pl a n t th a t gro w s v erti ca lly nee ds no

m a rk ed m odifi ca ti on of i ts i n t ern a l ti s su es s o l ong a s the ,
h eight it rea ch es i s v ery s m a ll A s we be f ore sa w the s pi ra l

.
,
or c yli n d ri c a l rolli n g u p o f a S impl e c ell u l a r f r ond or t h e ,
m ore b u lky g rowt h of a s impl e cell u l a r a xis m a y giv e the ,
requ i s it e s t ren gth ; a n d t he r equ i s i t e ci r cu l a ti on m a y be c a r
ri e d on th r ou gh t h e u n ch a ng e d cell u l a r ti s s u e B u t i n p ro
.
p orti on a s the h eight t o be a tt a i ned a nd the m a s s t o be

s u pp ort e d i n cr ea se th e s u pp or ti n g p a r t m u s t a cq ui re g rea t er
,
b u lk or g rea t e r d en s ity or b oth ; a nd s o m e m odifi ca ti on th a t

,
s ha ll fa c ilit a t e t h e t ra ns f er o f nu t ritiv e li q u id s m u st t a k e
pl a ce H enc e i n th e i nner ti s su es of pl a nts w e m a y exp ec t

.
,
t o fi nd th a t s t ru ctu ra l ch a ng es a ns weri ng t o th es e requ i re

m ent s b ec om e m a rk ed a s the growth of the a e ri a l p a r t
,
b ec om es grea t F a ct s c orres p on d with th es e exp ec t a ti ons

. .
A m ong t he h u mbl er Co rm ophyt e s whi c h c ree p o v er or

,
ra i se th em s e lv es bu t littl e a b ov e t he su r f a c es th ey fl ou ri s h
,
u p on ,
th ere i s s ca rc ely a ny in t erna l di fferenti a ti on : the
v a s cu l a r a n d w o ody s t ru ct u res if n ot i n a l l ca ses a b s olu t ely
,
u n rep re s en te d a re r a r ely a n d v ery feebly i n dic a t ed

,
Bu t .
a m on g t h e high er typ es — th e F e rn s a n d L y c op odi u m s
wh i c h r a i s e th ei r f ron d s t o con s id er a bl e h eight s th ere a re ,
v a s c u l a r bu ndl es a nd h a r d ti s su es lik e woo d ; a nd by the

T ree F ern s m a s s iv e a x es a re d ev el op ed
-
T h a t the rel a ti on
.
whi ch th u s s h ow s it sel f a m on g Cr ypt og a m s i s h a bit u a l a m ong

P h a n og a m s s c a rcely nee d s s a yi n g
,
.
P ha n og a m s h ow ev er a re n ot u niv ers a lly th u s ch ara c

, ,
t er i zed i n a d ecid e d wa y .B es id es th e c omp a ra tiv e w a n t of

w o ody ti ssu e i n fl ow eri ng pl a nts of h u mbl e g r owth a nd ,
64
2 74 P H Y S I OL OGI CA L D E V E L OP ME NT .
b es id es th e p a u city of v es s el s i n ordi na ry wa t er pl a nt s th ere -

,
a re c a s e s o f m u ch m ore m a rk ed div er g en ce fro m t h i s t ypi c a l
intern a l s t ru ctu re . T h ese ex cepti ona l c a s es occu r u n d er

ex cep t i on a l c on diti on s a n d a re highl y i n s t ru ctiv e
,
T h ey a re .
o f t w o ki n d s . One gr ou p of th em i s fu r ni sh ed by
c er t a i n pl a nt s w hi c h a r e p a r a s iti c on th e e xp os ed r oot s o f
t rees— p a ra s itic n ot p a r ti a lly a s th e Mi s tl et oe bu t t o th e

, ,
e x t en t o f s u b s i s t i n g w h o lly on th e s a p th ey a b so rb F u ngu s .
lik e i n c ol ou r a n d t ex tu re a nd h a vi n g s c a l es for l ea v es th es e
, ,
B a l a n op h or ae a n d Ra fil es i a ceae a re recogni za bl e a s P ha n og a m s
by s ca r cely a ny oth er t ra it s th a n th ei r fru cti fi ca ti ons A l ong .
with th ei r a b ort ed l ea v es a nd a b senc e of chl orophyll th ere ,
i s a g rea t d eg ra d a ti on o f t h os e i n t er na l ti s su es by w hi c h
P h a n og a m s a re com m onl v di s ti ngu i s h ed T h ou gh D r [ n ow
. .
S i r J ] H ook er ha s s h own th a t th ey a re not a s s om e b ot a ni s t s

,
th ou ght d ev oid of s pi ra l v es s el s ; yet a s S h own by the m i s

, ,
t a k e p revi ou s ly m a d e i n c l a s s i fyi ng th em th ei r a ppli a nces for

,
c i r cu l a ti on a r e ru di m en t a ry . A n d thi s t ra it g oes a l on g w ith a

grea t ly—s i m pl i fi ed di s tribu ti on of nu t rim ent I n the a b s enc e .
of l ea v es th ere ca n be bu t littl e d ow n cu rren t o f s a s u ch

p
-
,
a s l ea v es u s u a lly s u pply t o root s : th ere c a nn o t be m u ch
b ey ond a n u p wa rd cu rrent o f th e a b s orb ed j u i ces T he .
o t h er c a s es o cc u r wh e r e c i r cu l a ti on i s a rr es t e d or ch eck ed i n
a differen t w a y ; na m ely i n pl a n t s th a t a r e wh olly s u bm erg ed

"
, .
T h es e a re th e P odos tem a ceae .C l othi ng a s th ey do th e s u b

m erged rock s th ei r r oots pl a y th e p a rt of rhi z om es b ei ng
, ,
a t t a c h ed t o th e s u b s t ra t u m by h a i rs a nd o th er p r oc es ses a nd ,
h a vi n g the l ea f b ea ri ng a nd fl ower b ea ri ng s h o ot s on th ei r
- -
s u r f a ces. T he l a tt er sp rea d ou t m ore or l ess h ori z on t a lly a n d

a r e a ls o fix ed t o th e s u b s t r a t u m i n th e s a m e m a nn er a s th e
root s . Ob s erv e th en th e c onnexi on o f fa cts One of th es e .
P odos tem a ceae n ee d s n o i nt ern a l s ti ffeni ng s u b s t a nc e for it ,
exi s t s i n a m edi um o f i t s own s p ec ifi c g ra vity ; a n d b ei n g i n
a p os iti o n t o a b so r b wa t er o v e r i t s ent i re su r f a c e it h a s n o ,
nee d for a ci rcu l a ti on of cru d e s a p— nor i nd eed i n the

, ,
a bs ence of ev a p ora ti on fro m a ny p a rt of its s u rf a c e cou ld ,

T HE INNE R T ISS U E S OF P L A NT S .
2 75
a ny ac tive c ir cul a t i on t a k e pl a ce H ere a cc or di n gly t he

.
, ,
t ra c h ea l a nd m ech a n i c a l el em en t s a re u nd ev el op ed T h ou gh .
s pi ra l v es sel s a r e not e nt i r ely a b s en t yet th ey a re s o r a r e a s

,
t o d o n o m or e th a n v er i fy t h e i n fer ence of pha n og a m i c rel a

t i on s h i p d ra wn fro m th e fl ow ers .
T he m eth od o f a greem en t th e m eth o d o f di fferen ce a n d

, ,
t h e m eth od o f c on com it a n t v a r i a ti on s thu s u n it e i n p rovi n g

,
a di rec t r el a ti o n b et w een th e d e m a n d for s u pp or t a n d c i r cu
l a ti on a n d the e xi s ten c e of t h es e v a s cu l a r wo ody bu n dl e s

,
wh i c h th e high er pl a nt s h a bit u a lly p os s es s T h e qu es ti on .
whi ch we h a v e t o c on s id er i s — Und er w h a t i n flu enc es a re

th ese s t ru ctu res a n s weri ng t o th es e requ i rem en t s d ev el op e d ?
, ,
H ow a re th ese i n t ern a l di fferen ti a ti on s c a u sed ? T he i n qu i ry

m a y be c on v eni en t ly divid ed T h ou gh th e s u pp or ti n g ti s su es
.
a n d th e ti s s u es con cern e d i n th e c i rcu l a ti o n of liq u i ds a r e
c l o sely c o nn ec t ed a n d i n d ee d ent a n gl e d
,
w ith on e a n oth er , ,
w e m a y fi tl y d ea l with th em a p a r t L et u s t a k e fi rst t he
.
s u pp orting t i s su e .
§ 2 79 . Ma ny c om pl a ce f a cts i ndic a t e th a t th e m e
m on-
ch a n i ca l s t r a i n s t o w hi c h u p r ight gr ow i n g pl a nt s a r e exp os ed ,
th em selv es cau se i n crea s e of the d en s e d ep os i ts by whi ch su ch

pl a n t s a re en a bl ed t o res i s t su ch s t r a i n s T h ere i s the f a ct .
th a t th e m a s s iv enes s of a t ree t ru n k v a ri es a c c ordi ng t o the

-
s t r ess h a bit u a lly p u t u p on it I f th e c on t r a s t b etw een th e

.
sl en d er s tem of a t ree grow i ng i n a w ood a n d th e bu lky s t e m

o f a ki n d r e d t r ee g row i n g i n th e fi eld s be a scr ib e d t o di ffer ,
e n ce o f nu t r i t i on ra t h e r th a n di f fer en ce o f exp os u re t o wi n d s
t h er e i s s till th e f a ct th a t a t r ee t r a i n ed a g a i nst a w a ll h a s a
l es s b u lky s t em t h a n a t ree of th e s a m e ki n d g rowin g u n
su pp ort e d ; a n d t h a t b et w een t h e l on g w ea k b r a n ch es o f th e
o ne a n d th e s t i f f on es o f th e oth er t her e a r e d ecid ed con t r a s t s .
I f it be obj ec t ed th a t a t ree s o t r a i ne d a n d b ra n ch es s o b orn e

hav e rel a t iv ely l es s foli a ge a n d t h a t th ere f or e th es e u n lik e
,
s
'
n es s es a ls o a re d u e t o u n lik en es s es of g en er a l n u t riti on whi ch ,
m a y i n p a r t he t ru e; th ere a r
'
e s till su ch c a se s a s th os e o f
2 76 P H Y S IOL OGI CA L D E VE L OP ME NT .
g a rd en pl a nt s w hi ch wh en held u p by tyi ng th em t o s ti ck s
,
h a v e wea k er s tem s t h a n w h en t h ey a re u np ropp ed a nd si nk ,
d own if th ei r p rop s a re t a k en a wa y A g a i n th ere i s the .

,
e vid en ce s u ppli e d by roo t s T h ou gh t he c on t ra s t b et ween

.
t he fee bl e root s o f a s h el tered t ree a n d t h e s t r on g root s o f

a n exp os e d t ree m a y lik e th e c on t r a st o f th ei r S t ems be
, , ,
m a i n ly du e t o di ffer enc e of nu t r iti on a n d th ere fore s u ppli es

,
bu t d ou bt fu l evid ence w e g et t o l era bly c l ea r evid en ce w h ere

,
t rees g row i n g on i n c li n ed rocky su r f a ces s en d i nt o c revic es ,
th a t a ffor d littl e m oi s tu re or nu t ri m ent r oot s w hi ch nev er ,
t hel ess b ecom e thi c k w h er e th ey a re s o di rect ed a s t o b ea r

grea t S t ra i ns . S u s pi c i on th u s ra i se d i s s t rength ened
i nt o c onvicti on by s p eci a l evid ences occu rri n g i n th e pl a c es
w h er e th ey a re t o be exp ec ted T he C a ct u ses w i t h th ei r
.
,
s u cc u l en t g r ow th s th a t p a s s i n t o w oody gr ow th s s l ow ly a nd
i rreg u l a rly giv e u s the opp ort u nity of t ra ci ng the c onditi ons
,
u nd er w hi c h t h e w oo d i s f orm ed Good ex a mpl es occu r i n th e

.
g en u s C er eus a nd es p eci a lly i n f orm s lik e 0 cr enu l a tus

, . .
H ere f r om a m a s s iv e v erti c a lly g r ow i ng r od of fl eshy ti ss ue

,
-
,
t wo i nch es or m ore i n di a m et er th ere g r ow a t i nt erv a l s l a t er a l

,
r od s s i m il a rly b u lky w hi ch qu i ckly c u r vi ng th em selv es t a k e

, , ,
v ertic a l di recti ons One o f t hes e h ea vy b ra n ch es p u t s grea t

.
st ra i ns on i t s own s u b s t a n ce a n d th a t of t h e s t em a t th ei r
p oi nt o f j u n cti on ; a n d here b oth Of th em b ec om e b rown a nd
h a rd whil e th ey c on ti nu e g reen a nd su ccu l en t a l l a rou n d
,
.
S u ch di fferen ti a ti on s m a y be t ra ced i nt erna lly b efore th ey

a re vi s ibl e o n t h e s u r fa ce I f a j oi nt o f a n Op un ti a be s li ced
.
th rou gh l ongi tu di n a lly th e gr ea ter res i s t a n ce t o the kn i fe

,
a l l a r ou n d t h e na rr o w n ec k i ndi c a t es t h ere a l a r g er d ep os it
,
o f lig ni n t h a n e l s ewh er e ; a n d a s ec t i on o f th e ti s s u e pl a ce d
u nd er t h e mi c ros c op e e xhibit s a t the n a rrow es t p a rt a con

,
c en t ra t i o n o f t h e w o o dy a n d v a s c u l a r b u ndl es Clea r .
e vid en c e o f a n oth er ki nd h a s b een n o t e d by Mr D a rwi n i n th e .

,
or g a ns o f a tt a chm en t o f c li m bi n g pl a n t s S p ea ki ng of S ol a
.
n u m ja s m i n oi des h e s a ys Wh en the fl exibl e p eti ol e o f a

‘
h al f or a q u a rt er g rown l ea f h a s c l a s p ed a ny obj ect i n th ree

-
,
T HE INNE R T I S S U E S OF P L A NT S .
2 77
or f ou r d a ys it i nc rea s es m u ch i n t hi cknes s a nd a ft er s ev era l ,
wee k s bec om es w ond erfu lly h a rd a nd rigid ; s o th a t I c ou ld

h a rdly re m ov e on e from i ts s u pp ort On c om p a ri n g a t hi n
.
t ra n sverse s li ce of thi s p eti ol e wi th one f rom t he next or

o ld er l ea f b e nea th which h a d not c l a s p e d a nythi ng i ts
, ,
di a m et er wa s fou n d t o be fu ll y d ou bl ed a nd i ts s t ru ctu re ,
g rea tly ch a n ged . T hi s cl a s p e d p eti ol e h a d a ctu a lly

b ecom e t h i ck er th a n th e s t e m cl os e b en ea t h ; a nd thi s wa s
c hi efly du e t o th e g r ea t er thick nes s of t h e ri ng of w ood ,
which p r es ent ed b oth i n t ra n s ver se a n d l on git u di na l s ec ti ons

, ,
a cl os ely S imi l a r s t ru c t u r e i n t h e p e ti ol e a n d a xi s T he
“
a s s u m p t i on by a p eti ol e of thi s s t ru ctu re i s a s i ngu l a r

m orph ol ogi ca l f a ct ; bu t it i s a s till m ore si ng u l a r phy s i o ~
l ogi ca l f a ct th a t s o grea t a ch a n ge sh ou ld h a v e b een i n d u ced

”
by the m ere a ct of cl a s pi ng a su pp ort .
I f th ere i s a di rec t rel a ti on b et ween m ec h a n i ca l s t res s a n d

t he f or m a ti on of w oo d it ou ght t o expl a i n for u s t he i n t ern a l
,
di strib u ti on of th e w ood L et u s s ee wh eth er it d oes thi s

. .
When seeki n g i n m ech a nic a l a cti ons a nd rea cti ons the
c a u s e of th a t i ndu ra t ed s t ru ct u re whi ch form s the v erte
b ra t e a xi s 254 it wa s p oi nt ed ou t th a t i n a t ra ns vers e
l y s t ra i ned m a s s the grea t es t p res su res a nd t en s i on s a re
-
,
th row n on the m ol ec u l es of the c on ca v e a n d conv e x s u r f a ces .
H ence su pp os i n g th e t r a ns v ers ely s t ra i ne d m a s s t o be a cyl i n

,
-
der b ent b a c kw a r d s a nd forw a rd s n ot i n on e pl a n e bu t n ow

,
i n thi s pl a ne a n d n ow i n th a t i t s p er iph er a l l a y ers wi ll be

,
t h ose on whi ch the g rea t es t s tres s f a ll s A n ordi na ry di coty

.
l edon ou s a xi s i s s u c h a cyli n d er s o s t r a i n ed Th e m a i n .
t ena nce o f i ts a ttit u d e e ith er a s a l a t era l S h oot or a v ert i ca l

s h oot impli es s u bj ec ti on t o t h e b en di n g s ca u s e d by i t s own
,
weight a nd by t he ev er v a ryi n g wi n d -
T h es e b en di ng s
.
i m ply t ens i on s a nd p res s u res f a llin g m os t s everely fi rs t on

o n e S id e o f i t s ou t er l a y er s a n d t h en o n a n oth er A n d i f th e .
d ense su b s t a nce a bl e t o res i s t th es e t en si ons a nd p re s s u res i s

dep os it ed m os t wh ere th ey a re grea t es t we ou ght t o fi n d it,
t a ki ng the s h a p e o f a cyli n d ri c a l c a s i n g T hi s i s j u s t wh a t
.
278 P H Y SI OL OGI CA L D E VE L OP ME NT .
we do fi nd . On
tti n g a cr os s a s h oot i n c ou rs e of f or m a ti on
cu ,
we s ee i t s c ent ra l s p a ce e i th er u n occu pi ed or occu pi e d only

by s oft ti s s u e T h a t th e l a y er of h a r d t i s s u e su rrou ndi ng
.
thi s i s not the ou t erm os t l a y er i s t ru e : th ere li es b ey on d i t

,
th e c a mbi u m l a y er fr om which i t i s form ed th e phl oem

, , ,
a nd th e c or t e x B u t ou t s id e of th e s o ft phl oem th ere i s

.
f requ ently a no th er l a yer of d ense ti s s u e now kn ow n a s the

p ericyc lic fib res h a vi ng freq u en tly a t ena city grea ter ev en
,
th a n th a t of the w ood— a l a yer whi ch whil e it p r ot ect s the ,
c a mbi u m a nd O ffers a dditi ona l res i s t a n ce t o the t ra n s verse

st ra i n a d m its of b ei ng fi s s u red a s fa s t a s th e cyli nd er of
,
w ood thick ens T h a t i s t o s a y the d ep os it of res i s ti ng su b

.
,
st a nce i s a s com pl et ely p eriph era l a s the ex ogen ou s m od e of

gr owth p erm it s S o too i n g enera l a rra n gem ent i s it w i th
.
, ,
t he ordi n a ry m onocotyl ed on ou s s t em D i fferent a s i s h ere .
t h e i nt erna l s t ru c t u re th er e yet h old s th e s a m e genera l di s

,
t ri bu t i on of ti ss u es a n s weri ng t o th e s a m e m ech a n i ca l c on
,
d i ti ons T he v a s cu l a r wo ody b u n dl es m ore a b u nd a nt t owa rd s

.
,
t he o u t s id e o f the S t em th a n nea r th e c en t re p r od u ce a h a rd er ,
ca s i ng s u rrou ndi ng a s o ft er c or e I n th e s u pp orti n g

.
s t ru c tu res Of l ea v e s we fi n d s ign ifi c a n t d evi a t i ons f rom thi s
a rra n g em ent Whil e a x es a re on the a v era ge exp os ed t o

.
eq u a l s t ra i ns o n a l l s id es m o s t l ea v es s p rea di ng ou t th ei r
, ,
s u r fa c es h ori z on t a lly h a v e thei r p eti ol es s u bj ec t t o s t ra i n s

,
th a t a r e n ot a lik e i n a l l di recti on s ; a n d i n th e m th e h a r d
ti ss u e i s di fferently a rr a nged I t s t ra ns v ers e s ec ti on i s
.
not r i ng s h a p ed bu t c res cen t s h a p ed : t h e tw o h orn s b ei ng

- -
di rect ed t owa rd s th e u pp er s u r fa ce o f th e p eti ol e T h a t thi s .
a rr a n g em ent i s o ne w hi c h a ns we rs t o th e m ec h a n i ca l c on
d i ti ons i s n ot ea s y t o d em on s t r a t e : w e m u s t s a ti s fy ou rs elv es
,
by n oting th a t h ere wh ere the di s t ribu ti on Of forces i s di f

,
ferent the di s t rib u ti on o f r es i s ti n g ti s s u e i s di ffere nt

,
A nd .
th en sh owi ng c oncl u s iv ely the c onnexi on b et ween th ese di ffer

,
e n ces ,
w e h a v e t he f a c t th a t i n p eti ol es g r owi ng v er ti ca l l y
a n d s u pp or ti ng pe lt a t e l ea v es— p eti ol es whi c h a r e th er ef ore
TH E INN E R TI SS U E S OF P L A NT S .
Su bj ect t o q
e ua l t ra ns verse s t r a i n s ides— the v a s cu l a r
on a l l S
b un dl es a re a rra ng e d c yli n d ri c a lly

a s i n a x es ,
.
S u ch t h en a re s om e of t he r ea s ons for c on c l u di ng t h a t th e
, ,
d ev el op m en t of t he s u pp ortin g t i s s u e i n pl a n t s i s c a u s ed by ,
t h e i n cid ent for ces whi c h thi s ti s su e h a s t o res i s t T he .
i n divi d u a l s i n whi c h thi s di rec t b a l a nci n g of i nner a n d ou t er

a c ti on s p ro g ress es m o s t f a v ou r a bly a re th os e w hi c h
,
o t h er ,
thi ngs eq u a l a re m os t li k ely t o p r os p er ; a nd by h a bi tu a l

, ,
s u rviv a l of the fitt es t th ere i s es ta bli sh ed a s y s t em a ti c a nd

,
c on s t a n t di s t r ib u ti on o f a d e p osit a d a pt e d t o th e c i rcu m
st a nces of ea c h type .
2 8 0 T he fu n cti on
. of
ci rcu l a ti on m a y now be d ea lt w ith .
We h a v e to con s id er h ere by wh a t s t ru ct u res thi s i s di s

ch a rge d ; a n d wh a t conn e xi on exi s t s b e t ween t h e d em a n d
for th e m a n d th e genes i s of th em .
The c on t r a s t b etw een th e r a t es a t w hi ch a dye p a s s es

th r ou gh si m pl e cell u l a r t i s s u e a n d cell u l a r ti s s u e of w hi ch th e
u n it s h a v e b een e l on g a t ed i n di ca t es on e o f th e s t ru c t u ra l
,
ch a nges req u i red to fa c ilit a te ci r cu l a ti on I f pl a c ed wi th i ts .
c u t s u r fa c e i n a c ol ou re d liqu id th e p a r en c hy m a o f a p ot a t o
,
or th e m e d u ll a ry m a s s o f a ca bb a ge s t a lk w ill a b s orb t he
-
,
liqu id w ith ext rem e s l ow nes s ; bu t i f th e s t a lk of a fun gu s be

s i m il a r ly pl a c ed th e liqu id r u n s u p it a n d es p ec i a lly u p its
, ,
l oos e centr a l s u b s ta n ce v ery qu i c kly , On c o m p a ri n g t he

.
ti s s u es whi ch thu s b eh a v e s o di fferently w e fi n d th a t wh erea s ,
i n the one ca s e t h e c om p on ent c ell s p a ck ed c l ose t o geth er , ,
h a v e d evi a t ed fro m th ei r p ri m itiv e s ph eri c ity only a s mu ch a s

m u t u a l p r es su r e n eces s it a tes i n th e o th er ca s e th ey a re d ra wn
,
ou t i nt o l o n g t u b u l es with na rro w s p a ces a m on g t h e m— th e
g rea tes t di m ens i on s of th e tu bu l es a n d th e S p a ces bei ng i n the

di rect i on w hi ch the dye t a k es s o r a pi dly
-
T h a t w hi ch w e .
s h ou ld i n fer th en f r om t he l a w s of ca pill a ry a c ti on is
, , ,
e xp erim ent a lly s h ow n : liq u id m o vi ng th r ou gh ti s s u es foll ow s
th e li nes i n whi c h t he el em en t s o f th e ti ss u es a re m os t
280 P H Y S I OL OGI CA L D E VE L OP ME NT .
l
e on ga t ed I t d oes thi s for two r ea s ons T h a t na rrowi ng of
. .
t he cell s a n d i nter c ell u l a r s p a ces w hi c h a ccom p a ni es th ei r

e l on ga ti on f a c ili t a t es c a pill a r i t y
,
a nd a t th e sa m e t i m e fewer
o f t h e sep t a f or m e d by t h e j o i ned en d s of th e c ells h a v e t o be
p a s s ed t h rou gh i n a giv en di s t a nce . H enc e th e

g enera l fa ct th a t t he es t a bli s h m en t of a ru dim ent a ry v a s cu l a r
s y s t em i s the f orm a ti on of b u n dl es of c ell s l en gth ened i n th e
,
di rec ti on whi ch t h e liqu id i s t o t a k e T hi s we see v ery

.
obvi ou s l y a m ong t h e l owe r C or m ophyt e s I n one of th e .
li ch en lik e L iv erw ort s th e v eins whi ch b r a nchi ng th r ou gh

-
, ,
i ts fron d s erv e a s c om m u nic a ti ons w i th i ts s c a t t ered rootl ets

, ,
a r e f or m ed o f c ell s l on g er th a n th o s e com p os i n g th e g en er a l
ti ssu e of the f rond : t he l ength s of th es e cell s c orres p on di ng

i n th ei r d i recti on s w i t h th e l engt h s of t h e v ei n s S o t oo i s .
, ,
i t wi t h th e mid rib s o f s u c h f ron d s a s a s su m e m ore d efi nit e

S h a p es ; a n d s o t oo i s i t wi t h the c ree pi ng s t em s whi c h
, ,
u n it e m a ny su c h f r ond s T h a t is t o s a y the c u rrent whi c h

.
,
s e t s t ow a r d s the grow i n g p a rt f rom t h e p a rt w hi c h s u ppli es
c ert a i n m a t er i a ls for g row th s et s th rou gh a p orti on o f th e

,
ti s su es c omp os ed of u nits th a t a re l on ger i n th e li n e of the

c u rren t th a n a t right a ngl e s t o th a t li n e T h e lik e i s t ru e
.
o f P ha n og a m s Omitti ng a l l ot h er ch a ra ct er i s ti cs of th os e
.
p a rt s of t h em thr ou gh which chi efly th e c u rrent s of s a p

fl ow we fi nd the u n if or m fa c t t o be th a t t h ey c ons i s t o f cell s
,
a n d i n t er ce ll u l a r s p a ce s di s ti n gu i s h e d fro m o th er s by th ei r
l en gth s I t i s th u s w ith v ei n s a n d mid rib s a n d p eti ol es ;

.
, ,
a n d if w e wi s h p r o of th a t it i s thu s w ith s t em s we h a v e bu t
,
t o ob s er v e th e c ou rs e t a k en by a c ol ou red s ol u ti on i n t o whi ch
a s t em i s i ns ert ed .
Wh a t i s th e origi na l c a u s e o f thi s di fferen ti a ti on ? I S it

p os s ibl e th a t thi s m odifi ca ti on of cell s t ru ct u re whi ch fa v ou r s
-
t he t r a ns fer of liqu id t ow a rd s ea ch pl a ce of d em a n d i s i t s el f ,
c a u s ed bv th e c u r ren t whi ch th e d em a n d s et s u ? D es t h e
p o
s t rea m m a k e i ts ow n ch a n nel ? T h ere a re v a ri ou s r ea s on s

for thin ki n g th a t it d oes I n th e fi rs t pl a c e th e s i m pl es t a nd
.
,
ea r li est ch a nn el s s u ch a s w e s ee i n the L iv erwo rts

, do not ,
TH E INN E R T ISS U E S OF P L A NT S .
281
d evel op i n a ny s yst em a tic wa y bu t b ra n ch ou t i rregu l a rly

, ,
fo ll owi ng e very w h ere th e i r regu l a r l ob es of the f ron d s a s

th es e s p rea d ; a n d on ex a m i n i n g u n d er a m a g nifier t he pl a c es
a t whi c h th e v ei n s a r e l os t i n th e c ell u l a r ti ss u e it w ill be ,
seen th a t th e c ell s a r e th ere s lightly l ong er th a n t h os e
a rou n d : s u gg es ti n g t h a t th e l en gt h en ing o f th em w hi ch
p r od u ces a n ext en s i on of the v ei ns t a k es pl a ce a s f a s t a s
,
t h e growt h o f th e ti s s u e be y on d ca u s es a cu rrent t o p a ss
th rou gh th em I n the s econ d pl a c e a di s a pp ea ra n c e of the
.
,
gr a nu l a r cont ent s of th ese c ell s a cc omp a n i es thei r u ni on

in to a v ein— a resu lt which th e t ra ns mi s s i on o f a cu rren t
m a y n ot i m p rob a bly b ri n g a b ou t B u t be the s p ec i a l c a u s es
.
o f thi s di f feren ti a ti on wh a t th ey m a y th e evid enc e f a v ou rs

,
v ery m u ch th e c onclu s i on th a t th e g enera l c a u se i s the

s etti n g u
p o f a c u rr en t t ow a r d s a pl a ce w h er e t h e s a p i s
b ei ng c on s u m ed . I n the hi s t ol ogi ca l d ev el op m en t
o f th e high er pl a n t s we fi nd con fi r m a ti on T he m ore
.
fin i s h ed di s t rib ut in g c a n a l s i n P h aen oga m s a re f orm ed of c ell s

p revi ou s ly l ength ened A t p a rt s of which th e typ ic a l s t ru e
.
t u re i s fix ed a nd the d ev el opm ent di rect thi s f a ct i s not ea s y

, ,
t o t ra c e ; the cel l s ra pidly t a k e th ei r el ong a t ed s t ru ct u res i n

a nti c ip a t i on o f th ei r pr e d et er min e d fu n c ti on s
-
Bu t i n pl a c es
.
wh ere new v es sel s a re requ i red i n a d a pt a ti on t o a m odifyi ng

gr owth we m a y cl ea rly t ra ce thi s s u cces s i on T he s welli ng
,
.
r oot o f a t u rn ip ,
c on ti nu a lly h a vi ng i ts v a s cu l a r s y s t em
fu rth er d ev el op ed a n d the com p on en t v es s el s l en gth ene d a s
,
well a s mu ltipli ed giv es u s a n opp ort u nity o f w a t chin g th e

,
p r oces s I n it we see th a t th e reti cu l a t ed c ell s w hi ch u n it e

.
t o form d u c t s a ri se i n th e mid s t o f b u n dl es of c ell s th a t h a v e

,
p revi ou s ly bec om e el on ga t ed a n d th a t th ey a r i s e by t r an s
,
form a ti on of s u ch el on g a t ed c ell s a n d w e a l s o see th a t t h es e

b u n dl es of el ong a t ed cell s h a ve a n a r ran g em ent s u gges tiv e
o f th ei r for m a ti on by p a s s i ng c u r rent s .
A re th ere gr ou n d s for thi nki n g th a t th ese fu r th er t ra n s

f orm a t i ons by w hi c h s t ri ngs of el ong a t ed c ell s p a s s i n t o
v es sel s li ned w ith s pi ra l a nnu l a r reticu l a t ed or oth er
, , ,
28 2 P H Y S I OL OGICA L D E VE L OP ME NT .
f ra mework s a re a l s o i n a ny wa y d et ermi ned by the cu rrent s

,
o f s a p c a rr i ed ? T h ere a re s om e s u c h g roun d s .
A s j u s t i ndi c a t ed t h e on ly pl a c es w h er e w e m a y l ook
,
for evid enc e wi t h a ny ra t i ona l h op e of fi ndi ng it a re ,
pl a ces wh ere s om e l oca l requ i rem en t for v es s el s h a s a ri sen

i n c on s eq u enc e of s om e l oc a l d ev el o p m en t whi ch th e typ e
d oes n ot i n v olv e . I n th es e c a s es w e fin d s u ch evid ence .
Goo d ill u s t ra ti on s occ u r i n t h os e g en era o f th e C a c ta c eae ,
which s imu l a t e l ea ves lik e E p i phyl l u m a n d P hyl l oca c tus

, .
A b ra nc h of one of th ese i s ou tli n e d i n Fig 2 5 6 A s b efore

. .
expl a i n ed thi s i s a fl a t t en ed a xi s ; a n d t h e n ot ch es a l on g
i t s edg es a r e th e s ea t s of the a x i ll a ry bu d s Mos t of t hese

.
a xill a ry b u d s a re a rre s t ed ; bu t oc c a s i on a lly on e o f t h em

g r ow s. N ow i f t a ki ng a n E pi p hyl l u m s h oot whi c h be a rs a
,
-
l a t era l s h oot we comp a re the p a rt s of i t th a t a re n ea r th e

,
a b or t e d a xill a r y b u d s w i t h th e p a rt th a t i s n ea r th e de
vel op e d a xill a ry bu d w e fi nd a c ons pi cu ou s di f

,
feren c e In .
t h e neighb ou rh o od o f a n a b ort ed a xill a ry bu d th er e i s n o

e xt ern a l s i g n o f a ny in t ern a l di f ferenti a t i on ; a n d on h old
i ng u p t he b ra nch a g a i ns t th e ligh t the u n i for m t ra ns
,
l u cency sh ow s th a t th ere i s n o grea t er a m oun t of d ens e

ti ssu e nea r it th a n i n oth er p a rt s of the s u ccu l ent m a s s .
Bu t wh ere a n a xill a ry bu d h a s d ev el op ed a p r om i nen t rou nd e d

,
r idge j oi n s t h e mid r ib o f th e l a t er a l b ra nc h with t h e mid ri b
o f t h e p a rent b r a n c h I n th e m id s t o f thi s rou nd ed r idge

.
a n o p a q u e c ore m a y be s ee n A n d on cu tti ng th rou gh i t thi s

.
,
Op a qu e c ore p r ov es t o be f u ll of v a s c u l a r bu ndl es i m b edd e d

i n w oody d ep os it s . Cl ea rly th ese cl u s t ers o f v essel s imply
,
t ra ns for m a ti on s of th e ti ssu es c a u se d by th e p a s s a g e of i n
,
c r ea s e d c u r rent s o f s a p T h e v e ss els were n ot th ere wh en

.
t he a xill a ry bu d w a s f orm ed ; th ey w ou ld n ot h a v e de
vel op ed h a d th e a xill a ry bu d p r o v e d a b ortiv e ; bu t th ey
a r i s e a s fa s t a s gr ow t h o f th e a xill a ry bu d d ra w s th e s a p
a l on g th e li n es i n w hich th ey l i e Verifi ca ti on i s obt a i ned

.
by ex a m i n i ng th e i nt ern a l s t ru ct u res I f l ongit u di na l

.
s ecti on s be m a d e throu gh a growi ng bu d of Op u n tia or

T HE I NN E R T I SS U E S OF P L A NT S .
283
C er eu s , i t w ill be f ou n d th a t the v es s el s i n cou rs e of f or m a

'
ti on c onv erg e t ow a r d s t he p oi n t of gr ow th a s they wou ld ,
do i f th e s a p cu rr ent s d et erm i n e d th ei r f orm a ti on ; t ha t th ey

-
a r e m os t d ev e l op e d nea r t h e i r pl a ce o f conv erg en ce w hi c h ,
t h ey w ou ld be if s o p rod u ced ; a n d th a t t h ei r t erm i na ti on s

i n th e t i s s u e of th e p a rent sh oot a r e p a rti a lly f orm ed li ne s -
of i rr egu l a r el on g a t e d c e ll s lik e th ose ou t o f w hi ch the ve s

,
s els of a l ea f or bu d a re d evel op ed .
C oncl u di ng th en th a t sa p v es sel s a ri s e a l ong th e li nes of

, ,
-
l ea s t res i s t a nc e th rou gh w hi c h cu rrent s a re d ra wn or f orced

, ,
t h e qu es t i on t o be a s k ed i s —Wh a t phy s i ca l p r oc es s p ro d u ces
th em ? T h ei r com p on en t cell s u n it ed end t o end m o re or l es s

,
i rreg ul a rly i n w a y s d et erm i ned by t h ei r ori gi na l p os i ti on s ,
f orm a ch a nnel m u c h m ore p erm ea bl e b oth l ongi tu di na lly

,
a n d l a t era lly th a n th e ti s s u e a r ou n d
,
H ow i s th i s g rea t er
.
~
p erm ea bility ca u sed ? T he id ea fi rs t p r op ou n d ed

,
I b eli ev e by Wolff th a t the a dj oi ne d en d s o f th e c ell s a re

,
per f ora t ed or d es troy ed by th e p a s s i ng cu rrent i s one for ,
whi ch mu ch i s t o be s a id Wh eth er th ese s ept a a re di s so lved

.
by the liqu id s th ey t ra ns mit or wh eth er th ey a re b u rs t by

,
th os e su dd en gu sh es whi ch a s we s h a ll h erea ft er s ee m u s t
, ,
frequ ent ly t a k e pl a ce a l ong t h es e c a n a ls n ee d not be di s ,
c u sse d : i t i s s u ffici en t for u s th a t th e sept a do i n m a ny ca ses , ,
di sa pp ea r l ea ving i n t erna l ridges s h owi n g th ei r p os iti ons ;

,
a nd i n o th er c a s es
, b ecom e ex trem ely p or ou s T h ou gh it i s
, .
m a n i fes t th a t thi s i s n ot the p roces s of v a s cu l a r d evel op m en t

i n ti s su es t h a t u n fold a ft er p r e—d et er mi n ed typ es s i nc e i n , ,
th ese the d ehi s cen ces or p erfora ti on s o f s ept a occu r b efore

,
s u c h di r ec t a c ti on s ca n h a v e com e i n t o pl a y
; yet it i s s t ill
p os s ibl e th a t the di sa pp ea ra nces o f s ept a w hi ch now a r i se by
r ep etiti on o f th e typ e were es t a bli s h e d i n th e typ e by s u c h
di rect a cti ons . Be thi s a s it m a y h owev er a , ,
s i m u lt a n eou s ch a nge u nd ergon e by these l ongit u di na lly

u nit e d cell s m u s t be oth erw i s e ca u s ed F r a m e w or k s a r e
.
-
f ormed i n them — fra m e work s whi ch c l os ely fitti n g th ei r

-
,
i nner su rf a c es m ay c ons i s t eith er of s u cc es s ive ri ng s or con

, ,
2 84 P H Y S I OL OGI CA L D E V E L OP ME NT .
t i nu ou s spi ra l th rea d s o r net work s or s t ru ct u res b etween

, ,
s pi ra l s a nd ne t wor k s or net w or k s w ith o p en i ng s s o fa r

,
di m i ni s h e d th a t th e c ell s con t a i n i ng th e m a re di s ti ngu i s h ed

a s f enes t r a t ed .T h ei r di fference s om it ted h owev er th es e , ,
s t ru ct u r es h a v e th e co mm on ch a ra c t er t h a t w hil e s u pp or t ,
i ng the coa t s of t he v es s el s th ey a l s o giv e s p eci a l fa c iliti es

,
for th e p a s sa g e of liqu id s b oth th rou gh th e s id e s of t h e

,
t ra ns f orm e d cell s a nd th rou gh thei r u nit ed end s wh ere th es e ,
a re n ot d es t roy e d .
T o a tt em pt a ny phy s ic a l i nt er p ret a ti on of thi s ch a ng e i s

sc a rc ely s a fe : the c onditi on s a re s o c ompl ex T h ere a re .
r ea s on s for s u s p ectin g h ow ev er th a t it a ri s es f rom a v a c u o

, ,
l a ti on o f th e su b s t a nce d ep os it ed on the c ell wa ll I f ra pidly -

.
d ep os it ed a s it i s lik ely t o be a l ong li nes wh ere s a p i s freely

,
s u ppli ed t hi s m a y i n p a s s i n g fro m the s t a t e of a s ol u bl e

, ,
'
c oll oid t o th a t o f a n i n s ol u bl e c oll oid s o c ont ra ct a s t o l ea v e

,
u nc ov ere d s p a ces on th e c ell m em b ra ne ; a n d t hi s c h a ng e

-
,
o rigi na lly c on s eq u en t o n a phy s ic o c h emi ca l a c ti on -

m a y be ,
s o m a i n t a i ned a nd u tili z e d by n a t u ra l s el ec ti on a s t o res u lt ,
i n st ru ct u res of d efi nit e ki nd s re gu l a rly f or m ed i n gro wi n g

,
p a rt s i n a nti c ip a ti on of f uncti ons t o be a ft erwa rd s di s ch ar g ed .
B u t with ou t a ll egi ng a ny s p ec i a l ca u se for thi s m eta m or

,
ph os i s we m ay rea sona bly con c l u d e th a t it i s i n s om e wa y

,
c ons eq u en t u p on th e c a r ryi ng of sa p I f w e e x a m i ne t i s s u es
.
s u ch a s th a t i n th e i nt er i or o f a g row i n g t u rn ip th a t h a s n ot
e t b eco m e s t ri ngy w e m a y i n t he fi rs t pl a c e fi nd b u n dl es
y , , ,
o f e l on g a t ed cell s n ot h a vi n g yet d ev el op e d i n th e m th ose
f enes t ra t ed or reti cu l a t ed s t ru ct u r es by whi ch th e d u ct s a re

ev en t ua lly ch a ra c t eri zed A l ong th e c en t res o f a dj a c ent
.
b u ndl es we m a y fi nd i ncom pl et e li nes of su ch cell s— s om e

th a t a re p a rti a l ly or wh olly t ra ns form ed wi th s om e b et ween ,
th em th a t a r e not t ra n s for m ed I n oth er b u n dl es c ompl et ed

.
,
ch a i n s of su ch t ra ns form ed c ell s a re vi s ibl e A nd th en i n .

,
s till old er b u ndl es th ere a r e s ev er a l com pl et e c h a i n s ru n n i n g

,
sid e by s id e A l l which f a cts imply a m et a m orph os i s of the

.
T HE I NNE R T IS S U E S OF P L A N TS .
l
e on ga t ed c ell s i ndi rectly c a u s e d by
,
the c ontin u ed a cti on of
t he c u rrent s ca rri ed .
§ 2 8 1 H ere h owev er p resent s it s elf a f u rth er p robl em

.
, ,
.
T a ki n g i t a s m a n i fes t t h a t t h ere i s a typi ca l di s t rib u ti on of

s u pp ort i n g ti s s u e a d a pt ed t o m eet t h e m ec h a n i ca l s t r a i n s a
pl a n t i s exp osed to by i ts typi ca l m od e of g row th a nd a l s o ,
t h a t th ere g oes on s p eci a l a d a p t a ti on of th e s u pp or ti n g ti s su e

t o t he s p eci a l s t ra i ns th e i ndivid u a l pl a nt h a s t o b ea r ; a n d
t a ki n g it a s t ol era bly evid ent th a t th e s a p ch a nn els a re -
origi na lly d et ermi ne d by t h e p a ss a ge o f cu r rent s a l ong li nes
o f l ea s t r e s i s t a n c e ; t h er e s till rem a i ns t h e u ltim a t e qu es ti on
T h r ou gh w h a t phy s ic a l a c ti ons a re es t a bli s h ed th es e g en era l

a n d s p e c i a l a dj u s t m en t s o f s u pp orti n g ti s s u e t o t he s t ra i n s
b orne a nd thes e di s t ribu ti ons o f nu t ritive liqu id requ i red t o

,
m a k e p os s ibl e s u c h a dj u s t m en t s ? C l ea rly i f th e e xt ern a l

,
a ct i on s p ro d u ce i n t ern a l rea c ti on s ; a n d if thi s pl a y o f a c ti on s
a nd rea c ti on s res u l ts i n a b a l a nci ng of t h e s t r a i n s by t h e
res i s t a nces ; w e m a
y r a ti ona lly s u s p ect th a t th e i n cid en t
forces a re di rec tly con d u c iv e t o th e s t ru c t u ra l c h a nges by
whi ch th ey a re m et L et u s c on s id er h ow they mu s t w ork
. .
Wh en a ny p a rt o f a pl a n t i s b ent by th e wi nd the ti s s u es ,
on i t s c on v ex su r f a c e a r e s u bj ect t o l on git u di na l t ens i on a nd ,
th es e ex ten ded ou t er l a y er s c omp res s th e l a y ers b en ea th

t h em . S u ch o f th e v es s el s or ca n a l s i n th es e s u bj a cent l a y er s
a s cont a i n s a p m u s t h a v e s o m e o f thi s s a p e xp ell ed
, P a rt o f .
it will be s qu eeze d th r ou gh the m ore or l es s p orou s wa ll s o f

t he c a na l s i nt o th e s u r rou n di ng ti ssu e th u s su pplyi ng it with
,
a s s i m il a bl e m a t er i a l s ; whil e p a r t o f it ,
a n d p r ob a bly th e
l a rg er p a rt will be th ru s t a l ong th e c a na l s l ongit u di na lly

,
u p w a r d s a n d d ow n w a r d s Wh en th e b ra nch or t wig or
.
l ea f s t a lk recoil s th es e v ess el s reli eved f rom p ressu re exp an d

-
, , ,
t o t h ei r or igi n a l di a m et ers A s th ey exp a n d the s a p ru s h es

.
,
ba ck i nt o th em f ro m a b ov e a n d b el ow . I n w hich ev er of
th ese di rec ti ons l ea s t ha s b een exp elled by the c omp res si on ,
28 6 P H Y S I OL OGI CA L D E VE L OP ME NT .
fro m th a t di recti on m os t m u s t retu rn d u ri ng the dil a ti on ;

s eei n g th a t th e force which m ore effici ently res i s ted t h e
t h ru s ti ng b a ck of the s a p i s th e s a m e f orce whi c h u rg es it
i nt o the exp a nd ed v es s el s a g a i n wh en th ey a re reli ev ed f rom
,
p res s u re A t the next b end of the p a rt a f u rth er p or ti on of

.
sa n w ill be s q u eezed ou t a n d a fu r th er p or ti on t h ru s t fo
r
w a r d s a l ong t h e v es s e l s T his ru d e p u mpi ng p r oc es s thu s

.
s er v es for p ro p e lli n g th e s a p t o h eight s w hi ch it cou ld n ot
rea c h by c a pill a ry a c ti on a t the s a m e ti m e th a t it i nc id ent

,
a lly ser v es t o fee d th e p a rt s i n w hich it t a k es pl a c e It .
s t r en gth en s th em t oo j u s t i n p rop orti on t o th e s t ress t o be

, ,
b orne ; s i nce the m ore sev ere a n d the m ore rep ea t ed th e

s t ra i n s t h e g r ea t er m u st be t h e e x u d a ti on of s a p f r om t h e
,
v es s el s or d u ct s i nt o the s u rr ou ndin g ti s su e a nd th e grea ter ,
th e thick en i ng of thi s ti ssu e by s econd a ry d ep os it s By .
thi s s a m e a cti on th e m ov em en t of the s a p i s d eterm i ned

e ith er u p w a r d s o r d ow nw a r d s a c c or di n g t o t h e c on diti ons
, .
Whil e th e l ea v es a re a ctiv e a nd ev a p ora ti on i s g oi ng on f rom

th em th es e o s c ill a ti on s o f the b ra nch es a nd p eti ol es u rge
,
f orwa r d the s a p i nt o th em ; b ec au s e s o l ong a s the v es s el s of

the l ea v es a r e b ei ng empti ed th e s a p i n th e c om p ress ed
,
v es sel s of th e os cill a ti n g p a rt s will m eet w ith l es s res i s t a nce

i n th e di recti on o f the l ea v es th a n i n th e opp os it e di recti on .
B u t wh en ev a p ora ti on c ea s es a t nigh t this w ill n o l onger be ,
t he ca se .Th e s a p d ra wn t o the o s cill a ti ng p a r t s t o s u pply ,
th e pl a c e o f th e ex u d ed sa p m u s t c om e fro m the di rec ti on s

,
o f l ea s t res i s t a n ce A slight b r eeze w ill b ri ng it b a ck fro m

.
th e l ea v es i nt o th e gen tly s wa yi n g t wi gs a s t ron ger breeze

-
,
i nt o th e b en di ng b ra nch es a ga l e i nt o th e st ra i ned st em a nd
,
r oot s— ro ot s i n whi c h l o n git u di n a l t en s i on p rod u c es in a n ,
o th er wa y th e s a m e ef
,
fect s th a t t ra ns v ers e t ens i on d oes i n
th e b ra n ch es .
T w o p os s ibl e mi si nt erp ret a ti on s m u s t be gu a rd ed a g a i nst .
I t is n ot t o be su pp osed th a t thi s f orce p u mp a cti on ca u se s -
m ov em ent o f the s a p t owa rds one p oi nt r a th er th a n a noth er

i t i s simply a n a i d t o i ts m o v em ent F rom th e s t ock of s a p
.
THE INNE R T IS S U E S OF P L A NT S .
28 7
di s t ribu t ed th rou gh the pl ant m ore or l es s is ev erywh ere

,
b ei n g a b s t ra c t ed— h ere by ev a p ora ti on here by th e u nfold ,
i ng of th e p a rt s i nt o th ei r t ypi c a l s h a p es h ere by b ot h ,
.
T he r es u lt i s a t en s i on on the cont a i ned liqu id col u m n s w hi c h ,
i s g rea t es t now i n thi s di rec ti on a nd n ow i n t h a t T his .
t en s i on i t i s w hi ch m u s t be r ega r d ed a s th e force th a t de
t erm i nes th e cu rrent u p wa rd s or d ownw a rd s ; a nd a l l whi ch
th e m ec h a ni c a l a cti ons do i s t o f a cilit a t e th e t ra n s fer t o th e
pl a c es of g rea t es t d em a n d H ence it h a pp ens th a t i n a pl a n t
.
p rev ente d f rom os cill a ti n g bu t h a vi ng a typi ca l t end ency to

,
a ss u m e a ce r t a i n h eigh t a n d b u lk t h e d em a n d s s et u p by i t s
,
u n f oldi n g p a r t s w ill s till ca u se c u rren t s ; a n d th ere will still
be a lt ern a t e a s cent s a nd d es cen t s a ccor di n g a s th e v a ryi ng

,
c on di t i o ns c h a n g e t h e di r ec ti on of g rea t e s t d em a n d — th e
on ly di f fere n c e b e i ng th a t i n th e a b s ence o f os c ill a ti on s th e

, ,
g rowth w ill be l ess vig or ou s . S imil a rly it m u s t n ot ,
be su pp os ed th a t m ec h a ni ca l a c ti on s a re h ere a ll eg ed t o be
the s o l e c a u s es o f w oo d f orm a ti on i n th e i n divid u a l pl a n t
-
.
T h e t en d enc y of th e i ndivid u a l pl a n t t o for m w ood a t pl a ces

wh ere w ood h a s b ee n h a bitu a lly f orm ed by a n ces t ra l pl a n t s ,
i s m a ni fes tly a ca u s e a n d i n d eed th e chi ef c a u s e

, , , I n thi s .
,
a s i n a l l o th er c a s es , i nh erit ed st ru c t u res rep ea t th em s elv es

i rres p ec tiv e of th e ci rcu m s t a nc es of th e i n dividu a l : a b se nce
o f th e a pp rop r i a t e c on diti on s res u lti ng s imply i n i m p erfe ct
rep eti t i on o f t h e s t ru c t u res H ence th e f a ct th a t i n t ra i ned

.
t rees a n d h oth ou se s h ru b s d ens e s u b s t a nce i s s till l a rg ely

,
d ep os it ed ; th ou gh n ot s o l a r g ely a s w h er e the n or m a l m e
ch a ni ca l s t ra i n s h a v e a c t ed H en c e t o o th e f a ct th a t i n
“
.
, , ,
s u ch p l a n t s a s t h e E l eph a n t s foot o r th e Wel wi ts chi a m i m

’
-
bi l l s whi c h for u n t ol d g enera ti ons ca n h a v e u n d erg on e n o

,
os cill a ti on s th er e is a n e xt en s iv e f orm a ti on of w oo d
, ( th ou gh
n ot t o a ny c on s id er a bl e h eigh t a b ov e t h e gr ou n d
) i n rep eti ,
ti on of a n a n ces t r a l typ e : na t u r a l s el ecti on h a vi ng h ere m a i n

t a i n ed th e h a bit a s s ecu ri ng s om e oth er a dv a n t a g e th a n th a t
o f su pp ort .
S till it m u st be b or ne i n mi nd th a t th ou gh i nt ermitt ent

,
28 8 P H Y S I OL OGICAL D E VE L OP ME NT .
m ech a n i ca l s t ra i n s ca nnot be a s signed a s th e di rect c au s es of

th es e i nt erna l di fferenti a ti ons i n pl a n t s th a t a re a rt ifici a lly
s h e lt e r e d or su pp ort e d t h ey a re a s s i gn a bl e a s th e i n di rect
,
ca u ses ; s i n ce t h e i nh eri t e d s t ru c tu r es rep ea t e d a p a rt fr om

,
s u c h s t ra i n s a re t h e m s elv es in t er p ret a bl e a s a ccu m u l a t e d

,
res u lt s o f su c h st ra i n s a c ti n g on s u cces s iv e g en era ti on s o f
a nces t ra l pl a n t s T hi s will b ec om e c l ea r on com bi n i ng t he

.
s ev e r a l th rea d s o f t h e a r gu m en t a n d b r i nging it t o a cl os e ,
whi c h w e m ay now do .
§ 28 2 . T o p u t t he p era tiv e a
c o- o cti on s i n th ei r a ct u a l
or d er w ou ld r eq u i r e u s t o con s id er th em a s w orki n g on i n
,
d i vi du a l s s m a ll m odific a ti on s t h a t b eco m e con s pi cu ou s a n d

d efin it e only by i n h erit a n ce a nd gra d u a l i ncrea s e ; bu t it will
a i d ou r c om p reh ens i on with ou t l ea din g u s i nt o err o r i f w e ,
s u pp os e the w h ol e p roces s r es u m ed i n a si ngl e conti nu ou sly

exi s ti ng pl a n t .
A s th e pl a n t erect s th e in t egr a t e d s er i es of f rond s w h os e

u n it ed p a rt s for m i t s r u di m en t a ry a xi s t h e i n c rea s i n g a rea
,
’
o f fr on d su r fa ce exp ose d t o t h e s u n s r a ys ent a il s a n i n crea s
-
.
i ng d ra u ght u p on th e liqu id s c ont a i n ed i n t he ru dim enta ry

a xi s . The c u rren t s o f s a p s o p ro d u c ed onc e es t a bli s h ed a l ong
,
c erta i n li nes of c ell s th a t off er l ea s t res i s t a nce rend er th em ,
by th ei r c on ti nu ou s p a s s a g e m or e a nd m ore pe rm ea bl e T hi s .
es t a bli s hm en t o f c h a nn el s i s a id e d by t h e w i n d E a ch ben d .
p rod u ced by it w hil e yet th e ti s s u e i s u ndi fferenti a ted ,
s q u eezes t ow a r ds th e pl a c e of g r owt h a nd e v a p ora ti on t he
liqu id s th a t a re p a s s i ng by osm os e f r om cell t o cell ; a nd

w h e n th e li n es of m ov em ent b ec om e d efi ned ea ch b en d h elp s , ,
by forci ng th e liqu id a l ong th es e li nes t o rem ov e ob s t ru cti ons

,
a n d m a k e c on ti nu ou s c a n a l s A s fa s t a s this t ra n s fer o f sa p
.
i s fa cili ta ted s o f a s t i s the pl a n t ena bl ed fu rt h er t o ra i s e i t

,
s el f a nd a dd to i ts a s s imil a ti ng s u rfa c es ; a nd s o f a st do the

,
t ra ns v ers e s tra i ns bec omi ng gr ea ter giv e m ore effici ent a i d

, ,
.
T he c a n a l s thu s for m e d ca n be n eith er i n th e cent re of th e

ru dim ent a ry a xis nor a t its s u rf a c e : for a t neith er o f th es e
THE i NNE R TIS S U E S OF P L A NT S .
289
pl a c es c a n the t ra ns v ers e s t ra i ns p rod u c e a ny c on s id era bl e

c om p res s i ons T h ey m u s t a r i s e a l on g a t ra c t b e t ween th e
.
o u ts id e o f t h e a xi s a nd i t s c or e — a t ra ct a l ong which th ere
o cc u r th e sev er es t s qu eezes b etwee n th e s t re tch e d ou t er l a y er s
a nd t h e i nt ern a l m a s s Ju s t th a t di s t rib u ti on which w e fi nd

.
,
i s th e di s t ribu ti on w hich th es e m ech a nic a l a cti on s t en d t o

es t a blish .
A s th e pl a nt g a i ns i n h eight a n d a s t he m a s s of its f o li a ge
,
a cc u m u l a t es th e s t r a i ns th row n u p on i t s a xi s a nd es p ec i a lly
, ,
the l ower p a r t o f its a xi s ra pidly i n c rea s e , S u pp os i ng t he.
f orms t o rem a i n s i m il a r the s t ra i n s mu s t i nc rea se i n the ra ti o

,
o f t he c u b es of th e di m en s i on s ; or ev en i n a s om ewh a t high er
ra ti o. One c on sequ en c e mu st be th a t the c omp res s i ons to

whi ch t he v es sel s a t th e l ow er p a rt o f th e i n cipi ent s t em a re
s u bj ect be c o m e g rea t er a s fa s t a s th e h eight to w hich the s a p
,
h a s t o be ra i s ed b ecom es g rea t er ; a nd a n oth er cons equ enc e

m u s t be th a t th e l oc a l ex u d a ti on of s a p p rodu c ed by t he
p res s u re i s p r op orti ona t ely a u g m ent ed H en ce the m a t eri a ls
.
for i nt er s titi a l nu t r iti on b ei n g th ere su ppli ed m ore a bu n

da ntl y we m a y exp ec t thi ck en i ng o f the s u rr ou n ding ti s s u es
,
t o s h ow it self th ere fi rs t : i n oth er wor d s w o od w ill be ,
f orm ed rou nd the v ess el s of the l ower p a r t of th e i ncipi ent

s t ern
. Th e resu l ti ng g rea t er a bility of thi s l ower p a rt o f the
s t em t o b ea r s t ra i ns r end e r s p os s ibl e a n i nc r ea s e o f h eight
, ;
a nd w hil e a ft er a n i n c r ea s e o f h eight th e l owes t p a rt be
com es s till fu rth er s t ra i ned a nd still f u rth er thi c k en s

, th e ,
p a rt a b ov e it exp os e d t o lik e a cti on s u nd erg oes a lik e

, ,
thick eni n g T his i n d u ra ti on w hil e it s p rea d s u p w a rd s a l s o

.
, ,
s p r ea d s ou t wa r d s A s f a s t a s the ru d e cyli nd er o f d en s e
.
m a tt er f orm ed i n thi s w a y b egi ns t o i ncl os e t he origi na l

,
vess el s it b egi ns to pl a y the p a rt o f a res i s t a nt m a s s whi c h

, ,
m ore a n d m ore p rev ent s th e c on t a i ned v es sels f rom b ein g

s q u ee z e d ; w hil e b et w een it a n d th e ou t er l a y er s t h e grea t es t
comp res s i on oc cu rs a t ea ch b en d T hu s a t th e s a m e ti m e
.
t h a t th e origi n a l v es s el s b ec om e u s el es s th e p eriph era l c ell s

,
of the d ev el o i n g w o od bec om e th os e which h a v e th ei r liqu id

p
65
29 0 P H Y SI OL OGI CA L D E V E L OP ME NT .
c ont ents qu eezed ou t l ongit u di na lly a nd l a t era lly with i n

s
c rea s i ng forc e ; a nd c on sequ en tly a mid th em a re form e d

, ,
new s a p ch a nnel s fro m w hi c h th ere i s the m o s t a c tiv e l oca l

-
,
e x u d a ti on p r od u ci n g t h e g rea t es t d ep os it o f d en se m a tt er
, .
T hu s fu s i ng t ogeth er a s it were th e i n divid u a liti es o f

, ,
s u cc es s iv e g enera ti on s o f pl a n t s a nd rec og ni z i ng a s
, al l
imp ort a nt th a t f a cilit a ti on of th e p roc ess which n a t u ra l

s el e cti on h a s a l l a l ong giv en we a re e n a bl e d t o i n t erp ret t h e
,
chi ef i nt erna l di fferenti a ti ons of pl a nt s a s c on s eq u ent on a n

e qu ilib ra ti on b et ween i nn er a n d ou t er f or c es H ere i n d eed .
, ,
w e see ill u s t ra t ed i n a w a y m or e th a n u s u a lly ea sy t o f oll ow ,
the ev ent u a l b a l a nci n g of ou t er a cti ons by i nner r ea cti on s .
T he rel a ti on b et ween th e d em a n d for liqu id a n d th e f orm a ti on

o f ch a nnel s th a t s u pply liqu id a s w ell a s th a t b e t ween th e
,
i ncid enc e of st ra i ns a nd the d ep os it of su b s t a n ce which res i s t s

st ra i n s a re a m ong the cl ea rest sp ec i a l ex a m pl es of th e g enera l
,
t ru th th a t the m ovi ng equ ilib ri u m of a n org a ni s m i f n ot ,
ov erth row n by a n i ncid en t f orce m u s t ev en tu a lly be a dj u s t ed

,
to it .
T he p roc ess es h ere t ra c ed ou t a re of c ou rs e n ot t o be , ,
t a k en a s the only differenti a ti ng p roc es ses to which the i nner

tis su es of pl a nts h a v e b een s u bj ect B es id es th e chi ef ch a ng es
.
we h a v e c ons id ered v a ri ou s l es s c onspic u ou s ch a ng es h a v e

,
t a k en pl a c e T h es e mu st be p a s sed ov er a s a ri s i ng i n wa ys
.
t o o i nv olv ed t o a dmit of sp ecific i nt erp ret a ti on s ; e v en s u p

p os i ng them t o h a v e b een p rod u c ed by c a u ses of th e ki nd
a s s ign ed . Bu t the p rob a bility or ra th er i nd eed th e c ert a i nty
, ,
i s th a t s om e of th em h a v e not b een s o p r od u ced H ere a s .

,
i n nea rly a l l oth er c a s es i ndi rect equ ilib ra ti on h a s w ork ed i n

,
a i d of di rect eq u ilib ra ti on a n d i n m a n y c a s es i n di rect equ i l i
b r a ti on h a s b een th e s ol e a gency B es id es a s c ribi ng t o

.
n a t u r a l s el ecti on the r i s e of v a r i ou s i nt ern a l m o difi c a ti on s
o f o th er cl a ss es th a n th os e a b ov e t rea t ed we m u s t a s c rib e ,
s om e ev en of th es e t o na t u ra l s el ecti on I t i s s o w ith th e .
d ens e d ep os it s whi ch f orm th orn s a nd th e s h ell s of nu t s

thes e c annot h a v e res u lt ed f rom a ny i nner rea cti ons imm e
T HE I NNE R T IS S U E S OF P L A NT S .
di a tel y c a ll ed f orth by ou t er a cti ons ; but mu s t h a v e res u lt ed

m edi a t ely th rou gh the eff ect s of su ch ou t er a cti on s on the
sp ec i es
. L et it be u nd erst ood th eref ore th a t t he di ffer
, ,
enti a t i ons t o which t h e f oreg oi ng i nt erp re t a ti on a ppli es a re

,
on ly th o s e m os t c on spic u ou s on es which a re di rectly re l a t e d
t o the m os t c ons pi cu ou s i n cid ent f or ce s. T h ey m u st be t a k en

a s i n st a nc es on th e st rength o f w h i c h we m a y c on c l u d e th a t
oth er i nt ern a l di fferen ti a ti ons h a v e h a d a n a t u r a l g enesis ,
th ou gh in w a ys th a t we c a nn ot t ra c e .
C H A P TE R V .
PH Y S IO L O GI CA L I N TE GRA T I ON IN P L A N TS .
28 3 . A OO D d ea l h a s b een impli ed o n this t opic i n the

G
p rec edi ng ch a pt ers H ere h owever we m u st for a b ri ef

.
, ,
sp a ce t u rn ou r a tt enti on imm edi a t ely to it .
P l a nt s do not displ a y i nt eg ra ti on i n s u ch disti nct a nd

m u ltipli ed wa ys a s do a nim a l s Bu t i ts a dv a nce m ay be
.
t ra c ed b oth di rectly a nd i ndi rectly— di rectly i n the i ncrea s

i ng c o—ordi na ti on of a cti ons a nd i ndi rectly i n the effect of
,
thi s u p on the p ow ers a nd h a bits .
L et u s g rou p th e fa ct s u nd er th es e h ea d s : a sc endi ng i n
b oth c a s es f rom the l ower to the high er typ es .
§28 4 . The i n f er i or A l g ae a l
,
on g w ith littl e u nlik en ess of
p a rt s sh ow u s littl e m u tu a l d ep end enc e of p a rt s H a vi ng
,
.
s u r f a ces s imil a rly ci rcu m s t a nc ed ev erywh ere m u ch phy s i o

,
l ogi ca l divi s i on of l a b our c annot a ri s e ; a nd th eref ore th ere

c a nno t be m u ch phy s i ol ogic a l u nity . A m ong the s u p eri or
A l gae h owev er the di fferenti a ti on b et ween th e a tt a ch ed p a r t
, ,
a nd the f ree p a r t i s a cc omp a n i ed by s om e i nt eg ra ti on T h ere .
i s evid en tly a c ert a i n t ra n sfer o f m a t eri a l s which i s d ou btl ess

,
f a cilit a t ed by th e el ong a t ed f orms of the cell s i n the st em ,
a nd p rob a bly l ea ds t o th e f o rm a ti on o f d en se ti s s u e a t th e
pl a ces of g rea t es t st ra i n i n a wa y a ki n t o th a t rec ently ex

,
pl a i ned i n oth er c a s es And wh ere th ere i s thi s co ordi na

.
-
ti on of a cti ons th e p a rt s a re s o fa r m u t u a lly d ep end ent th a t

P H Y S IOL OGICAL INT E GRA T ION IN P L A NTS .
ea ch di es i f d et a ch ed f rom the oth er T h a t th ou gh the .
o r g a ni z a ti on i s s o l ow neith er p a rt ca n r ep ro d u ce t h e o th er
a nd s u r viv e by s o d oing is p rob a bly du e t o the ci rc u m s t a n ce

,
tha t ne ither p a rt c ont a i ns a ny c ons id era bl e s t ock of untra ns

f orm ed p r ot opl a sm ou t of which new ti ss u es m a y be pr o
,
du ced .
F ungi a nd L ich en s
p res ent no v ery sign ific a nt a dv a nc es
o f i n t e gra ti on . We will th eref ore p a ss a t once t o the
A rch eg o ni a tes . I n th os e o f th em which eith er a s s i ngl e ,
fr on ds or st ri ng s o f f ron d s S p r ea d ov er s u r f a c es a n d w hich
, , ,
roo ti n g th em selv e s a s th ey sp rea d do n ot n ee d th a t ea c h p a rt

,
s h ou ld rec e iv e a i d f ro m r em ot e p a r t s th er e i s n o d ev e l o p ed
,
v a scu l a r sy s t em s ervi ng to f a c ilit a t e t ra ns fer of nu t ri m ent :

th e p a r t s be i ng littl e di fferenti a t ed th ere i s bu t lit t l e in t eg ra
ti on B u t a l ong wi th a ssu mpti on of the u p right a ttit u d e a nd
.
the a c c omp a nyi ng s p eci a li z a ti on s p rod u ci ng v ess els for di s

,
t rib u ti ng s a p a nd h a r d ti ss ue for givi ng m ech a n ic a l su pp ort ,
there a r i s es a d ecid ed phy s i ol ogic a l divisi on of l a b ou r ren d er

i ng the a e r i a l p a rt d ep en d ent on the imbedd ed p a r t a nd the
i m bedd ed p a rt d ep end ent on the a eri a l p a r t H ere i n .
,
d eed a s el s ewh ere th es e c oncom it a nt ch a nges a re bu t two

, ,
a s p ect s of t he s a m e ch a ng e A lw a ys th e ga i n of p ower t o di s
.
c h a rg e a s p ec i a l f un c ti on i n v olv e s a l o s s of p ower t o p erfor m
o th er fu n cti on s ; a n d a l wa y s th eref or e i nc rea se d m u t u a l de

, ,
p en d en c e c on s tit u ti n g phy s i o l o gic a l in t eg ra ti on m u s t k eep ,
p a ce with th a t i nc rea s ed fitti ng of p a rticu l a r p a rt s t o pa rt i cu

l a r d u ti es w hich c on s tit u t es phy s i ol ogic a l di fferen ti a ti on .
Ma ki n g a grea t a dv a nc e a m ong the A rc h eg on i a t es thi s ,
phy s i ol ogic a l i nt egr a ti on rea ch es its clim a x a m ong P haeno

ga m s I n th em we i nt er d ep end enc e throu gh ou t
“
. see
m a sses th a t a re imm ens e A l ong with s pe ci a li z ed a p pl i

.
a n ces for s u pp ort a n d t ra n sfer we fi nd a n e xch a ng e o f a i d a t

,
gr ea t di s t a nces We see roo ts g ivin g the v a st a eri a l grow th

.
a h o ld t ena ci ou s enou gh t o withst a n d vi ol ent wi n d s a nd ,
s u pplyin g w a t er en ou gh ev en d u ri ng
pe r i od s o f d r ou ght ; w e
s ee a s t em a nd b ra n ch es o f c orres p onding st rengt h for u

p
294 P H Y S I OL OGI CA L D E VE L OP ME NT .
h oldi n g t he a ss m g a ns u nd er ordi na ry a nd extra or

i il a ti ng or
di na ry s t ra i ns ; and i n th es e a s simil a ti ng org a ns we s ee

el a b or a t e a ppli a nces for yi eldi ng t o t he s t e m a nd r oo t s the
m a t er i a ls ena bli ng th em t o fu lfil th ei r offic es A s a c on .
s equ ence of w hi ch g rea t er i nt egra ti on a ccomp a nyi ng the

g rea t er di fferen ti a ti on th ere i s a bility t o m a i nt a i n life ov er
,
a n imm en s e p er i o d u n d er m a rk e d vici ss it u d e s .
E v en m ore c on s picu ou s ly ex emplifi ed i n P haan oga m s i s ,
th a t phys i ol ogic a l i nt egra ti on whi ch h old s t ogether the fu nc

ti on s n ot of the i ndivid u a l only bu t of th e sp eci es a s a wh ol e .
T he org a n s o f rep ro d u cti on b oth i n th ei r rel a ti on s t o oth er

,
p a rt s of the i ndivid u a l b ea ri ng th em a nd i n th ei r rel a ti on s to

c orres p on din g p a rts of oth er i n dividu a l s S h ow u s a ki nd of ,
i nt egra ti on c ond u ci n g t o th e be tt er p res erv a ti on of the ra c e ;

a s th os e a l rea dy sp ecifi ed c on d u c e t o the b ett er p res er v a ti on of
th e i n divid u a l I n the fi rs t pl a c e this grea t er c c ordi na ti on

.
,
-
o f fu n cti on s j u st d es c rib ed i t self ena bl es P haenoga m s t o be

,
q u ea t h t o t h e g er m s th ey c a st off st ores of nu t ri m en t
, p o
r ,
tect i ve env el op es a n d m ore o r l ess of org a ni za t i on : s o givi ng

,
them grea t er ch a nces o f rooti ng th em selv es I n the s ec ond .
pl a ce cert a i n di fferenti a ti on s a m ong th e p a rt s of fru cti fica

,
ti on the m ea ni ng o f which Mr D a rwin h a s s o a dmi ra bly ex

,
.
pl a i ned give to th e i ndivid u a l s of th e s p eci es a ki nd of i nt e

,
ra ti on th a t m a k es p o s s ibl e a m u t u a l a id i n th e p ro d u cti on of
g
vig orou s offs p ri ng A nd it is i nt eres ti ng t o ob s erv e h ow i n
.
,
th a t dim orph i s m by which i n s om e c a s es thi s m u t u a l a i d i s

m a d e m ore effici ent the g rea t er d egree of i nt egra ti on i s
,
d ep end ent on the g rea t er d eg ree of differenti a ti on— not s imply

different i a ti on of th e f ru ctifyi ng org a n s f rom oth er p a rt s o f
th e pl a nt b ea r i ng th em bu t di ff eren ti a ti on of th es e f ru c tify
,
i ng o rg a ns f rom t he h om ol og ou s orga n s of neighb ou ri ng i ndi

vi du a l s o f th e s a m e r a c e . A n oth er f orm of this
co or di na ti on o f f u n cti on s w hich c o n d u ces t o th e m a i n t en a nc e
-
o f th e s p ec i es m a y be h ere n a m e d— p a rt ly for i t s i n t ri ns ic
,
i nt erest I refer t o the s t ra nge p roc ess es of mu ltiplica ti on

.
occ u rri n g i n th e g enu s B ryOp hyl l u m I t i s well kn own th a t

.
P H YSI OL OGI CA L I NT E GRA TI ON IN P L A NT S .
t he l
s u cc u en t l ea v es of B ca l yci n u m b orne on f oot s t a lks
.
,
-
s o b r ittl e t h a t th ey a r e ea s ily s n a p pe d by t he w i n d s en d ,
f ort h f rom th ei r ed ges wh en th ey f a ll t o the g rou nd bu d s ,
whi ch root t h em s elv es a n d g r ow i nt o i nd ep e nd ent pl a nt s T he .
corr el a ti on h ere obvi ou s ly f u rt h eri ng the p res erv a ti on o f th e
r a ce i s m ore d e fi n i te ly es t a bli s h e d i n a n oth er s p e ci es o f th e

,
g en u s— B pr ol iferu m T his pl a nt s h ooti ng u p t o a c ons id er

. .
,
a bl e h eigh t a n d h a vi n g a s t e m c ont a i n i ng bu t littl e w oody

,
fib re h a bi tu a lly b rea k s nea r t he b ott o m whil e s till i n fl ower ;

,
a nd i s th u s g ener a lly p rev ent ed f r om ri e ni n g i ts s ee d s The

p .
mu l tipli ca ti on i s h owever s ecu red i n a n oth er wa y B efore

, ,
.
the s t em i s b ro k en y ou ng pl a n t s h a v e b u dd ed ou t f r om the
p edi cel s of th e fl owers a nd h a v e grown to c ons idera bl e l ength s ;
,
a nd on t h e fa ll o f t h e p a ren t t h ey f or thwith c om m en c e th ei r
s ep a r a t e liv es H ere na t u r a l s el ec ti on h a s es t a bli s h e d a

.
rem a r k a bl e ki n d of c o o r di n a ti on be t ween a s p ec i a l h a bit of

-
g rowth a nd d eca y a n d a s p eci a l h a bit of p roli fera ti on

, .
§ 28 5 The a dv a nce o f phy s i ol ogic a l in t eg ra ti on a m ong

.
pl a n t s a s w e a scend to the high er typ es i s i m pli ed by th ei r ,
g rea t er c ons ta ncy o f s t ru ct u re a s well a s by the s t r ict er li m i

,
ta t i on s o f th ei r h a bit a t s a nd m o d es o f li fe C o mpl exity of .
s t ru ctu re i s g ener a lly a ccom p a ni e d w ith a gr ea t er t en d ency

”
t o per m a n en ce i n form s a ys D r
, [ n ow S i r J ] H ook er ; or
.
,
“ ”
c onv er sely ,the l ea s t c o mpl ex a re a l s o the m os t v a r i a bl e .
T hi s i s th e s ec on d a s p ec t u nd er whi ch we h a v e t o c ont em
pl a t e the fa ct s .
The di fferen ces b et w een the s i m pl er A l g ae a n d F u ngi a re

s o feebly m a rk e d th a t b ot a ni s t s h a v e h a d gr ea t diffi cu lty i n
fra mi n g d efi niti on s o f th es e cl a s ses T hi s s t ru c t u r a l i ndefi

.
n i ten es s i s a cco m p a n i ed by f u nc ti ona l i n d efi ni ten es s A l gae .

,
whi ch a re m os tly a qu a ti c i n cl u d e m a ny s m a ll f or m s th a t
,
f requ en t the d a mp pl a ces p referred by F u ngi A m ong F u nga .

,
th er e a re ki n d s whi ch l ea d s u bm erged liv es lik e th e A l ae

g .
Be s id es thi s i n di s ti n ct nes s o f the cl a s ses th ere i s g rea t v a ri a

,
bi l i ty i n th e s h a p es a nd m od es of life o f th ei r s p eci es— a v a ri

29 6 P HY SIOL OGICA L D E VE L OP ME NT .
a bility s o g rea t th a t wh a t w ere a t fi rs t t a k en t o be di fferent

s p ec i es
,
or di ffe re nt g en er a ,
or ev en di f fer ent o r d ers h a v e ,
p r ov ed t o be m erely v a r i eti es of one s p eci es S o i n con s t a nt .
i n s t ru ct u re a re the A l gae th a t S chl eid e qu ot es w i t h a pp rov a l

n
th e opi ni on o f Ku tzi ng th a t
,
th er e a re no s p eci es bu t m erely
”
form s o f A l g ae z a n Opi n i on w hi c h t h ou gh n ow re j ect ed
suf fi c i en tly impli es ext rem e i ndefini t enes s I n a l l whi c h

.
fa ct s w e s ee t h a t t h ese l owes t typ es of pl a nt s littl e di ffer ,
en t i a ted a r e a l so bu t li t tl e i n t eg ra t e d
,
.
A rch eg oni a t es p res ent a lik e rel a t i on be t ween t he s m a ll

sp eci a li z a ti on of f un cti ons whi ch con s tit u tes phy s i ol ogi ca l
di fferenti a ti on a nd the s m a ll com bin a ti on of fu ncti ons which
,
Moss es s a y s Mr
“ ”
con s t i tu t es phy s i ol og ic a l i nt eg ra ti on.
, .
Berk el ey ,
a re n o l es s v a ri a bl e th a n oth er c ryp t og a m s a n d ,
a re th er efor e f requ ently v ery di ffi c u lt t o di s ti n gu i s h N ot .
on ly will t h e s a m e s p eci es exhibit gr ea t div ers ity i n th e s i ze ,
m od e o f b r a nchi ng f orm a nd nerv a ti on of the l ea v es bu t the

, ,
”
ch a ra ct ers of even the p eri s t om e it s elf a re not cons t a nt .
A nd c onc erni ng the cl a s s ific a ti on o f t he rem a i ni ng g rou p ,
F i l i ca l es h e s a y s
,
N ot only i s th ere g rea t di ffi cu lty i n
a rra ngi ng fer n s s a ti s f a ct orily bu t it i s ev en m or e diffi cu l t t o
,
d etermi ne th e limit s of s p eci es .
A fter thi s v a gu eness of s ep a ra ti on a s well a s i nc ons t a n cy

o f s t ru c t u r e a nd h a bit a m on g th e l ow er pl a nt s the s t a bility ,
o f s t ru c t u re a nd h a bi t a n d divi s ibili ty o f g rou p s a m o ng th e
high er pl a nt s a pp ea r rel a tiv ely m a rk ed T h ou gh P h aenoga m s

,
.
a r e m u c h m or e v a r i a bl e th a n m os t b ot a ni s t s h a v e u ntil
l a t ely a ll owed y et the d efiniti ons of s peci es a n d genera

,
'
m a y be m a d e with fa r g rea t er p rec i s i on a nd th e fo r m s a re

,
fa r l es s ca p a b l e o f ch a n ge th a n a m ong C rypt og a m s
,
A nd .
this com p a ra tive fi xi ty of typ e i m plyi ng a s it d oes a cl o ser

, , ,
c ombi na ti on of the c omp onent f u ncti ons we s ee t o b e the , o
a ccomp a nim en t o f th e g r ea t er di f fer enti a t i on o f th os e f un c

ti ons an d o f the s t ru c t u res p er f ormi n g th em T h a t th es e .
ch a ra ct er s a re c or rel a tiv e s i s f u r th er s h own by th e f a c t th a t
th e high er p l a nt s a re m or e res t ri c t ed i n th ei r h a bit a ts th a n

P H Y S IOL OGICAL INTE GRA T ION I N P L A NT S .
t he l ower pl a nts b oth i n s p a ce a nd t ime

, T he m u c h .
”
na rrower d elimit a t i on i n a rea o f a n i m a l s th a n pl a n t s sa y s ,
S i r J H ook er “
. a nd gre a t er r es t ri c ti on o f F a u n a s th a n
,
Fl ora s sh ou ld l ea d u s t o a nticip a t e th a t pl a n t typ es a r e geo

,
-
,
l ogic a lly s pe a ki ng m ore a nci ent a nd p erm a n en t th a n the

,
higher a ni m a l typ es a re a nd s o I bel i eve th em t o be a nd I

, ,
wou ld ext en d th e d oc t ri ne ev en t o pl a n ts o f highly com pl ex

”
s t ru ct u re T h os e cl a s ses a nd ord ers whi ch a re t he l ea st
.
”
c om pl ex i n org a ni za t i on a re the m os t wi d ely di s t rib u te d .
28 6 . T hu s
th a t whi c h the gen era l d oct ri ne of ev ol u ti on
l ea d s u s to a nticip a t e w e fi nd i m pli ed by the f a ct s T he
, .
physi ol ogic a l divi s i on of l a b ou r a m ong p a rt s ca n g o on only ,
i n p rop orti on t o th e m u t u a l d ep end enc e of p a rt s ; a n d the

m u t u a l d ep end en c e o f p a r t s ca n p ro gress on ly a s f a s t a s th ere
a ri s e s t ru c t u res by w hi c h th e p a rt s a r e ef fi c i ently c o m bi ned ,
a n d th e m u t u a l u tili z a ti on o f th ei r a ct i on s m a d e ea s y .
T o s a y d efin it ely by wh a t p roces s i s b r ou ght a b ou t thi s

co ordi na ti on of f u nc ti on s whi c h a c c om p a n i es th ei r s p ec i a l i za
-
ti on i s h a rdly p ra ctic a bl e D i rect a nd i n di rect eq u ilib ra ti on

,
.
d ou btl ess c o op era t e i n es t a bli s hi ng it

-
We m a y s ee for
.
,
ex a mpl e th a t e v ery i n c rea s e of fit nes s for fun c ti on p r o d u ce d

,
i n the a eri a l p a r t of a pl a nt by light a s w ell a s ev ery i n c rea s e

,
of fit nes s for fu n c ti on p ro d u c e d i n i ts imb edd ed p a rt by t he
di rect a cti on of th e m oi s t ea rth mu s t c ond u c e t o a n i nc rea sed

,
c u rrent of the liqu id ev a p ora t ed f r om the one a nd s u ppli ed

by the oth er— mu s t s erv e theref ore t o a i d th e form a ti on of
, ,
sa ch n n l i n t h w a y s a l a dy d es c r ib e d t h a t i s — m ust
p a e s e r e ;
-
s erv e t o d ev el o p th e s t ru ct u res th r ou gh w hi c h m u t u a l a i d
o f th e p a rt s i s giv en : th e a dditi on a l di f feren ti a ti on t end s

i m m edi a t ely t o b ri ng a b ou t the a dditi on a l i n t egr a ti on C on .
t ra ri wi se it i s obvi ou s th a t a n i n t er d ep en d enc e s u c h a s w e
,
-
s ee b et w een th e s ec r eti on o f h oney a n d th e f ertili z a ti on o f
germ s or be t ween th e d ep os it of a lb u m en i n th e cotyl ed ons

,
o f a n e m b ry o pl a nt a n d i t s s u b s eq u en t s t riki n g r o o t
-
is a ,
ki n d of i nt egr a ti on i n th e a cti ons o f th e i ndivid u a l or of the

298 P H Y SI OL OGICA L D E VE L OP ME NT .
sp eci es which no di fferenti a ti on h a s a di rec t t end ency t o

,
ini ti a t e H enc e we m u s t reg a rd the t ot a l res u lt s a s du e t o a

.
pl exu s of i nfl u ences a cti ng s i m u l ta neou s ly on the i ndivid u a l

a nd on t he s p eci es : s om e c hi efly a ffecti ng th e one a nd s om e
chi efly a ffecti ng the other .
[ N OT E — I n N a tu r e for Ju ne 1 1 , 1 8 9 6, D r . Ma xwell M a s
t ers ,
in y on P l a nt B reedi ng n a m es a n i n s t ru ctiv e
a n es s a ,
fa ct con cern i n g th e p r od u ct i on of v a r i eti es by s el ecti on o f

'
slightly div ergent f orm s H e s a y s .
T o the u nt r a i ne d eye the p ri m or di a l diff erenc es not ed a re

,
oft en v ery s light ; ev en th e b ot a ni s t u nl es s hi s a t t en t i on be ,
s p ec i a lly di rec t e d t o th e m a tt er f a il s t o s ee mi nu t e di ffer en ces

,
whi c h a re p erc eptibl e enou gh t o th e ra i ser or h i s wor k m en .
N or m u s t it be th ou gh t t h a t th es e v a ri a ti ons di ffi cu lt a s th ey ,
a r e t o r ec ogn i se i n t h e b egi nn i ng a re u nimp o rt a n t On th e

,
.
contra r y t h ey a r e i n t er es ti n g phy s i ol o gic a lly a s t h e p ot enti a l

, , ,
o rigi n of new s p eci es a nd v e r y o ften t h ey a re c o m m er ci a lly

,
v a l u a bl e a l s o T h ese a pp a ren tly t rifli ng m orph ol ogi ca l dif

.
feren ces a re o ften a s s oci a t ed w ith phy s i ol ogi ca l v a ri a ti ons

whi c h r en d er s o m e v a ri eti es s a y of wh ea t m u ch b ett er
, ,
ena bl ed t o r e s i s t m ild ew a n d di s e a se g en era lly t h a n o th er s .
S om e a g a i n p r ov e t o be b ett er a d a pt ed for cert a i n s o il s or

, ,
for s om e c li m a t es th a n oth er s ; s om e a re l es s li a bl e t o i n j u ry
”
f rom p red a t ory bi rd s th a n oth ers a nd s o on , .
T h u s we a re s h ow n th a t t o a m u c h g rea t er d eg ree t h a n
,
might be s u pp osed mi nu te ch a nges of f or m s a nd fu ncti ons

,
i n on e p a r t o f a pl a nt a re c or rel a t ed w i th ch a ng es o f fo rm s
a nd f u n c ti on s th rou gh ou t it T he i nt er d ep end ence— th a t i s
.
-
t o s a y th e phy s i ol ogi ca l i nt eg ra ti on— i s v ery cl os e a t the

,
s a m e ti m e th a t i t i s v er y co m pl ex .
H ere whil e na m i n g th es e fa c t s i n ill u s t ra ti on o f phy s i o

l ogic a l i nt eg ra ti on i n pl a nt s I n a m e th em beca u s e thev
ill u s t ra t e a n imp ort a n t t ru th b ea ri ng u p on the genera l q u e s
ti on o f h eredity whi c h I h a v e d ea l t w i th i n A pp endix G a nd ,
to whi c h I n ow es p eci a lly dr a w a t ten ti on ]

CH A P T E R VI .
I
D F F E RE N T A T I IO N S BE T W E E N T H E O U TE R A ND INNE R
T I SS UE S O F A N I M A L S .
§ 28 7 WH A T wa s s a id res p ecti ng the p rim a ry physi ol ogic a l

.
d iff erenti a ti on in pl a n t s a ppli es with littl e b ey ond ch a nge o f

,
t er m s t o a ni m a l s A m ong P r otoz oa a s a m ong P r otop hy ta

.
, ,
the fi rs t d efi nit e c ont ra s t of p a rt s i s t ha t b et wee n ou tsid e

a nd i n s id e T he sp eck of j elly or s a rc od e whi c h a pp ea rs
.
t o c on s tit u t e t he s impl es t a n im a l p rov es on cl oser ex a mi n a

, ,
ti on t o be a m a ss of s u b s t a nce c on t a i ni ng a nu cl eu s— a
,
p eripl a st i n th e mid s t of w hich there i s a mi nu t e end opl a s t ,
consi s ti ng of a s ph er i c a l m emb r a ne a n d i t s co nt en t s .
T hi s p a r a ll el only j u s t t ra c ea bl e a m on g th e s e Rhi z op od s
, ,
whi ch a re p er pe t u a lly c h a ngin g the di s t r ibu ti on o f t h ei r ou ter

s u b s t a n ce bec om es a t o nce m a rk ed i n t h ose high er P r o tozoa
,
whi ch h a v e fix e d sh a p es a nd m a i nt a i n c on s t a n t r el a ti ons
,
b et ween thei r s u r fa ces a n d th ei r envi r onment s I nd eed th e .
Rhiz op od s th ems elv es on p a ss i ng i nt o a s t a t e of qu i es cen c e

,
i n whi c h th e rel a ti ons o f ou t er a n d i nn er p a rt s a re fix ed ,
bec om e ency s t ed : there i s for med a h a r d ened ou t er c oa t

di fferent f rom the m a t ter whi ch i t c ont a i ns A nd w h a t i s .
h ere a t emp ora ry ch a ra ct er a ns weri ng t o a t em p or a ry defi ni te

ness o f c on diti on s i s i n th e I nfus ori a a c on s t a n t c h a ra ct er
, ,
a n sw eri n g t o d efi nit e con diti on s t h a t a re con s t a n t E a c h of .
t h ese m i nu t e crea t u res th ou gh n ot coa t ed by a di s ti nc t

,
m em b ra ne ha s a n ou t er l a yer of exc ret ed s u b s t a nce f or mi ng

,
a d elic a t e cu ti cl e .
28 8 . T he ea r ly est a bli s hm ent of thi s p rim a ry t

c on ra s t of
2 99
3 00 P H YS IOL OGI CA L D E VE L OP ME N T .
ti s su es a ns werin g t o t his p r i m a ry c ont ra st of c onditi ons i s ,
no l es s cons pi cu ou s i n a ggr eg a t es of th e sec ond or d er T he .
f eebly i nt eg ra t ed u nit s of a S p onge with i ndivid u a liti es s o

-
,
lit tl e m erg ed i n th a t of the wh ol e th ey f orm th a t m os t of

th em s t ill ret a i n thei r s ep a ra t e a ctiviti es neverth el es s sh ow
,
u s i n th e u n lik enes s th a t a ri s es b etween th e ou t er m os t l a y er

,
a n d t h e c o n t a i n ed m a s s th e effect o f c on v erse with un lik e

,
c on di t i on s
. T hi s ou t erm os t l a yer is c omp o s ed of u nits s o m e
w h a t fl a tt ened a nd u ni ted i n t o a c onti nu ou s m e mb ra ne— a
ki nd of ru di m ent a ry s kin .
S ec on d a ry a gg rega t es i n which th e liv es of th e u nits a re

m ore su b ordin a t e t o the life of th e wh ol e c a rry thi s di s ,
ti ncti on fu r th er T h e l ea di ng physi ol ogic a l t ra it o f ev ery

.
c oel ent era t e a ni m a l i s the divi s ibility of i ts s u b s t a nc e i n t o

en d od er m a nd ect o d er m— the p a rt n ext the f oo d a n d th e
p a rt next the envi ronm ent Fig 1 4 7 . . rep resenti n g a
p orti on of the b ody wa ll of a H ydra s een in s ec ti on giv es

-
,
s om e id ea o f t hi s fun d a m en t a l diff erenti a ti on T h e c rea t ure

.
con s i s t s of a s impl e s a c th e c a vity o f w hi ch i s i n c om m u ni

,
c a ti on with the s u rr oun di ng wa t er ; a nd h ence the u n lik e

n es s b et ween th e o u t er a nd in n er l a y ers h a s n ot bec o m e
grea t T he es s ent i a l c ont ra s t i s th a t bet ween the di fferen

.
t i a ted p a rt s of wh a t wa s origi na lly the s a m e p a rt — a u nif orm

m emb ra ne c omp os ed of j u xt a p os e d c ells .
For h ere i nd eed we a re s h o wn u nmist a k a bly h ow the

, ,
p rim a ry con t ra s t o f s t ru ct u res foll ows u p on the p rim a ry con

t ra s t o f c onditi ons .T h e or di n a ry f or m from which l ow
t y p e s of th e M e ta zoa s et ou t i s a h oll ow sph ere form ed of
,
cell s p a ck e d s id e by sid e — a bl a s t u l a a s it i s c a ll ed : a l l th ese

,
cel l s bei ng simi l a rly exp os ed t o t he envi ronm ent T he .
bl a stu l a p res ently ch a ng es i nt o wh a t is c a ll ed a ga s t ru l a— a

f or m res u lti ng f rom the i nt r oversi on o f one of the sid es o f
the bl a s t u l a . I f th ere be t a k en a s m a ll b a l l of vu l ca ni zed
i ndi e rubb er s a y a n i nch or m ore i n di a m et er an d h a vi ng a
-
, ,
h ol e i n i t th r ou gh whi ch the a i r m a y es ca p e a nd if one s id e ,
o f it be th ru s t i nwa r ds s o a s t o p rod u c e a cu a nd if t he
p ,
T HE OU TE R A ND I NNE R T ISS U E S OF A NIMA L S .
3 01
w id e Op eni ng of the cu p be s u pp os ed to c ont ra ct thu s ,
b ecomi ng a na rrow op en i ng th ere will res u lt s om ethi ng lik e

,
the g a s t r u l a f orm Ma ni fes tly th a t p a rt of the origi na l l a yer

.
whi c h h a s b ec om e i n t erna l i s di fferently con diti oned f ro m

the res t whi c h re m a i ns ext ern a l : th e one c onti nu in g t o h old
con v er s e w ith t h e f or c es of th e en vi r on m en t whil e th e o th er ,
b egi ns t o h old c onv ers e w ith the nu t r itiv e m a tt er s t a k en

i nt o the sa c f orm ed cha mbe r— the a rchenteron or p ri mitiv e
-
st om a ch I nt eres ti ng evid enc e of the p rimitiv e ext erna lity

.
of the dig estiv e ca vity i s yi eld ed by t h e f a ct th a t w h erea s
the bl a s t u l a co ns i s t ed of cili a t ed cell s a n d wh erea s the cili a

,
ti on p ers i s ts th rou gh ou t li fe on the ou t er l a yer or p a rt s of it , ,
i n s u nd ry l ow typ es— ev en in s om e C hae t op o d s— i t p ers i s ts

a l s o on th e a lim en t a ry t r a ct o f s u n d ry l ow typ es : n ot on ly
i n the H ydm bu t c o mm only i n N em erti nes i n s Om e P l a ty ,
hel m i n thes a n d e v en i n s om e l ee ch es
,
.
Bes id es b ei ng ena bl ed thu s to u nd ers t a n d h ow a n a ggre

g a t e of A m aebu f or m u ni t s ori gi na lly c on sisti ng of a si ngl e
-
,
l a y er m a y p a s s i nt o a n a ggrega t e con si s ti ng of a d ou bl e
,
l a yer ; we m a y a l s o u nd ers t a nd u nd er wh a t i nfl uenc es th e

t ra n s iti on t a k es pl a ce I f the h a bit which s o m e o f t he
.
p rim a ry a gg reg a t es h a v e of wra ppi ng th ems elv es r ou n d

,
m a s s es o f nu t rim ent i s f oll owed by a secon d a ry a ggrega t e

, ,
there will na tu ra lly a ris e j u s t th a t re diff erenti a ti on whi ch ~
the H ydra s h ow s u s .
28 9 . cc ou nt o f th e p rim a ry differenti a ti on c a rri es

T hi s a
u s on ly h a l f w a y t ow a r d s a t ru e c on ce pti on o f the di s ti nc ti on
-
b et ween ou t er a nd i nner ti s su es T h ou gh u s i n g w ord s i n

.
,
thei r cu rrent s enses thi s i nt rov ert ed p a rt of the p ri m itive

,
l a yer h a s b ec om e i n t ern a l i n cont ra s t with th e rem a in d er ,
which c onti nu es ext erna l yet thi s i nt r overt ed p a r t h a s not

,
b ec om e i nt ern a l i n th e s t rict phy s i ol ogi ca l s en s e F or it .
r em a i ns s u bj ect t o th e a cti on s of th os e en vi r oni n g m a tt er s
whi ch a r e t a k en i n a s food : s u ch en vi r on i n g m a tt ers wh en ,
th ey h a pp en t o be m ovi ng p rey a cti ng u p on it mu ch a s th ey

,
3 02 P H Y SIOL OGICA L D E VE L OP ME NT .
might a ct up on th e ext eri or S o th a t thi s i nt rov ert ed p a r t

.
h a s a qu a s i ext erna lity

-
I t h a s n ot the s a m e a b sol u t e i n
.
t erna l i ty a s h a v e th os e p a r t s whi ch nev er c om e i n cont a ct

wi th p rod u c ts of th e ou t er w orld H er e we m u s t b ri efly .
reco gn i z e th e di s ti nc ti on b et ween th e s e p a r t s a nd th e p a r t s
thu s fa r c ons id er ed .
Rev er ti ng t o ou r s ymb ol th e in di a ru bb er b a ll it will be

,
-
,
s een th a t th e i nt rov ers i on m a y be s o c ompl et e th a t the c a vi t y
i s oblit era t ed with th e resu lt th a t th e i n t erna l s u r f a c es o f

,
the ou t er a n d i nn er l a y ers c om e i n c on t a ct T hi s i s t he .
s t a t e rea c h ed i n t h e si m pl es t c oel en t era t e a ni m a l th e H ydr a , .
there b ei ng i n it n othi ng m ore th a n a thin s t ru ct u rel es s

l a m ell a b et ween the ect od er m a nd end od erm a s s h own i n ,
Fig 1 4 7 T hi s l a m ell a rep res ents a l l th a t th ere is of s t rictly

. .
i nt erna l ti ss ues Bu t th e i n t rov ers i on i ns t ea d of b ri ngi ng

.
,
the i nn er s u rf a c es of th e b a ll i n t o c ont a ct m a y be s o fa r i n ,
co mpl et e a s t o l ea v e a s p a c e a nd i n v a ri ou s c rea t u res a nd

,
em b ry os o f oth er s s ymb oli zed by thi s a rra ng em en t this s p a ce

, ,
b ec om es occ u pi ed by a ti s s u e f ormed f rom one or oth er or

b oth of the tw o p rim a ry ti s su es— th e m es obl a st or m es o
d er m T hi s i nt ermedi a t e l a y er s om etim es a s i n th e Medus a
.
, , ,
growi ng i nt o a m a s s of j elly servi ng a s a fu lc ru m for th e

c rea t u re s c ont ra cti ons or a s i n th e S p onge givi ng a p a s s iv e
’
, , ,
b a s i s t o the a ctiv e ti ss u es b ec om es i n high er anim a l s the

,
l a yer ou t of whi ch the st ru ct u res th a t s u pp ort the b ody and

m ov e it a b ou t a s well a s th os e th a t di s t ribu t e p rep a red
,
nu t rim en t a re d ev el op ed
, F rom it a ri s e the b ones the
.
,
m u s cl es a nd th e v a s cu l a r s yst em — th e m a s s es of di ff eren
,
ti s t e d ti s s u e whi ch a re t ru ly i nt erna l a nd occ u py wh a t is

c a ll ed the b ody c a vity or p er i visc er a l sp a c e
- -
.
I n the high er typ es o f a nim a l s this sp a c e c om es t o be

p a rti a lly occu pi ed by a s t ru ct u re th a t m a y be d esc ribed a s a
c a vity withi n a c a vity— the c oel om Mos t zool ogi st s rega rd .
t hi s a s a ri s i ng by a re i nt rov ers i on of the a r chen ter on or

-
p rim a ry a lim ent a ry sa c I t i s ea sily t o be p erceiv ed th a t

.
a ft er the int rov ers i on which p r od u c es thi s diges tiv e c a vity the ,
T HE OU TE R A ND I NNE R T ISS U E S OF A NI MA L S .
3 03
w a ll of vity m a y be a g a i n i n t r ov ert ed i n su ch w ay a s t o
th e ca
i nt ru d e i nt o the p eri vi s cera l s p a c e T he c oel om thu s f orm ed

-
.
i s s u b sequ ently s h u t off B ecom i ng i n c l u d e d a m ong t he m ore

.
t ru ly i nt erna l st ru ct u res a nd i n p a rt giving or igi n t o c ert a i n

,
li ni ng m emb ra nes it h a s for i ts chi ef fu nc t i on th e form a ti on

,
o f o rg a n s for the e xc reti on a nd e m i ss i on o f ni t r og enou s w a s t e
a nd o f t h e g enera tiv e p r o d u ct s : s om e p or ti on s o f it ret a i n i n g ,
a s a cons equ en ce i ndi rec t c onn exi ons w ith the envi ron m ent
,
a nd c h a ra c t er s u s u a lly a cc om p a n yi n g su c h connexi on s .
H ere w e a re n ot c on c erned w i t h fu rt h er d et a il s : th e a i m
bei ng s imply t o i ndi ca t e the w a y i n whi c h ou t o f the origi n a l
l a y er wh olly ext ern a l th ere a ri s e by p rim a ry a nd s ec ond a ry
, , ,
i nt r ov ers i ons a nd the f or m a ti on of i nt erm edi a t e m em b ra nes

,
a n d s p a c es the c hi e f c ont r a s t s b et w een ou t er a n d i n ner t i s

,
a n d h ow th ere s im u lt a neou s ly
s u es , g o on th e diff erenti a
ti ons a c c omp a n yi ng di fferent c onditi on s .
§ 28 9a . A n oth er al l -
imp ort a n t differenti a ti on b et ween
ou t er ti s s u es a n d i nner ti ss u es h a s n ow t o be set f orth— th a t
by whi ch the nerv ou s sy s t em b ecom es es t a bli s hed a nd di s

ti ngui sh ed S t r a ng ely enou gh lik e the one a b ov e d es crib ed
.
, ,
it i s s equ ent u p on a n i nt rov ers i on : th e n erv ou s s y s t em is

p ri m a r ily a s ki n s t ru c tu re a nd d ev el op s by the i nf oldi ng of
-
thi s s ki n—s t ru ct u re .
I n c rea t u res p os s ess i ng the ea rli es t ru dim ents o f nerv es

thes e exi s t i n c ert a i n su p erfici a l cell s E a c h h a s a s m a ll .
t u b u l a r orific e f rom w hich p r oj ect s a mi nu t e h a i r a nd ea ch ,
ha s on i ts u n d er s id e p r oc es ses r u nn i n g i nt o th e ti s s u e
be l ow a nd s ervi ng a s it s eem s t o c on d u c t imp res s i ons f ro m
, , ,
the p r oj ecti ng h a i r wh en it i s di s t u rb e d by cont a ct s with

for eign b odi es A pl exu s of fib r es b r i n gi ng th e i nner p ro
.
ces s es o f s u c h c ells i nt o c om m un ic a ti on a r i ses a n d f or ms ,
s om ethi ng lik e a n er v ou s l a y er ca p a bl e of p r op a g a ti n g i m
pu l ses i n a l l di recti on s A t a su b s equ ent s t a ge s om e of the

.
su pe r fici a l c ell s c ea s i ng t o be th ems elv es th e rec ipi ents of

,
ext ern a l s tim u li s i n k i nwa rds a nd bec om e g a ngli on c ells con

,
-
3 04 P H Y S I OL OGICA L D E V E L OP ME NT .
n e c ted wi t h the n erv ou s pl ex u s— a gent s a s we m u st s u ppo s e

, ,
for th e rec ep t i on m u l t iplic a ti on a nd di ffu s i on of t he i m p u l s es

, ,
r e c e iv e d fro m t h e o u t er c ell s .
A s th u s fa r d ev el op ed t he n erv ou s st ru ct u re i s one fitt ed

,
o n ly for a v a g u e stim u l a ti o n o f di s p ers ed cont ra c til e fib res ,
c a u s i n g m ov em ents o f a n u n di r ect ed ki nd A concent ra ti on .
o f th es e s u p erfi c i a l nerv ou s s t ru ct u re s is a p roba bl e p relim
i na ry t o th e nex t ch a n ge — a n a l l imp ort a n t c h a nge

-
F or a .
p a rt of th e s u rf a ce b egi n s t o si nk i nw a r d s f orm i ng i n the , ,
Ver tebr a ta a g roov e ; a nd f ro m the li ni ng cell s of thi s groov e

, ,
whi c h p res ently c l oses ov er th e ce nt ra l p a rts of th e nerv ou s

,
s y s t em a ri se : d efin it e nerv es h a vi n g m ea n tim e a s we m a y ,
s u pp os e b een d ev el op e d ou t of t h e i n d efi n it e n er v ou s pl exu s
,
.
N egl ec ti ng wh a t th ere i s i n thi s Of a sp ecu l a tiv e na t u re it ,
i s s u ffi ci ent for th e p re s en t p u rp os e t o recogni z e th e u n

d ou bt ed fa ct th a t the nerv ou s s y s t em i s devel op ed f rom the
ec t od erm a n d th a t or igi n a lly e xt ern a l it i s m a d e i n t ern a l by
, , ,
a p roc es s of si n ki n g in o r by a p r oc ess of d efini t e i nt rov ers i on .
§290 . Wh eth er di rect equ ilib ra ti on i n di re ct eq u ilib

orra
ti on h a s ha d the grea t er s h a r e i n p rod u ci ng th es e fu nd a menta l

cont ra s t s b e t ween th e i nner a n d ou t er ti s s u es o f a nim a l s ,
mu s t be l eft u nd ecid ed The tw o c a u ses h a v e al l a l on g c o

.
o p era t ed— m odi fi c a ti on of th e i n divid u a l a cc u mu l a t e d by

i nh erit a nc e p r ed omi na ti ng i n s om e c a s es a nd i n oth er c a s es ,
m odific a ti on of th e ra c e by s u rviv a l of the in c id ent a lly fitt est .
On the oth er h a n d th e a cti on of the m edi u m on th e orga nism

,
c a nn ot f a il t o ch a n g e its s u rf a c e m ore th a n its c ent re a nd s o ,
diff erenti a t e the tw o ; whil e on th e oth er h a nd the s u rf a ces

, ,
o f or g a ni s m s i nh a bi t i n g the s a m e m ediu m di s pl a y ext rem e
u n lik eness es which c a nn ot be du e t o th e imm edi a t e a cti on s
o f th ei r m edi u m L et u s dwell a m om en t on th e antith es i s

. .
We h a ve a bu nd a nt evid en c e th a t a nim a l p rot opl a sm is

ra pidly m odifi ed by light h ea t a i r wa t er a nd the s a lts
, , , ,
c ont a i ned i n wa t er— c oa gul a t ed t u rned f rom s ol u bl e i nt o i n

,
s ol u bl e p a rti a lly cha n ged int o is om eric c omp ou nds or other

, ,
“
TH E OU TE R A ND INNE R T ISS U E S oE A NI MAL S .

3 05
wise che mic a lly a lt ered I m m edi a t e met a m orph ose s of thi s
.
kin d a re o ft en Obvi ou s ly p r odu ced i n ova by ch a nge s of th ei r

m edi a . A t the o u t set th ere f ore b efore yet th ere e xi s t ed
, ,
a ny s u c h di ffer enti a ti on a s th a t whi c h n ow u su a lly a r i s es by
i nh erit a n ce th ese envi r oni ng a g enci es m u s t h a v e t end ed t o

,
o rigin a t e a p r ot ec t iv e en v el op e F or a m o difi c a ti on p r od u ced

.
by th em on the su p er fi ci a l p a rt of the p ro t opl a s m m u s t ,
eith er h a v e b een a d ecom p o s iti on or e ls e t h e f orm a ti on o f a
c o m p ou n d w hi c h r em a i n e d s t a bl e u n d er th e i r s u b sequ en t
a cti on. T h ere w ou ld be g enera t e d a n ou t er l a y er of s u b s t a nce

tha t wa s s o m ol ecu l a rly imm obil e a s t o be in ca p a bl e of fu rth er
m et a m orph os es whil e it w ou ld s hi e ld the c ont a i ned p r ot o
,
pl a s m f rom th a t t oo g rea t acti on of ext ern a l forces w hi ch by

-
,
ra pidly c h a ngi n g th e un s t a bl e equ ilib r i u m o f i t s m o l ecu l es
i nt o a rel a tiv ely s t a bl e eq u ilib r i u m w ou ld a rres t d ev el op m ent

,
.
E vid ently o rga ni c ev ol u ti on wh eth er i n divid u a l or g enera l

, ,
mu s t a lwa y s a nd everywh ere h a v e b een s u b or di na t e t o th ese

physic a l n eces s iti es . T h ou gh n a t u ra l s el ec ti on b egi nni n g ,
w ith mi nu t e p ort i on s o f p r ot opl a s m m u s t a l l a l on g h a v e ,
t end ed t o es t a bli s h a m ol ecu l a r c om p os iti on a pt t o u nd erg o

thi s di fferenti a ti on of su r f a ce from cen t re t o th e m os t f a v ou r
a bl e e xt en t, yet i t m u s t a l l a l on g h a v e d one s o whi l e con
t roll ed by thi s p roc es s of di rect eq u ilib r a ti on .
C on t r a riwi se th e m a n y a n d g rea t u n lik enes s es a m ong th e

,
d erm a l s t ru ct u res of c rea t u res i nh a biti n g th e s a m e el em ent ,
c a nno t be a s c rib ed t o a ny s u ch c a u s e T he c ont r a s t s be t ween .
na k e d a nd s h ell e d Ga s t r op od s b et ween m a r i ne Wor ms a n d

,
Cru s t a cea ns b et w een s o ft —s ki nne d Fi s h es a nd Fi s h es i n

,
a rm ou r lik e th e P ter i ch th s m u s t h a v e bee n p rod u ce d en ti r el v

y ,
by na t u ra l s el ecti on E n vi roni ng f or ces a re a s b ef ore th e

.
, ,
u l t i m a t e c a u s es ; bu t th e force s a re n ow n ot s o m u ch t h os e
e x erc i s e d by t h e m e di u m a s th os e ex er c i s e d by th e oth er i n
h a bit ant s of th e m ediu m ; a nd th ey d o not a ct by m odifyi n g

th e su r fa ce o f th e i n divid u a l bu t by killi n g off i n divid u a ls
,
wh os e su r fa ces a re l e a s t fitt ed t o th e requ i rem en ts : thu s

s l owly a ffe cti ng th e s p ec i es S till the d erm a l s k el et on b ri s tli ng
.
66
3 06 P H YS I OL OGICAL D E VE L OP ME NT .
with s pi nes whi ch p rot ects the D i odon or the Cycl i ch thgs
,
f ro m enem i es it c ou ld not es ca p e c om es withi n the g enera l

,
f ormu l a of a n ou t er ti s s u e di fferenti a ted fro m i nner ti s su es

by the ou ter a cti ons t o whi ch the c rea t u re i s exp os ed : th e
differenti a ti on h a vi ng g one on u ntil th er e i s eq ui lib riu m
b et ween the d es t ru ctiv e forces t o be m et a nd the p rot ectiv e
forces whi c h m eet th em .
I f we v ent u re t o a pp orti on th e res p ectiv e s h a res whi ch

m edi a t e a nd imm edi a t e a c t i on s h a v e h a d i n diff erenti a ti n g
o u t er f ro m i nn er ti s s u es w e s h a ll p rob a bly n o t be fa r w rong

,
i n a scrib i n g th a t p a rt of th e res u lt w hi ch i s a lik e i n a l l

a nim a l s ,
m a i nly t o th e di rec t a c ti ons o f th ei r m edi a w hil e ,
we a s c r ibe the m u ltit u di nou s u nlik enes s es of th e resu lts i n

v ari ou s a nim a l s p a rtly t o the i ndi rect a cti ons o f the m edi a
, ,
a n d p a rtly t o t h e i ndi r ec t a c ti o ns o f o th er a n i m a l s by w hich
the m edi a a re i nh a bit ed T h a t i s t o s a y whil e a s sign i ng th e

.
,
s p eci a liti es o f t h e di f fer enti a ti on s t o th e s p eci a li ti es of c on

v ers e w i th the a genc i es i n th e envi r onm ent m os t of th em ,
o rg a n ic ,
we m a y a s s ign t o t he c on s t a nt a nd u n iv ersa l c on
v ers e w ith its i n org a n ic a genci es the u niv ersa l ch a ra ct eri s ti c
,
o f t e gu m en t a ry s t ru c t u r es— th e i r g r ow th ou t w a r d s fr om a
l a y er lyi ng b el ow the su rf a c e whi ch c onti nu a lly p rod u c es new

s u b s t a n c e t o repl a c e th e s u b s t a nc e w o rn a wa y or ca s t of f .
H ere l et m e a dd a pi ec e of evid ence whi ch s t rength ens

the genera l a rgu m ent a t th e s a m e tim e th a t it j u s tifi es thi s
,
a pp orti on m en t . Wh en u lc era ti on h a s g one d eep en ou gh t o

d es t roy th e t egu m ent a ry s t ru ct u res th ese a re nev er rep ro
,
du ced . T h e p u c k ered s u r f a ce f orm ed wh ere a n u lc er h ea l s ,
or w h ere a s er i ou s b u rn h a s d es t r oy ed t h e s ki n cons i s t s of ,
m odifi ed c onnectiv e ti s su e which a s th e h ea li ng g oes on

, , ,
s p r ea d s i nw a r d s fro m th e edges o f th e u lc er : s om e o f it ,
p erh a p s g row i ng f ro m the p orti ons of c onnectiv e ti s s u e th a t

,
dip d own b et ween th e m u s cu l a r bu ndl es T hi s c onnec tiv e

.
tis su e i s nor m a lly c ov ered by the epid ermis a n d th u s s h elt ered

f rom envi roni ng a cti on s Wh a t ha s h a pp ened to i t ? I t h as
.
n ow bec om e the ou t er m os t l a yer A n d h ow d oes it c omp ort

'
.
TH E OU TE R A ND INNE R TISS UE S OF A NIMAL S .
3 07
its elf u nd er its new c onditi ons ? I t p rod uc es a s u p erfici a l

s u b s t a n ce w hi c h pl a y s t h e p a r t o f th e epid erm i s a nd g r ow s
ou t wa rdly F or s i nc e th e s u rfa c e s u bj ec t t o f ri c ti on a nd
.
,
exf oli a ti on h a s t o be c on ti n ua lly renew ed

,
th ere m u s t be a
,
c ontinu a l rep rod u cti on of a n ou t er m os t l a yer f rom a l a yer

benea th T h a t is t o s a y t he cont a ct of thi s d eep sea t e d ti ss u e
.
,
-
w ith ou t er a g enci es p ro d u c es i n it s om e a pp r oa ch t owa rd s

,
t h a t ch a ra ct er which w e fi nd u ni v er s a lly ch a ra ct er i zes ou t er

ti s su e . B u t whil e we s ee u n d er thi s exp os u re t o th e c on
di ti ons c omm on t o a l l i nt egu m ent a t en den cy t o a s su m e
,
t he s t ru ct u re c om m on to a l l i nt e gu m ent w e s ee n o t end ency

,
t o a s su m e a ny of th e s p eci a liti es o f t egu m ent a ry s t ru ct u re :

n o ru di m en t s of gl a n d s or h a i r s a c s m a k e th ei r a pp ea ra n c e .
A na l og ou s c onc l u s i on s m a y be d ra wn res p ecti ng the pro

c es s es of di fferenti a ti on by whi c h f rom the ou t er l a y er
nerv ou s ti s s u e a nd fi na lly a ner v ou s sy s t e m a re e v olv ed .
H ere a l so b oth di rec t a n d i n di rec t equ ilib r a ti on a pp ea r t o

, ,
.
h a v e op era t ed T w o rea s ons m a y be a ssigned for th e b eli e f

.
th a t the t ran s form a ti on of c er t a i n s u p erfi c i a l cells i nt o s ens i

tiv e cell s wa s i niti a t ed by exp os u re t o ext erna l stim u li T he .
fi r s t i s th a t ext rem ely u nst a bl e a s p r ot opl a s m i s di s t u rb

, ,
a n ec s receiv e d by th e ou t er s id e o f a s p ec i a lly exp os ed c ell -
c ou ld s ca rcely fa il t o c a u s e ch a ng es p a ss i ng th rou gh it
t ow a rd s the i nt eri or m a ss of the b ody a nd th a t p erp et u a l
,
r ep etiti on o f s u ch ch a ng es w ou ld t en d t o g ener a t e ch a nn els
o f ea s y t r a n s mi s si on th rou gh th e p r ot opl a s m T he s ec on d
.
rea s on i s t h a t i f w e do n ot a s s u m e thi s p r oces s of i n iti a ti on

,
bu t a s su m e th a t s u rviv a l of th e fitt es t w a s the s ol e a g ency ,
th en n o rea s on ca n be a ssigned why th e ner v ou s s y s t em

s h ou ld n o t h a v e b ee n a t th e ou t s et f or m e d i n t e r na lly i ns t ea d
of b ei n g i n iti a t e d ext er n a lly a n d th en t r a ns ferr ed t o th e i n
teri or : the r ou n d a b ou t p r oc es s w ou ld be i nexpli ca bl e At .
the s a m e ti m e th e p rod u cti on of a c ent ra l nerv ou s sy s t e m by

i nt r ov er s i on of su p erfi ci a l s ens itiv e cells ca nnot be a s crib ed
to the differen ti a ti n g effects of ext ern a l s ti m u li bu t m u st be
,
a s crib ed t o n a t u r a l s el ecti on N O p er pe t u a l rep etiti on of

.
P H Y S IOL OGICA L D E VE L OP ME NT
.
t
o u er di s t u rb a nces wou ld c a u s e the s i nki ng i nwa rd s a nd ,
c ov eri n g u p o f th e s p eci a lly s en s i t iv e a re a a nd th e pl exu s

,
-
b el ow i t B u t it i s m a ni fes t th a t s i nce these nerv ou s s t ru e

.
t u res a t once a l l i m p ort a nt a nd ea s ily i nj u red wou ld be

,
-
,
s a fer if r em o v ed f ro m t h e s u r fa ce s u r viv a l of t he fitt es t c on

, ,
t i nu a l l y p reservi n g th os e i n whi ch th ey w ere m ore d eeply

s ea t e d w ou ld t en d t o p r o d u c e a n a rra ng em en t i n w hi ch a l l
,
p a rt s bu t the a ct u a l rec eivers of ext ern a l s t im u li b eca me

i nt erna l .
H enc e c ont empl a ti ng g enera lly th es e two f un d a m ent a l

,
diff erenti a ti ons of i nner f rom ou t er ti s s u es we m ay concl u d e

,
th a t th ou gh thei r fi rs t st a ges res u lt ed from di rect eq u ilib ra

ti on th ei r s u b s equ ent a n d high er st a ges res u lt ed from i n
,
di rect equ ilib ra ti on .

C H A P T E R VI I .
I
D FFE RE N T A T I IO N S AMO N G T H E O UT E R I
T SS UE S OF
AN IM A L S .
§ 29 1 . TH E t
ti s s u es Of an im a l s origi na lly h om o
ou er ,
g e n eo u s o v er th ei r w h ol e s u r fa ce s p a s s i n t o a h et er o g en eity
,
whi c h fi t s thei r res p e ctiv e p a rt s t o th ei r r es p ec tiv e c on diti on s .
S o nu m erou s a n d v a ri e d a re t he i m pli ed di fferenti a ti ons th a t ,
i t i s i m p ra cti ca bl e h er e t o d ea l with th em a l l e v en i n ou tli ne .
T o t r a c e th em u p t h rou gh c l a sses of a n i m a l s o f i ncr ea s i ng

d egrees of a ggr eg a t i on wou ld ca rr y u s i n t o u ndu e d et a il
, .
D i d s p a ce p er m it it w ou ld be p os s ibl e to p oi n t ou t a m on g
,
th e P r o tozoa v a r i ou s ca s es a n a l ogou s t o th a t of th e A r cel l a

,
whi ch m a y be d esc r ib ed a s lik e a m i c ros copi c L imp et h a vi ng ,
a s a r c od e b ody o f w hich th e u pp er s u r fa c e h a s b ecom e h orn y ,
whi le th e l ow er s u r fa c e with i ts p rot ru di ng p seu d op odi a ,
r et a i n s th e p r imitiv e j elly lik e c h a r a c t er

-
T h a t differenti a
.
ti ons of thi s kin d h a v e b een gr a d u a lly est a bli s h ed a m on g

th ese mi nu t e c rea t u res th rou gh th e u nlik e rel a ti on s o f t h ei r
p a rt s to th e en vi ron m en t i s a n i nferen ce s u pp or t ed by a
,
for m whi c h w hil e th e r es t o f t he body h a s a sc a rc ely di s

,
“
a g r ees with A r c e l l a a n d D i l u ia i n
fi g
‘
t i ng ui sh a bl e c oa ti ng ,
h a vi n g the p s eu d op odi a p rot ru s ibl e f rom one ext remity only

”
of the bodv .
Ma n y p a r a ll el s p ec i a liz a ti on s of s u r f a ce a m on g a gg reg a t es
of t he s ec on d ight be i n s t a n ced fr om th e C ael en tera ta
or d er m .
I n th e H ydr a th e ect od er m p res ents o v er i t s w h ol e a rea n o

,
c on s pi cu ou s u n lik enes s es bu t th ere u s u a lly e xi s t i n th e

hyd roid p olyp es of su p eri or typ es decid ed c ont ra s t s be t w een
,
309
310 P H YS IOL OGICA L D E VE L OP ME NT .
the high er a nd l ower p a rts Whil e th e high er p a rts ret a i n

.
th ei r origin a l ch a ra ct ers th e l ower p a rt s exc ret e h a rd ou t er

,
l a yer s yi eldin g s u pp ort a nd p r ot ecti on Va r i ou s s t a g es o f .
”
I n H ydra c ti n ia “
the different i a ti on might be f oll ow ed .
,
s a y s P ro f Green thi s h orn y l a y er

.
,
b ec om es el ev a ted a t i n
terva l s t o f or m nu m er ou s r ou gh p roces ses or spi n es whil e ,
o v er th e g en era l s u r f a c e of th e ec t o d er m i t s p res en ce is
”
a lm os t imp erc eptibl e I n o th er typ es a s i n C or dg l op hora
.
, ,
’
it sp rea ds p a rt wa y u p the a ni m a l s s id es endin g i ndefi nit ely ,
.
I n E i m eria it e xt en d s it s elf s o a s t o en cl ose the enti re b ody
of ea ch p olypit e l ea vi ng b a re on ly the m ou th a nd tip s of the

,
t ent a cl es .
”
Whil e i n Ca m p a nu l a ri a it h a s b ec om e a p a rti a lly
d et a ch ed ou t er cell i nt o whi ch the c rea t u re ca n ret ra ct its
,
e xp os ed p a rt s .
But it i s a s needl es s a s it w ou ld be w ea ri s om e t o t ra c e
th rou gh the s ev era l su b ki ngd om s th e ri s e of th es e mu ltif orm
-
c on t r a s t s with th e vi ew of s ee ki n g i nt erp ret a ti on s o f th em

, .
I t will s u ffi ce if we t a k e a few g rou ps of the ill u st ra ti ons

f urni sh ed by th e high er a nim a l s .
292 . We b egin with th os e m odifi ca ti ons of su r fa c e

m ay
whi ch s u bs erv e res pi ra ti on T h ou gh we ordi na rily thin k of

.
res pi ra ti on a s the q u it e s p ec i a l fun cti on of a q u it e s p ec i a l
or g a n yet or igi n a lly it is not s o

, L ittl e d ev el op ed a ni m a l s
.
-
p a rt with th ei r c a r b onic a cid a n d a b s orb oxygen th r ou gh the ,
g enera l s u rf a c e of the b ody E v en i n the l ower typ es of the

.
high er cl a s s es the genera l s u r f a c e of the b ody a id s l a rgely i n

,
a er a ti n g th e bl ood ; a n d th e p a rt s whi c h di sc h a rg e the g rea t er
p a rt of thi s f un cti on a re s u b s t a nti a lly nothi ng m ore th a n

s lightly a lt ered a nd e xt en d ed p orti on s of th e s ki n .
S u ch differenti a ti ons m ark ed i n v a ri ou s d egrees a re t o be

, ,
s een a m on g M ol l us ca I n th e P ter op oda th e only m odific a ti on

.
which a pp ea rs t o f a cilit a t e respi ra ti on i s the mi nu t e vas cu ,
l a ri ty of one p a rt o f th e ski n H igh er typ es p os s ess s peci a l

.
ski n d ev el opm ents

-
The D ori s h a s a pp end a g es d ev el op ed
.
i nt o el a b ora t ely b ra nch ed f orms —s m a ll t rees of bl ood v ess els

- -
T HE OU T E R T IS S U E S OF A NI MA LS .
31 1
c ov ered by s lightly ch a nged d er m a l ti ss u es

-
A nd th es e
.
a rb ores cen t b r a nchi ae a re g a th ere d t og eth er i n t o a si ngl e

cl u st er T hu s th ere i s evid enc e t h a t l a rg e ext erna l res pi ra
.
t ory orga ns h a v e a r i sen by d egree s f ro m simpl e s ki n : a s ,
i nd ee d th ey do a r i s e d u ri n g th e d ev el opm ent of ea c h i ndi

,
vi du a l h a vi ng t hem Ju s t a s g ra d u a lly a s i n t he em b ry o
.
a s i m pl e bu d on t h e i nt egu m en t w ith i t s c ont a i ned v a s c u l a r

,
l oop p a s s es by s ec ond a ry bu ddi ng s i nt o a t ree lik e g rowt h

,
-
p enet ra t ed everyw h ere by divid i ng a n d s u b dividi ng bl ood -
v es sel s ; s o gra d u a lly h a s th ere p r ob a bly p roc eed ed the

differenti a ti on whi c h h a s t u rn ed p a r t o f the ou t er su r f a ce
i nt o a n org a n for ex cretin g c a rb oni c a cid a nd a b sorbi ng
oxyg en .
Cert a i n i n feri or v ert eb r a t e a nim a l s p resen t u s wi th a li k e

m et a m orph os i s of ti s s u es T h ese a re t he A m p hi bi a
. The .
b ra nchi ae h ere d evel op e d f rom the s ki n a re cov ered w ith cel

,
l u l a r epid erm i s n ot m u c h thi nner th a n th a t c ov eri ng th e res t

,
of the b ody L ik e it th ey h a v e th ei r s u rfa c es s p eckl ed w ith

.
pigm en t c ells ; a nd a re n ot even c ons picu ou s by th ei r ext ra

-
v a s cu l a rity— wh ere th ey a re t emp or a ry a t l ea st T h ey f a cili .
t a t e th e ex ch a nge of g a ses i n s ca r cely a ny other wa y th a n by

a ffor di n g a l a r g er a rea o f c ont a c t with th e wa t er a nd i n t er ,
p os i ng a ra th er thi nner l a yer of ti s s u e b et ween th e wa t er

a nd th e bl oo d v es s el s -
T h os e v ery s i m pl e br a nchi ee of th e
.
l a rv a l A m p hi bi a th a t h a v e th em but for a s h ort tim e ,
gr a du a t e i nt o the m ore com pl ex ones of th ose th a t h a v e them

for a l on g tim e or p erm an ently ; s h owi ng a s b efore the s m a ll
, ,
s t a g es by whi c h thi s h et erog eneity o f s u r f a c e a cc omp a nyi n g
h eter og eneity of fu ncti on m a y a ri se .
I n wh a t wa y a r e s u ch differen ti a ti on s est a bli s h ed ? Ma i n

l y n o d ou bt by na t u ra l s el ecti on bu t a l s o t o s om e d egree I
, , ,
t hi nk by th e i n h erit a nc e of di rect a d a pt a ti on s
,
T h a t a p or
.
ti on of th e i nt egu m ent a t which a era ti on i s fa v ou red by l oc a l

c on diti ons s h ou ld th ereby be l e d t o g r ow i n t o a
,
l a rger
s u rf a ce o f a era ti on a pp ear s imp rob a bl e
,
S u rviv a l of th ose
.
i ndividu a ls which h a pp en t o h a ve thi s p orti on of the i ntegu

312 P H YS I OL OGI CA L D E VE L OP ME NT .
m ent s om ewh a t m ore d ev el op ed , s ee m s h ere t he on ly lik ely

c a u se .
29 3 . A m ong the c ons picu ou s m odific a ti ons by whi ch the

origi na lly u ni for m o u t er l a y er i s r en d er ed m u l t i form
-
a re th e ,
p rot ectiv e s t ru ctu res L et u s l o ok fi rs t a t the few ca ses i n

.
which t he f orm a ti on of th es e i s a s c r ib a bl e m a i nly t o di rect

equ ilib ra ti on .
A l rea dy referenc e h a s b een m ore th a n once m a d e t o th ose

thi ck eni ngs th a t occu r wh ere th e s ki n i s exp osed t o u nu su a l
p ressu re a nd fri cti on A r e th es e a d a pt a t i ons i nh eri ta bl e ?
.
a n d m a y th ey by a c cu m u l a ti on th rou gh m a ny g en era ti on s
, ,
p r od u ce p er m a nent d er m a l s t ru ct u res fitt ed t o p erm a nent or

freq u ently recu r ri ng s t ress ? T a k e for i n s t a nce th e ca ll os i
-
, ,
ti es on the k nu ckl es of t he Gori l l a w hi ch a re a d a pt ed to ,
i ts h a bit o f p a rti a lly su pp ort i ng it s elf on i ts cl osed h a n d s

when m ovi ng a l ong th e g r ou n d S h a ll we s u pp os e th a t th es e
.
d efen s ive thi c k eni ng s a re p rod u ce d a fres h i n ea ch i ndividu a l

by the di rect a cti ons ; or th a t th ey a re i nh erit ed m odi fi ca
ti on s c a u s ed by su ch di rect a cti ons ; or th a t th ey a re wh olly
du e t o t h e n a t u ra l sel ecti on of s p on t a neou s v a ri a ti ons ? T he
.
l a s t su pp os iti on d oe s n ot s eem a p rob a bl e one S u ch thi ck en .
i ng s i f s p ont a neou s w ou ld be no m ore lik ely t o occu r on the

, ,
knu ckl es th a n on a ny oth er of th e hu nd red equ a l a rea s form

i ng th e s ki n s u rf a c e a t l a r ge ; a nd th e c h a n c es a g a i n s t th ei r
-
s imu lt a ne ou s o cc u rren ce on a l l eight k nu ckl es w ou ld be i n
c a l cu l a bl e Moreov er the impli ca ti on wou ld be th a t th os e

.
,
s light e xt ra thick nes se s o f s ki n on the k nu c kl es w ith which ,
we m u s t s u pp os e th e sel ecti on t o h a v e c om m enc ed w ere s o ,
a dv a nt a geou s a s t o c a u s e s u rviv a l s Of the in divid u a l s h a vi ng
th em i n p res enc e of oth er s u p eri oriti es p os s es sed by oth er

,
i ndivid u a l s T h en th a t s u rviv a l s s o c a u s ed if th ey ev er
. ,
o cc u rr ed a t a l l sh ou ld h a ve occ u rred w ith the f req u ency

,
r eq u i s i t e t o es t a b l i s h a nd i nc r ea s e t h e v a r i a ti on i s h a rdly ,
s u pp os a bl e A nd if w e rej ect a s a l s o u nlik ely th e rep ro

.
, ,
d ucti on of th es e ca ll os iti es al e n ovo i n ea ch i ndivid ua l ( for

T HE OU T E R TIS SU E S or A NIMA L S .
313
thi s wou ld imply th a t a ft er a th ou s a nd g enera ti on s ea ch

y oun g g orill a bega n with kn u ckl es h a vi ng s ki n no thi ck er
th a n el s ewh ere ) there rem a i ns only th e i n ference th a t t h ey
,
ha v e a r i s en by the t ra ns mis s i on a nd a ccu m u l a ti on of fu nc

t i ona l a d a pt a ti ons . A noth er c a se whi c h s ee m s
i nt erp ret a bl e only i n a n a na l og ou s wa y i s th a t of the s pu rs,
th a t a re d evel oped on the wi n g s o f c er t a i n bi rd s— on th os e

o f t h e C h a j a s c rea m er for ex a mpl e T h ese a re w ea p ons of
.
o ffen c e a nd d e fence I t i s a f a m ili a r f a ct th a t s om e bi rd s

.
s t rik e with th ei r w i n g s o ft en givi ng s ev ere bl ow s ; a nd i n th e

,
bi r d s na m ed the bl ow s a re m a d e m ore formid a bl e by the

,
h orn y d a gger S h a pe d growt h s s ta n ding ou t f rom th ose p oi nt s

,
-
on t he wi ng s whi c h d eliv er th em A r e th es e s p u rs di rec tly

.
o r i ndi rectly a d a ptiv e ? T o c onc l u d e th a t na t u r a l s el ec t i on

o f s p on ta neou s v a ri a t i ons h a s c a u se d th e m i s t o concl u d e
,
tha t with ou t a ny l oca l s timu l u s th ick en i ng s of the s kin

, ,
o cc u rred s ymm et ri ca lly on the two w i n g s a t th e pl a c es

req u i red ; th a t s u ch thick en i ng s s o l o c a li z ed
,
h a pp ened t o
,
a ri s e i n bi r d s
g iv en t o u s in g th ei r wi n g s i n fight ; a n d th a t
on thei r fi rst a pp ea ra n c e the thi ck eni n g s were d ec id ed
en ou gh t o g iv e a pp r eci a bl e a dv a n t a g es t o t h e i ndivid u a l s di s
t i ngu i sh ed by th em— a dv a n t a ges i n b ea r i ng th e r ea cti on s o f

th e bl ow s if not i n i n fli c ti n g th e bl o w s B u t t o c onc l u d e thi s
.
i s I thi n k t o c onc l u d e a ga i n s t p r ob a bility

, , C on t ra r i w i se
.
,
if w e a s su m e th a t th e thi c k eni ng Of th e epid er m i s p r odu ced

by h a bit u a l rou gh u s a ge i s i nh eri ta bl e th e d ev el op m en t of
,
th es e s t ru ct u res p res ent s n o di fficu lty T he p oi nt s of imp a ct

.
wou ld bec om e i n du r a t ed i n wi n g s u s ed fo r s t r iki n g w ith

u nu su a l f req u en cy The c a ll os iti es o f s u rfa c e th u s gener a t ed
.
,
r end er i n g th e p a rt s l es s sens itiv e w ou ld ena bl e th e bi rd i n

,
whi ch th ey a r os e t o giv e w i th ou t i n j u ry t o i t self m ore

, ,
vi ol ent bl ows a nd a g rea t er nu mb er o f t h em : s o i n s om e ,
c a ses h elpi n g it t o c onq u er a n d m u ltiply

, A m on g i t s d es cen d
.
a nt s i nh eriti n g th e m o difi ca t i on a n d th e a ccom p a n yi ng h a bit

, ,
the thi ck eni n g w ou ld be fu rt h er i n c rea sed i n th e s a m e w a

y
s u rviv a l o f th e fitt e s t t endi n g e v er t o a c c el era t e th e p r oc e s
s .
P resently th e h orny
d f orm ed hith ert o d ef ensiv e only
n o es s o ,
i n th ei r effect s w ou ld by th ei r p r om i n enc e b ecom e offen s iv e

, , ,
— wou ld m a k e t h e bl ows giv en m ore h u rt fu l A n d n ow .
n a t u ra l s el ec ti on a idi n g m ore a c tiv ely

,
w ou ld m ou ld th e ,
n od es i nt o s p u r s : t h e i n divid u a l s i n whi ch t he no d es were
m os t p oi n t ed wou ld be a pt to s u rviv e a n d p r op a g a t e ; a n d the

p oi nt ed nes s g enera ti on a ft er g enera ti on thu s increa s ed w ou ld ,
en d i n th e well a d a pt ed s h a p e we s ee
-
.
B u t if i n th es e c a ses th e di fi er ent i a ti ons w hi ch fit p a r

t i cu l a r p a r t s o f the ou t er ti ss u es t o b ea r rou gh u sa g e a re
c a u s ed m a i nly by t he di r ec t b a l a n c i n g o f e x ter na l a ct i on s by
i nt erna l rea cti on s then we m a y s u s p ect th a t the like i s t ru e

,
o f o th er m odific a ti on s th a t oc cu r w h er e S p ec i a l s t r a i n s a nd
a b ra s i on s h a v e t o be m et P os s ibly it i s t ru e of s u n d ry p ar t s
.
th a t a re f orm ed of h a r d ened epid er mi s s u ch a s the na il s , ,
cl aws h oof s a nd h oll ow h orn s of M a m m a l s ; a l l of w hi ch

, ,
”
,
s a ys P r o f H u xl ey . a r e con s t ru ct ed on essen ti a lly t h e s a m e

,
pl a n b ei n g div erti cu l a of the wh ol e i nt eg um en t the ou t er

, ,
”
l a y er of w h os e ecd eron h a s u n d er g one h orny m et a m or ph os i s .
L ea vi n g op en h ow ev er th e qu es ti on w h a t t eg u m ent a ry
, ,
s t r u c t u res a r e d u e t o di rect equ ilib r a t i on fu r th ered a n d c on ,
t roll ed by i n di rect equ ilib ra ti on it i s t ol era bly cl ea r th a t,
di rec t equ ilib ra ti on ha s b een one of the f a ct ors .
§ 29 4 . ct
D er m u res a o f
l a n o th
s t ru
er c l a s s a r e d e v el o p e d
m a i nly i f not w h oll y by th e a cti ons of ext ern a l c au ses
, ,
on s p eci es ra th er th a n on i ndivid ua l s T h es e a re the .
v a ri ou s ki n d s o f c l othi ng— h a i rs f ea th ers qu ill s sc a l es , , , ,
s cu t es T h ou gh it i s no l ong er th ou ght a s a t one ti m e th a t

.
a l l th es e v a r i ou s t egu m en t a ry s t ru c t u res a re h om ol og ou s wi th
one a n oth er yet it i s u nq u es ti on a bl e t h a t s u nd ry o f th e m or e

,
c on s pic u ou s ones a re T h o s e whi ch a re e xt rem ely u nlik e

.
m a y be seen li nk ed t og eth er by a l on g s eri es o f g ra d u a t ed
f or m s A ret rogra d e m et a m or ph os i s f rom fea th ers t o a p

.
p en d a ges th a t a re a lm os t sc a l e lik e i s well s een i n the c oa t

-
,
o f t h e P engu i n T h ere i s m a nifes t a t ra ns iti on f r om the

.
T HE OU TE R TISS U E S OF A NI MA L S .
31 5
bi rd —lik e c ov eri ng t o the fi sh l ik e c overi ng— a t ra n s i ti on s o

-
g ra d u a l th a t n o pl a ce c a n be f ou nd wh ere a n a pp reci a bl e
b rea k occu r s ; a nd if the s ca l e lik e a pp end a ges a re n ot -
t ru ly s c a l es yet th ey ex em plify a n ext rem e m et a m orph os i s .
L e s s st ri ki n g pe rh a ps bu t s c a rc e ly l es s s ign ifi ca nt a re
, , ,
the m odifi ca ti ons t hrou gh w hi c h we p a s s from fea th ers to

h a i rs on the s u rf a c es of the Os tri ch a n d the C a s s owa ry
,
.
T he s ki n of t he P orc u pi ne S h ow s u s h a i r s a nd qu ill s un it ed
by a s eri es of i nt erm edi a t e st ru ct u res di fferi n g f rom one ,
a not h er i n a pp r ec i a bly E v en m ore rem a r k a bl e a re cer t a i n

.
o th er a lli a n c es of d er m a l s t ru c t u res I t m a y be t a k en a s
“
”
cert a i n I thin k s a y s P r of H u xl ey th a t the s ca l es pl a t es
, , .
, , ,
a nd s pi n es o f a l l fi s h es a r e h om ol o g ou s org a ns ; n or a s l es s s o
tha t the t egu m ent a ry s pi n es Of the P l a gi os t om es a re h om o

l og ou s with th ei r t eeth a nd th ence w ith th e t eeth of a l l
,
vert eb ra t a .
Fu rt her d et a il s c oncerni n g th es e t eg u m ent a ry st ru ctu res

a r e n ot nee dfu l for p r es en t p u r p os es a n d a re i n d eed bu t i n,
di rec tly rel ev a nt t o the su bj ec t of phy s i ol ogic a l d evel opm en t .
H ere th ey a re of i nt eres t t o u s only by inv olvi n g th e g enera l

qu es ti on— Wh a t ph y s i ca l i n fl u ences h a v e b r ou ght th em i nt o
exi s t en c e ? S till w i th a vi ew t o d efi nit e p r es en t a ti on of th e
p robl em it will be well to c ont em pl a t e the m od e o f de
,
vel op m ent c o mm on t o th e m os t f a mi li a r o f th e m .
S u pp ose a s m a ll pit t o be f or m ed on the p revi ou s ly fl a t

S ki n ; a nd s u pp o s e th a t t h e g r o wth a n d c a s ti n g Off o f h or ny
c ell s w hi c h g oe s o n o v er th e s ki n i n g en er a l c on ti nu es t o ,
g o on a t th e u su a l ra t e ov er th e d ep ress ed s u rfa ce of
thi s pit Cl ea rly th e qu an tity of h orn y m a tt er p r od u ced
.
withi n thi s h oll ow will be g rea t er th a n th a t p r od u ced on a

,
l evel p or ti on o f the s kin s u bt endi n g a n equ a l a rea o f th e

S u pp ose su ch a pit t o be d eep en ed
’
a nim a l s ou tsid e .
u ntil it be c om es a s m a ll s a c I f th e ex foli a t i on g oes on a s

.
b ef ore the r esu lt will be th a t th e h orny m a tter exp ell ed a s

, , ,
it m u s t be th rou gh th e m ou th Of the s a c whi c h n ow b ea rs

, ,
a sm a ll p r op orti on t o t h e i nt er na l s u r f a c e o f th e s ac will be ,
31 6 P H YS IOL OGICA L D E VE L OP ME NT .
l a rg e i n qu a ntity c omp a red with th a t ex foli a t ed f rom a

p orti on of th e s ki n equ a l i n a rea t o the m ou th of th e s a c :
t h er e w ill be a c on spi cu ou s th ru s t i n g forth of h orny m a tt er .
S u pp os e onc e m ore th a t the s a c i ns t ea d of re m a i ni n g s i m pl e

, ,
h a s i ts b ott om p u s h ed u p i nt o i ts i n t eri or lik e th e b ott om Of ,
a w i ne b ot t l e — the i nt rov er si on b e i ng c a r ri ed s o fa r th a t t h e
-
i ntrov ert ed p a rt r ea ch es nea rly t o th e ext erna l Op en i n g a nd ,
l ea v es s ca rc ely a ny sp a ce be tween the i nt r ov ert ed p a rt a nd

the wa ll s of t he s a c I t i s ea s y t o s ee th a t th e ex foli a t i on
.
conti nu i ng f ro m th e s u rfa c e of th e i nt r ov ert e d p a r t a s w ell a s ,
f rom the i ns id e of the s a c genera lly the h orny m a tt er ca s t ,
Off wi ll f or m a d ou bl e l a y er ; a nd w ill c om e ou t o f t h e s a c
i n the s h a p e of a t u be h a vi n g w ithi n i ts l ower en d th e i nt ro

v erted p a r t a s the c ore on w hi ch it is m ou lded a nd f rom th e
, ,
a p ex o f whi c h i s c a s t off th e su b s t a n c e filli ng l ess d ens ely , ,
i ts i nt eri or T he st ru c t u r e resu lti ng w ill be w h a t we k now

.
a s a ha i r .Ma nif es tly by p rogres s iv e enl a rgem en t o f the s a c ,
a nd f u r th er c omplic a ti on o f th a t i nt r ov ert ed p a rt on wh i c h
th e exc ret ed s u b s t an c e is m ou ld ed th e p rot ru di n g g rowth m a y

,
be ren d ered l a r ger a n d m ore i nv olv ed a s w e s ee i t i n qu ill s ,
a n d fea th ers .S o th a t i ns en s ibl e s t ep s thu s i ndic a t ed i n

,
p ri ncipl e c a rry u s f rom the exf oli a ti on of epid er m is by a fl a t

,
s u r fa ce t o t h e e xf oli a ti on o f it by a h oll ow s i m pl e s a c a n
, ,
i nt rov ert ed s a c a n d a s a c f urther com pli ca t ed ; ea c h of whi ch

,
p rodu ces i ts m odifi ed ki n d of t egu m ent a ry a pp end a g e .
Bu t n ow a ft er c ont empl a ti ng thi s typic a l ill u s t ra ti on

, ,
we r et u rn t o th e g enera l qu es ti on Wh a t a re the a genci es

.
w hi c h h a v e b een Op er a tiv e i n d ev el opi ng th es e s ki n

s t ru c t u res ? I ndi rect eq u ilib ra ti on m u s t h a v e work ed a lm os t
a l on e i n p r o d u c i n g th em N o di rec t i ncid en ce o f f orces ca n
.
h a v e d ev el op ed the ena m ell ed a rm ou r of the L epi dos teus or

the t es s el a t ed pl a t es o f the Gl yp todon a n d i ts m od ern a lli es .
S u rviv a l of th e fitt es t m u s t h ere a nd i n mu ltit u di n ou s oth er

c a s es be reg a rd ed a s th e s ol e c a u s e .
§ 29 5 . A m on g m an y o th er di fferenti a ti ons o f t he ou er t
TH E OU T E R T IS S U E S OE A NI MA L S .
317
t i s s u es , t he m os t w orthy t o be noti c e d in th e
p a c e th a t re
s
m a i ns a re th os e by whi ch orga n s
,
of s e ns e a re f o r m e d We .
w ill b egin with the s impl es t a nd m os t c l os ely a lli ed t o the

f oreg oi ng .
E very h a i r th a t i s not t oo l ong or fl exibl e t o c onv ey t o its

r oo t e d end a s t ra i n p u t u p o n i t s f ree en d i s a ru di m en t a ry ,
t a ct u a l org a n ; a s m a y be rea dily p r ov ed by t ou chi n g on e o f

th os e gr ow i ng on the b a ck o f th e h a nd I f th en a crea t u re .
, ,
h a s c ert a i n h a i rs s o pl a c ed th a t th ey a re h a bit u a lly t ou ched

by the Obj ec t s with whi ch it d ea l s or a mid whi ch it m ov es , ,
a n a dv a nt a ge i s lik ely t o a cc ru e if th es e h a i r s a re m odifi e d
i n a wa y th a t en a bl es th em the be tt er t o t ra n smit the i m

p res s i ons d eriv ed S u ch m odifi ed h a i rs we h a v e in the
.
”
vi bri s s ae or a s th ey a re c o mm only c a ll ed t he whi s k ers
“
, , ,
p os s es sed by C a ts a nd feli ne a nim a l s genera lly a s well a s by ,
S ea l s a nd m a ny Rod en ts T h es e h a i rs a re l on g en ou gh t o
.
rea ch obj ect s a t c on s id era bl e di s t a n c es ; th ey a re s o stiff th a t
f orc es a ppli ed t o th ei r f ree end s c a u s e m ov em ents of th ei r

,
imb edd ed ends ; an d the s a cs c ont a ini ng th ei r imb edd ed ends

b ei ng well cov ered wi th ner v e fibres th es e d ev el op ed h a i rs
-
,
s erv e a s i n st ru m ents of expl ora ti on By c ons t a n t u s e of th em

.
the a nim a l l ea rn s t o j u dge of th e rel a tiv e p o siti on s of obj ects

p a s t which or t ow a rd s which it is m ovi ng Wh en st ea lthily
, , .
a pp roa chi n g p r ey or st ea lthily es c a pi ng enemi es s u ch a id s t o ,
perc epti on a re Obvi ou s ly imp ort a nt : in d eed th ei r imp ort a nce

h a s b een p rov e d by th e dimi ni s h ed p ow er of s elf gu id a nc e i n -
th e d a rk th a t res u lts f rom cu tti ng th e m off T h es e th en a re

,
.
, ,
d erm a l a pp en d a ges origi na lly s ervi ng the p u rp os e of cl oth

i ng bu t a ft erw a r d s di fi er enti a ted i nt o s ens e org a ns
,
-
.
T h a t ey es a re e ss en ti a lly d erm a l S t r u ct u res s eems sc arc ely

c onceiv a bl e Y et an ex a mi na ti on of th ei r ru dim ent a ry typ es
.
,
a n d of th ei r g enesis i n c rea t u res th a t h a v e th em w e ll d ev e
l op ed sh ow s u s th a t th ey rea lly a ri s e by s u cc es s ive m odi fi ca

,
ti ons of the d ou bl e l a yer com p os i ng the i nt egu m ent T h ey .
m a k e th ei r fi rs t a pp ea ra nc e a m ong the simpl er a nim a l s a s

S p ecks of pigm en t c ov ered by p orti on s of epid ermi s slightly
,
318 P H Y SI OL OGI CA L D E VE L OP ME NT .
con ev x and littl e more t ra ns p a rent th a n th a t a rou nd i t

a .
H ere th ei r fu n d a m ent a l com mu n ity of st ru ct u re with th e

s ki n is ea s y t o t ra ce ; a n d th e f orm a ti on of th em by di ff eren
ti a ti on of it p res en ts n o di ffi cu lty N ot s o fa r
.
i n a dv a nc e of th es e a s m u c h t o ob s cu re the rel a ti on ship a re ,
t he ey es whi c h th e C ru s t a cea ns p ossess

'
I n ev ery fi sh .
’
m onger s s h op we m a y s ee th a t th e eyes of a L ob s t er a re
ca r ri ed on p edicl es a n d w h en th e L ob s t er c a st s i t s sh ell t h e ,
ou t er c oa t of ea ch eye b ei ng c on ti nu ou s wi th th e epid ermi s

,
o f i t s p edicl e i s th rown Off a l on g w ith th e rest o f th e exo

,
s k el et on . Benea th the t ra ns p a rent epid erm ic l a y er th ere ,
exi s ts a g rou p o f ey es o f th e ki n d whi ch we see i n a n
i ns ect ; a nd th es e a c cor di ng t o a high a u th ority a re i ncl os ed

, ,
i n th e d erm a l sy s t em D es c r ibi ng th e a rra ngem en t o f th e

.
p a r t s M Mil ne E dw a r d s writ es
,
. Bu t th e m ost rem a rk a bl e
ci rcu m s t a nce i s th a t the l a rge c a vity withi n which the wh ol e
,
o f th es e p a r a ll el c ol u m n s ev ery one o f w hi c h i s its elf a p er

,
feet eye a re c ont a ined i s cl os ed p os t eri orly by a m emb rane

, , ,
w hi c h a pp ea r s t o be n eith er m ore n or l es s th a n th e middl e

t egu m ent a ry m em b r an e pi erce d for the p a s s a ge of the optic
,
nerv e ; s o th a t t he o cu l a r ch a mb er a t l a rg e res u lts f rom
th e s ep a ra ti on a t a p oi nt of th e tw o ext ern a l l a y er s of th e
genera l env el op e . T hu s t oo i s it i n th e m a i n , ,
ev en w ith th e highly d ev el op ed ey es of t h e Ver tebra te

-
.
T he th ree p a i r s of s en s or y org a n s a pp ert a i ni ng t o th e

“
high er s en s es s a y s P rof H u xl ey
,
th e n a s a l sa c s the ey es
.
, ,
a d t
n h e e a rs — a r i s e a s S impl e c oec a l i nv ol u ti on s o f th e ex
t ern a l i nt egu m ent of the h ea d of the emb ry o T h a t su ch .
i s th e c a s e s o fa r a s th e olfa ct ory s a cs a re c onc erned is

, ,
obvi ou s a nd it is n ot diffi cu lt t o obs erv e th a t th e l en s a n d

,
th e a nt eri or c h a m b er of th e eye a re p ro du c e d i n a p erf ectly

s imil a r m a nner I t is n ot s o ea s y t o s ee th a t th e l a by ri nth
.
of the ea r a ri se s i n thi s w a y a s th e sa c res u lti n g f ro m th e

,
i nv ol u ti on of the i nt egu m ent i s s m a ll and rem a i ns op en bu t ,
a v ery sh ort tim e ,

B u t I h a v e s o f req u ently v erifi ed
. .
’
H u s chk e s a nd Rem a k s s t a t em en t th a t it d oes s o a ris e th a t
’
,
T HE OU T E R TI SS U E S OF A NI MA L S .
I t t i
en er a n n o d ou bt wh a t ev er of th e f a ct T he ou t er en d s .
o f th e o lf a ct o ry s a c s rem a i n op en bu t th os e of t he ocu l a r
,
a u dit ory sa c s ra pidly c l ose u p

.
a nd s h u t of
,
f th ei r c on tent s
”
f rom a l l di rec t com m un i ca ti on w ith the ext er i or T h a t is .
t o s a y t he eye c on s id ered a s a n o pti c a l a pp a ra t u s i s p r o

,
du ced by m eta m orph oses o f th e s ki n : th e only p a rt s of it

n ot thu s p ro d u c ed b ei n g th e m em b r a nes lyi ng b et wee n t he
,
s cl e ro ti c a n d th e vit re ou s h u m ou r i ncl u di ng th os e reti na l

,
s t r u c t u r es for m e d i n th em A l l i s t egu m en t a ry s a v e th a t
.
whi c h h a s t o a pp reci a t e th e i m p res s i on s whi ch th e m odifi ed

i nt egu m ent c oncen t ra t es u p on it .
T h u s a s P r o f H u xl ey h a s s om ew h ere p oi nt ed ou t th ere
,
.
,
is a s u b s t a nti a l p a ra ll eli s m b et wee n a l l the s ens ory org a ns i n

thei r m od es Of d ev el opm ent ; a s th ere i s t oo b et ween th ei r , ,
m od es of a cti on A vi bri s s a m a y be t a k en a s th ei r c o m m on
.
typ e . I n crea sed i m p res s ibili ty by a n ext ern a l s timu lu s ,
req u i res a n i nc rea s ed p eriph era l e xp a n s i on o f th e nerv ou s
s y s t em on w hich t he s tim u l u s m a y fa ll ; a nd thi s i s s ec u re d
by a n i nt rov ers i on of the i nt egu m en t f ormi ng a s a c on the ,
wall s of whi c h a n erv e m a y ra m ify T h a t th e m ore ext end ed

.
sen s ory a r ea t h u s con s tit u t ed m a y be a c t e d u p on th ere ,
r eq u i res s om e a pp a ra t u s c on v eyi ng t o it f r o m w ith ou t the
a pp r op ri a t e s ti m u l u s ; a nd i n th e ca s e o f th e vi bri s sa this ,
a pp a ra t u s i s th e epid ermi c gr ow th w hi ch u n d er th e f or m o f ,
a h a i r p rot ru d es f rom the s a c

, A n d th a t th e gr ea t est
.
s ens itiveness m a y be Obt a i ned the ext erna l a cti on mu st be

,
e x a gg er a t ed or m u l t ipli ed by th e a pp a ra t u s w hi c h c on v ey s it
t o th e recipi ent nerv e ; a s i n th e c a s e o f th e vi bri ss a it i s by

, ,
the d ev el op m en t of a h a i r i n t o a n el a s tic l ev er th a t t ra n s ,
f o rm s the slight force a cti ng th rou gh c onsid er a bl e S p a c e on

i t s exp os ed end i nt o a g rea t er forc e a c ti ng th rou gh a s m a ll er
,
sp a ce a t i ts root e d end S i m il a rly w ith the org a n s of the

.
hi gh er s enses I n a ru dim en t a ry eye th e s lightly m odified

.
,
s en se cell h a s bu t a ru dim ent a ry nerv e t o t a k e c ogni za nce of

th e imp ressi on ; a nd t o c on cen t r a t e th e i m p res si on u p on it ,
there is nothi ng b ey ond a thick eni ng of th e epid ermis i nt o a

320 P HY SI OL OGIC A L D E VE L OP M E NT .
l ens s h a p e
-
. Bu t t h e
d ev el op e d eye s h ows u s a t ermi na ti on
o f th e ner v e gr ea tly e xp a n d ed a n d divid e d t o re ce iv e th e
ext ern a l s ti m u l u s I t s h ows u s a n i nt r ov ert ed p orti on of the

.
i ntegu m en t cont a i ni ng th e a pp a ra t u s by whi ch the ext erna l

s ti m u l u s i s c on v ey e d t o t he rec ipi en t n er v e T h e s t ru ct u re .
d ev el op ed i n t h i s s a c not only conveys th e s timu l u s bu t a l so , ,
lik e i t s h om ol ogue c oncent ra t es it ; a nd i n th e one ca s e a s

,
i n the oth er th e s t ru ct u re whi ch d oes t hi s i s a n epid erm i c

,
g row th fr om th e b ott om of th e s a c E v en w ith the ca r it i s .
th e s a m e . A g a i n we h a v e a n i nt r ov ert ed p orti on of the i n t e
gu m ent on t he w a ll s o f which th e n er v e is di s t ribu t e d i n

,
t h e p r i m itiv e ea r .T h e ot olith es c ont a i ned i n the s a c thu s

f orm ed a re b odi es whi ch a re set i n m oti on by the vib ra ti ons
,
o f th e s u rr ou n di ng w a t er a n d c on v ey th es e vib ra ti on s i n
,
a n e x a gg er a t e d for m t o the ner v es A nd th ou gh it i s not .
a ll eg e d t h a t th es e o t o lith es a re d ev el op ed f rom th e epid ermic
li n i n g of the ch a m b er yet a s if not s o d ev el op ed they a re

, , ,
c oncreti on s f rom th e c ont ent s of a n epid ermi c sac th ey mu st ,
still be reg a rd ed a s epid ermi c p r odu cts .
Wh eth er th ese di fferen ti a ti on s a r e du e wh olly t o i ndi rect

eq u ilib r a ti on or w h eth er di rec t equ ilib ra ti on h a s h a d a s h a re
,
i n w orki ng th em a re qu es ti on s th a t m u s t be l eft op en
,
.
’
P os s ibly a s h ort h a i r s o pl a c ed on a m a m m a l s f a ce a s t o be
v ery o ft en t ou ch ed m a y by c onv eyi ng e x cit a ti ons t o the
, ,
n erv es a n d v es sel s a t i t s root c au s e ext ra g rowth o f th e

,
bu lb a nd i ts a pp end a ges a nd s o th e d evel opm ent of a vi bri ss a

,
m a y be f u rth ered P os sibly t oo th e light it self to whi ch

.
, , ,
th e ti ssu es of s om e i n f eri or a n im a l s a re ev erywh ere s ens itiv e ,
m a y a i d i n s etti n g u p c er t a i n o f th e m odifi ca ti on s by whi ch
th e n erv ou s p a rt s of vi su a l org a ns a re f or m ed : p rod u ci ng a s ,
it m u st the m os t p ow erfu l effect s a t th os e p oi nt s on th e s u r

,
fa c e whi ch th e m ov em ent s Of th e a n i m a l exp ose t o th e grea t es t

a n d m os t frequ ent c ont ra s ts of light a nd sh a d e ; a nd prop a
g a ti ng f rom th os e p oi nt s cu rrents of m ol ecu l a r ch a nge

th rou gh the org a nism But it s eem s cl ea r th a t the c om pl exiti es
.
T HE O U T E R T IS S U E S OF A NI MA L S .
3 21
of th e s ens ory o rg an s a re n ot th u s expli ca bl e T h ey m u s t

.
h a v e a ri sen by the na t u ra l se l ec ti o n of f a v ou ra bl e v a r i a t i o ns .
§ 296 . A
g rou p of f a cts s ervi ng t o el u c id a te th ose pu t
,
t oget h er i n the sev era l f oreg oi n g s ecti ons h a s t o be a dd e d , .
I h a v e res erv e d thi s grou p t o the l a s t p a r tly b eca u se i t i s,
t ra n s i ti on a l— li n k s th e di fferenti a ti on s of the li tera lly ou t er

ti s s u es with t h ose of the t ru ly i nner t i ss u es T h ou gh phy s i .
ca lly i n t e rn a l th e m u cou s c oa t o f th e a lim ent a ry c a n a l h a s

,
a u a i e x t ern a lity from a phy s i o l ogi c a l po i n t o f vi ew As

q s -
.
w a s p oi n t ed o u t i n t he l a s t c h a p ter th e S ki n a nd t he a s s imi
,
l a ti n g s u rfa ce h a v e t hi s i n c om m on th a t they c om e in di rec t

,
c on t a c t w ith m a tt er s n ot be l on gi n g t o t h e o r g a ni s m ; a n d
we s a w th a t a l on g w i t h thi s c omm u nity of rel a ti on t o a li en

s u b s t a nces th ere i s a cert a i n commu nity of st ru ctu re a nd
,
d evel opment T he lik e h old s with th e li ni n gs of a l l i nt erna l

.
c a viti es a n d c a n a l s t h a t h a v e e xt ern a l o p en ing s .
T he t r a n s iti on f rom the lit er a lly ou t er ti s s u e s t o th os e

ti s su es whi c h a re i nt erm edi a t e b et ween t h em a nd th e t ru ly
i nner t i ss u es i s vi s ibl e a t a l l t h e orifi c es of th e b ody ; w here
,
S ki n a n d m u cou s m e m b ra ne a re co nti nu ou s a n d t h e one ,
p a ss es in sen s ibly i nt o th e oth er T hi s vi s ibl e conti nu ity i s

.
a ss oc i a t ed n ot s i m ply w ith a g rea t d e gree o f m or ph ol ogic a l
c onti n u ity bu t a l s o w ith a g rea t d eg ree of phy s i ol ogi ca l con

,
t i nu i ty . T h a t i s to s a y th es e lit era lly ou t er a nd qu a s i ou t er

,
-
l a y ers a re ca p a bl e O f ra pidly a s su m i n g on e a noth er s s t ru e ’
t u res a n d fu nc ti on s wh en s u bj ec t t o on e a n oth er s conditi on s

’
.
Mu cou s su rfa c es norm a lly k ept cov ered bec om e skin lik e i f
, ,
-
exp os e d t o th e a i r ; bu t res u m e m or e or l es s f u lly t h ei r
n o rm a l c h a r a c t er s w h en res t or ed t o th ei r n orm a l p os i t i on s .
T h es e a r e t ru t h s f a m ili a r t o p a th ol ogi s t s T h ey conti nu a lly

.
m eet w ith p roofs th a t p erm a nen t ev er si on of th e m u cou s

m em b r a ne ev en w h ere it i s by p r ol a p s e o f a p a rt d eeply
,
sea t e d w ithi n th e b o dy i s foll o w ed by a n a d a pt a ti on event u

,
a lly a l m os t com pl e t e : or igi n a lly m oi s t t en d er t o t h e t ou ch

, ,
a n d i rr it a t e d by th e a i r th e s u r f a c e g ra d u a lly b ec om es c ov ered
,
67
3 22 P H Y SIOL OGICAL D E V E L OP ME NT .
w i th thick dry cu ti cl e ; a nd i s then s c a r cely m ore s ensitiv e

a ,
th a n ordi na ry i nt egu m ent .
Whet her t hi s eq u ilib r a ti on b et ween new ou t er f orces a nd

r ea c tiv e i nn er f or c es whi c h is thu s di rectly p r od u ced i n i n
,
di vi du a l s i s S imil a rly p r od u ce d i n ra c es m u s t rem a i n a s a

, ,
qu es ti on n ot t o be a ns wered i n a p os itiv e w a y On th e one .
h a nd we h a v e the f a c t th a t a m ong the higher a nim a l s th ere

,
a r e ca s es o f ua si ou t er ti ss u es w hi c h a r e i n one s p ec i es
-
q
h a bit u a lly ens h ea th ed whil e i n a noth er s p eci es they a re not
,
e n s h ea t h ed ; a n d th a t th es e tw o ti s s u es th ou gh u nqu es ti on ,
a bly h om o l og ou s diff er a s m u c h a s s ki n a nd m u c ou s m em
,
b r a ne di ffer On t he oth er h a n d th ere a r e c ert a in a n a l og ou s

.
,
c h a n ges of s u r f a ce a s on t h e a bd om e n of th e H erm i t C r a b
,
-
,
which giv e wa rra nt t o the su pp os iti on th a t s u rviv a l of the

fi ttes t is th e chi ef a gen t i n es t a bli s hi ng su ch di fferen ti a ti ons
s i nc e th e a bd om en o f a H er mit C ra b b a t h e d by w a t er w ithi n
-
,
t he s h ell it oc cu pi es i s n ot e xp os ed t o p h y s i ca l con diti on s

,
th a t di rectly t end t o di fferenti a t e i t s su rfa ce from the s u rfa ce

o f th e th or a x Bu t th ou gh i n c a s e s lik e thi s l a s t we m u s t
.
,
a s s ign th e r esu lt t o the n a t u r a l s el ecti on o f v a ri a ti ons a r i s i n g
i nc id ent a lly ; we m a y I t hi nk l egiti m a t ely a s s ign the res u lt

, ,
t o th e i m m edi a t e a cti on of c h a nged c on di ti on s wh ere a s i n ,
c a s es lik e th e fi r s t w e s ee th es e p rod u c i ng i n th e i ndivid u a l

, ,
ef f ects of th e ki n ds ob s er v ed i n the ra ce .
H ow ev er thi s m a y be t h e f orc e o f the g en era l a rgu m en t

,
rem a i n s th e s a m e I n th es e ex ch a nges of s t ru ct u re a n d
.
fu ncti on b et w een the ou t er a nd qua s i ou t er ti s su es w e get -

,
u n d en i a bl e p r oo f th a t t h ey a r e ea s ily di ffer enti a bl e A nd .
s eei n g thi s w e a r e en a bl ed th e m or e cl e a r ly t o s ee h ow th e re
,
h a v e i n c ou r se o f tim e a ri s en th ose e xt rem e a nd m u lti

, ,
t u di n ou s di fferenti a ti ons of the ou t er ti s s u es which h a v e b een

gl a n ced a t .
C H A P T E R VI I I .
I
D FFE RE N T A T I IO N S AMON G T H E IN N E R T SSI UE S OF
A N IMAL S .
h a ng e f r o m th e lips tt id e of
o th ei r th e
§29 7 . TH E c ou s
in s id e i nt r od u ces u s t o a new s er i es o f i nt eres ti ng a n d i n

,
s t ru c tiv e fa c t s j oi n i n g on t o th o se wi t h w hi c h t h e l a s t c h a pt er
,
cl os ed T hey con cern th e di fferenti a ti on s of t h os e c o a t s of

.
t h e a li m ent a r y ca n a l w hi ch a s w e h a v e s een a re phys i ol ogi

, ,
ca ll y ou t er th ou gh p hy s i c a lly i nner
,
.
T h es e c o a t s a re g rea tly m odi fi ed a t difl eren t p a rt s ; a nd

th ei r m odific a ti on s v a ry g rea tly i n di fferen t a n i m a l s In .
th e l ow er typ es w h ere th ey c om p ose a s i m pl e t u b e ru nni n g

,
from en d t o end o f th e b ody th ey a r e a l m os t u n i for m i n th e i r

,
h i s t o l o gi c a l ch a ra ct er s ; bu t on a s cen di n g from th es e typ es ,
we fi n d t h e m p resenti ng a n i ncrea s i ng v a r i ety of mi n u t e

s t r uctu r es b et w een th ei r t w o en ds T h e a r gu m ent w ill be
.
a d eq u a t ely e n for c e d i f w e limit ou rs elv es t o th e l ea di n g
m odifi ca ti ons they di s pl a y i n s om e of th e high er a n i m a l s .
The su cc es s iv e p a rt s of t he a li m ent a r y ca na l a re s o pl a ce d
with r es p ect t o i t s c ont ent s th a t t h e phy s i ca l a n d ch e m i c a l
,
ch a n ges u n d er g on e by i t s con t ent s w hil e p a s s i n g fro m on e
en d t o th e o th er i n evit a bly t en d t o t r a n s for m i t s or igi n a l l y

,
h om og ene ou s su rfa ce i nt o a h et er ogen e ou s s u r fa ce Cl ea rly .

,
th e effect p r od u c ed on th e fo o d a t a n y p a r t O f th e c a n a l bv
t r it u ra ti on by a ddi n g a s ec r et i on or by a b s orbi ng i t s nu t ri
, ,
tiv e m a tt er s i m pli es th e d eliv ery of th e food i n t o th e next

,
p a rt of the c a na l i n a s t a t e m ore or l es s u nlik e i ts p revi ou s

3 23
t t s— impli es th a t th e su rfa c e with whi ch it now c om es i n

s a e
c on t a ct i s di ff erently a ffect e d by it f ro m th e p rece di n g s u r
fa c es — i m pli es th a t is a di fferenti a ti ng a c ti on
, ,
T o u se c on .
c rete l a ngu a ge f ood th a t i s b rok en d own i n the m ou th a cts

-
o n th e oes oph a gu s a n d s t om a ch i n a w a y u nlik e th a t w hi c h
it w ou ld h a ve d one h a d it b een s wa ll owed wh ol e ; th e m a s ti

ca t ed foo d t o w hich c ert a i n s o lv ents or ferm ent s a r e a dd ed
, ,
b ec om es t o the i ntes ti ne a di fferen t s u b s t a nc e f ro m th a t whi ch

i t m u s t h a v e o th erwis e b een ; a n d the a lt ered f oo d res olv ed ,
by th es e a dditi ons i nt o its p r oxim a t e p ri ncipl es c a nnot h a v e ,
th ose p roxim a t e p r inc ipl es a b s orbe d i n th e next p a rt of th e

i nt es ti ne with ou t the rem ot er p a rt s being a ffect ed a s th ey
,
wou ld not h a v e b ee n i n the a b s enc e o f a b sorpti on I t i s t ru e .
th a t i n d ev el op e d a li m ent a ry c a na l s su ch a s the rea s oni ng

,
,
h ere t a citly a s su m es th es e m a rk ed s u ccess iv e di fi erenti a ti on s

,
o f th e f oo d a r e th e m selv es th e res u lt s o f re es t a bli s h e d

p
-
differenti a ti ons i n the s u c c es s iv e p a rt s of the c a na l Bu t it i s .
a l s o t ru e th a t a cti on s a nd rea c ti on s lik e t h ose h ere s o defi ni te
l y m a rk ed mu s t g o on i n d efi nit ely i n a n u n d e v el op ed a l i m en
,
t a ry ca na l I f th e f oo d i s ch a nged a t a l l i n the c ou rs e of i ts
.
t ra ns i t which i t m u s t be if the c rea t u re is to liv e by it th en it

, ,
c a nnot bu t a ct di s s i m il a rly on th e su cces s iv e t ra cts of the

a li m ent a ry c a na l a n d c a nn ot bu t be di s s i m il a rly rea ct ed on
,
by th em I nevit a bly th eref ore the u nif orm ity o f the s u r fa ce

.
, ,
m u s t l a ps e i nt o g rea t er or l es s m u lti formity : the differenti a

ti on of ea ch p a rt t en di ng ever t o i n iti a t e differenti a ti ons of
o th er p a rts .
N ot i nd eed th a t th e impli ed p roc es s of di rect equ ilib ra

, ,
ti on ca n be rega rd ed a s the s ol e p r oces s I ndi rect eq u ilib ra .
ti on a ids ; a n d, d ou btl ess th ere a re s om e of th e m odific a ti on s

,
whi ch only i ndi rec t equ ilib ra ti on ca n a c c ompli s h B ut w e .
h a v e h ere one u n qu es ti ona bl e c a u s e— a c a u se th a t i s k nown

t o work i n i ndivid u a l s c h a n ges o f th e ki n d a ll eged
, Wh ere .
,
fo r i ns t a nc e ca n cerou s di s ea s e of the oesoph a gu s s o na rr ows

,
th e p a s s a g e i nt o the s t om a ch a s to p rev ent ea sy d es c ent

o f th e f o o d th e oes oph a gu s a b ov e the Ob s t ru c ti on bec om es
,
T HE INNE R T I SS U E S OF A NI MA L S .
3 25
l g d i nt o a ki nd of p ou ch ; a nd the i nner s u rf a c e of thi s

en a r e
p ou ch be gi n s t o s ec ret e j u ice s th a t p rod u ce i n the food a ki n d

of ru d e dig es ti on A g a i n s t rict u re of the int es ti n e w h en i t
.
, ,
a ri s es g ra d u a lly is f oll owe d by hyp ert rophy of the m u s c u l a r

,
co a t of the i nt es ti ne a b ov e the c ons t r ict ed p a rt : the ordi na ry

pe ri sta ltic m ov em ent s bei ng i ns uffic i en t t o force the f ood
f orwa rd s a nd t he l odged food s ervi ng a s a c ons t a n t s timu lu s
,
t o c ont ra cti on th e m u s c u l a r fib res h a bit u a lly m ore ex erc i sed

, , ,
be c om e m ore bu lky T he d ed u cti on f ro m genera l p ri ncipl es

.
bein g thu s i nd u c tively enfor ce d we c a nn ot I think re s i s t

, , ,
t he c onc l u s i on th a t the di rect a cti on s a nd rea cti ons be t wee n

the f oo d a nd the a lim ent a ry c a na l h a v e bee n l a rgely i n s tru
m ent a l i n es ta bli s hi n g the c ont ra s ts a m ong its p a rt s A nd .
we s h a ll h old thi s vi ew with the m ore c onfid en c e on obs erv

i ng h ow s a ti s fa ct orily i n p u rs u a nc e of it we a re ena bl ed t o
, ,
e xpl a i n one Of th e m os t s t riki n g o f th es e differenti a ti on s ,
whi c h we w ill t a k e a s a typ e of the c l a ss .
T he gi zza rd of a bi r d i s a n exp a nd ed p orti on of th e a l i m en

t a ry c a na l s p ec i a lly fitt ed t o giv e th e f oo d th a t t rit u ra ti on
,
w hi c h t he t oothl es s m ou th o f a bi r d ca nnot giv e B esid es .
h a vi ng a g rea t ly d ev el oped m u s c u l a r coa t thi s gr i ndi ng

-
,
c h a mbe r i s li ne d w ith a thick h a r d c u ti c l e ca p a bl e o f bea r

, ,
i ng the f ri cti on o f the p ebbl es s wa ll owed t o serv e a s g r i nd

s tones . T hi s di fferenti a ti on o f the m u c ou s coa t i n t o a ridged
a n d t u be rc l ed l a y e r Of h or n y m a tt er— a diff eren ti a ti on w hich
,
i n the a na l og ou s org a n s of ce rt a i n M ol l us ca is c a rri ed t o th e ,
e xt ent of p rod u c i ng f ro m this m emb r a ne c a rtil a gi n ou s pl a t es

,
a nd ev en t ee th— v a r i es i n bi r ds o f differen t ki n d s a c c or di ng
,
t o th ei r f ood . I t is m od era t e i n bi rd s th a t fee d on fle s h a nd

fi sh a nd ext rem e i n gra n iv or ou s bi rd s a nd o th ers th a t liv e
,
on h a r d s u b s t a n c e s H ow d oes this imm ens e m odific a ti on of

.
th e a li m ent a ry c a na l ori gi na t e ? I n th e st om a c h
o f a m a mm a l th e m a cera ti n g a nd s olv ent a cti on s a re u n it e d
,
wi th th a t t ri tu r a ti ng a cti on which fi nish es wh a t the t ee th

h a ve m a i nly d on e ; bu t i n th e bi rd u na bl e to m a s ti ca t e thes e
, ,
i nt ern a l fun cti on s a re s pe ci a li zed a nd whil e the crop i s the

,
3 26 P H Y SIOL OGI CA L D E VE L OP ME N T .
m a cera t n i g ch a m b er th e
gi zza rd b ecom es a ch a m b er a d a pt ed
,
t o t rit u ra t e m or e effectu a lly T h i s a d a p t a ti on requ i res s i m ply

.
a n e x a gg era t i on of c ert a i n s t ru ct u res a nd a ct i on s w h i c h
ch a ra ct eri ze s t om a ch s i n genera l a n d i n a l es s d eg ree , , ,
a li m en t a r y c a n a l s th rou gh ou t th ei r w h ol e l en gth s T he .
m a s s iv e m u s c l es o f t h e gi zz a r d a r e s i m ply e xt r em e d ev el o p
m en t s of th e mu s c u l a r tu ni c whi ch i s a l rea dy cons id era bly

,
d ev el op ed ov er th e s t om a ch a n d i n c l oses a l s o th e oes oph a g u s

,
a nd t h e i nt es ti ne T h e i n d u r a t ed li n i ng Of t he gi zza r d
.
,
t hi ck ened i nt o h orny b u t t on s a t th e pl a ce s of s ev eres t p res

s u re ,
i s n ot hi ng m ore th a n a grea tly s t rengt h ene d a n d
m odifi ed epith eli u m A n d the g ri n di ng a ct i on o f the gi zza rd
.
i s bu t a s p ec i a li ze d form o f th a t rhy th m i ca l cont ra cti on by

w hi ch a n ordi n a r v st om a c h k nea d s th e c ont a i n e d f ood a n d ,
w hi c h i n th e oesoph a g u s effect s th e a c t o f s w a ll ow i ng whil e ,
i n the i nt es t i ne it b ec om es th e p eri s t a lti c m oti on A lli e d a s .
th e gi zza r d th u s c l ea rly i s i n s tru c t u re a n d a c ti on t o the

s t om a ch a n d a li m en t a ry c a na l i n g enera l ; a n d c a p a bl e o f
b ei n g gra d u a ll y di ff erent i a t ed f rom a s t om a c h w h ere a g row

i ng h a bit o f s w a ll ow i n g f oo d u nm a s tic a ted enta il s m ore
t rit u r a ti on t o be p er for m ed b efore th e f ood p a s s es th e pyl oru s ;
the qu es ti on i s — D oes thi s ch a n ge o f s t ru c tu re a ri se by di rect
a d a pt a ti on ? T h ere i s w a rra nt for the b eli e f th a t it d oes .
Bes id es s u ch c o ll a t era l e vid enc e a s th a t m u cou s m em b r a ne

b ec om es h orn y on the t oothl es s gu m s of ol d p eopl e w h en ,
s u bj ect t o c onti nu a l r ou gh u sa g e a nd th a t th e m u s c u l a r co a t
,
o f th e i nt e s ti ne thick en s w h ere u nu su a l a ctivity i s d e m a nd ed
o f it ,
w e h a v e th e di rec t evid en ce o f exp er i m ent H u n t er .
h a bit u a t ed a s ea gu ll t o fee d on g ra i n a nd f ou n d th a t th e
-
,
lin i ng of i ts gi zza rd b eca m e h a rd ened whil e th e gi zz a rd ,
m u s cl es d ou bl ed i n thicknes s A lik e ch a n g e i n the di et o f

.
a ki t e w a s foll ow ed by lik e res u lt s C l ea rly if di fferenti a ti on s

.
,
s o p r o d u ced i n th e i n divid u a l s of a r a c e u nd er ch a n ged h a bit s ,
a re i n a ny d e gr ee i nh er it a bl e a s t ru ct u r e lik e a gi zz a r d w ill
,
o rigi na t e th rou gh th e di re ct a c ti on s a n d r ea cti ons b et wee n
t he f ood a n d th e a li m ent a ry c a na l .
THE I NNE R T I SS UE S OF A NI MA L S .
327
A n oth er c a se— a v ery i nt eres ti ng one s om ewh a t a lli ed t o ,
t h i s— i s p res ente d by the ru m i n a ti n g a n i m a l s H ere s ev er a l .
dil a t a ti ons of the a li m ent a ry ca na l p reced e the t ru e s t om a ch ;

a nd i n th e m l a rg e q u a n ti t i es o f u n m a s ti c a t e d f oo d a r e s t or ed ,
t o be a ft erwa rd s ret u rned t o t he m ou t h a n d m a s ti ca ted a t

l ei su re Wh a t con di ti on s h a ve m a d e thi s S p eci a li za ti on
.
a dv a n t a g eou s ? a n d by w h a t p r oc es s h a s it b e en e s t a bli s h ed ?
T o b oth th es e qu es ti ons th e fa c t s i ndi ca t e a ns w ers w hi c h a re

n ot u nsa ti s f a c t o ry . C rea tu res th a t o bt a i n th e i r
f ood v ery i rregu l a rly— n ow h a vi n g m ore th a n t h ey c a n con
s u m e a n d n ow b ei n g for l on g p er i od s w i t h ou t a n — m u s t
, y ,
i n th e fi rs t pl a ce be a pt w h en v ery h u ng ry t o e a t t o t he
, , ,
ext re m e li m it s o f th ei r ca p a c i t i es ; a n d m u s t i n t h e s e c on d ,
pl a ce p r ofit by p ecu li a riti es which ena bl e t h em t o c omp ens a t e

,
t h em se lv es fo r l ong f a s t s p a s t a nd fu t u re
, A p erch w hi ch .
,
when its s t om a ch i s f u ll of y ou n g f rog s g oes on filli ng i ts ,
oes oph a gu s a l s o ; or a t rou t w hi ch ri s i n g t o th e fi s h er m a n s

’
,
fly p rov es wh en t a k en off th e h ook to be fu ll of wo rm s a nd

,
i ns ect l a r vae u p t o the v ery m ou th g a i n s by i ts a bili ty to t a k e

-
,
i ns u ch u nu su a l s u ppli e s o f food w h en it m ee t s w ith th em

o bvi ou s ly th r iv es b ett e r th a n it wo u ld do c ou ld it nev er ea t
m ore th a n a s t om a ch fu l T h a t thi s a bility t o feed grea tly i n

.
ex ces s o f im m edi a t e r eq u i re m en t i s one th a t v a r i es i n i ndi

,
v i du a l s Of th e s a m e ra c e we s ee i n th e m a rk e d c ont ra s t
,
b etw een ou r own p ow ers i n thi s res p ec t a nd th e p ow ers of ,
u n c ivili zed m en ; wh os e f a s t i n g a n d g or gi n g a re t o u s s o
a st on i s hi n g Ca rryi n g wi t h u s th es e con s id er a ti ons we s h a l l

.
,
n ot be s u rp r i s ed a t fi ndi n g dil a t a ti on s o f t he oesoph a g us i n
vu lt u res a nd ea gl es whi ch get th ei r p rey a t l ong i nt erv a ls

,
i n l a r ge m a ss es ; a nd w e m a y n a t u ra lly l oo k for th em t oo i n , ,
bi rd s lik e pigeons w hi ch c omi ng i n fl ock s u p on occa s i ona l

, ,
s u ppli e s of gra i n i ndividu a l l y p r ofit by d ev ou ri ng the

,
g rea t es t qu a ntity i n a giv en ti m e N ow wh ere th e t rit u ra ti on

.
o f th e f oo d i s a s i n th es e c a s es c a rr i ed on i n a l ow er p a rt o f
, ,
th e a li m ent a ry c a na l n othi ng f u rt h er i s req u i red th a n th e

,
st oring ch a mber ; bu t for a m a m m a l h a vi ng its g ri nding

-
,
3 28 P H Y S IOL OGI CA L D E V E L OP ME NT .
a pp a ra t u s i n i ts m ou th t o ga i n by th e h a bi t of hu rri edly
,
s w a ll ow i ng un m a s t ic a t e d foo d it m u s t a l s o h a v e th e h a bit o f
,
regu r git a ti n g t h e f oo d for s u b s equ en t m a s ti ca ti on T hi s .
c or rel a ti on of h a bi t s with th ei r a ns weri ng s t ru c t ures m a y , ,
a s we s h a ll see ,
a r i s e i n a v ery s impl e w a y T he
.
s t arti ng p oi nt of th e e xpl a n a ti on is a fa m ili a r f a ct— the f a ct
th a t indiges ti on O ft en res u lti ng f rom e x ces s of food i s a pt

‘
, ,
t o ca u s e th a t rev ers ed p eri s t a ltic a cti on kn own a s v omiti ng .
F rom this we p a ss t o the f a ct a l s o w i t hi n the exp eri enc e o f

,
m os t p ers ons th a t d u ri n g s light i ndiges ti on the s t om a ch

,
s om eti m es q u i etly reg u r git a t es a s m a ll p a rt o f i t s c on t ents a s
fa r a s the b a ck o f th e m ou th— givi n g a n u n pl ea s a n t a c qu a i nt

a n ce wi t h t he t a s t e of t h e g a s t r ic j u i ce s E xc epti ona l f a ct s o f
.
t h e s a m e c l a s s h elp t he a r gu m en t a s t e p f u rt h er T h ere a re
c ert a i n i ndividu a l s wh o a re ca p a bl e of ret u rni n g a t will a , ,
g rea ter or s m a ll er p orti on of th e c ont ent s of the di ges ti ng

s t om a c h i nt o t h e c a vity o f th e m ou th .I n s om e of th es e
c a ses the exp u l s i on of the food ha s req u i re d a vi ol ent effort
,
.
I n t h e m a j ority it h a s b een ea s ily e v ok e d or s u pp res sed .
Whi l e i n o thers it h a s b een a lm os t u ncont roll a bl e ; or its

,
n o n oc cu rrenc e a t th e h a bit u a l ti m e h a s b e en fo ll owed by a

-
”
p a i nfu l feeli ng of fu l nes s or by the a ct of v omiti ng
, .
H ere w e h a v e a c ert a i n phy s i ol ogic a l a c ti on oc c a s i on a lly ,
h a pp eni ng i n m os t p ers on s a nd i n s om e d ev el op ed i nt o
a h a bit m ore or l ess p ron ou nc ed : i n dig es ti on b ei ng th e
h a bit u a l a nt ec ed ent . S u pp os e th en th a t g rega r i ou s
, ,
a nim a l s livi ng on i nnu t r itiv e f ood s u ch a s g r a s s a re s u bj ect

, ,
t o a lik e phy s i ol ogic a l a c ti on a n d a re c a p a bl e o f lik e v a ri a

,
ti ons i n the d eg ree of it Wh a t will na t u ra lly h a p pe n ?

.
T h ey w a nd er i n h erd s now ov er pl a ces wh ere food i s s c a r c e

,
a nd now c omi n g t o pl a ce s wh ere it i s a b u nd a n t S om e m a s .
t i ca te th ei r f ood c om pl et ely b ef ore s wa ll owi ng it whil e s om e ,
m a s tic a t e it i nc om pl et ely I f a n oa s i s p res ently b a red by

.
,
th ei r gr a zi n g h a s not s u ppli ed t o the w h ol e h erd a fu ll m ea l

, ,
th en the i ndivid u a l s w hich m a s tic a t e c ompl et ely will h a v e

h a d l ess th a n th os e whi ch m a s tic a t e i nc ompl et ely— will n ot
THE IN N E R T I S S U E S OF A N I MA L S .
3 29
h a v e ha d enou gh T h ose w hi ch m a s ti ca t e i ncom pl et ely a nd

.
di s tend t h ei r s t om a ch s w i t h food di ffi cu lt t o digest will be ,
li a bl e t o t h es e regu rgit a ti on s but if th ey re m a s tic a t e wh a t -
i s th u s re t u rned t o t he m ou th ( a nd w e kn ow th a t a n im a l s
O ft en ea t a ga i n w h a t th ey h a v e v omit ed ) then the ext ra ,
q u a nti ty o f food t a ken ev entu a lly m a de diges tibl e wi ll yi eld

, ,
th em m ore nou ri s hm en t th a n i s obt a i ned by th os e whi c h

m a s ti ca t e c ompl et ely a t fi rs t T he h a bit i ni ti a t e d i n this
.
n a t u ra l wa y a n d a idi n g s u r viv a l w h en foo d is s c a rce will

, ,
be a pt t o c a u s e m odific a t i ons o f the a l i m enta r v c a na l .
We k now th a t dil a t a ti ons of c a n a ls rea di ly a ri se u nd er

h a bit u a l di s t en s i ons We know th a t ca na l s h a bit u a ll y
.
di s t end ed b ec om e g ra d u a lly m ore t ol era n t of the cont a i ned

m a s ses th a t a t fi rs t i rrit a t ed th em A nd we k n ow th a t .
th ere c om m only t a k e pl a ce a d a ptive m odifi ca ti ons of thei r

s u r f a ce s. H enc e if a h a bit Of th i s ki n d a nd the st ru ct u ra l
ch a nges resu lti ng f rom it a re in a ny d egr ee i nh erit a bl e it is
, ,
cl ea r th a t i nc rea s i n g i n s u cces s ive genera ti ons b oth imm e

, ,
d i a t el y by the c u m u l a tiv e efi ect of r ep etiti on s a n d m edi a t ely

by s u rviv a l o f the i n divid u a l s i n w hi ch they a re m ost decided ,
they m a y g o on u n til th ey end i n the p ec u li a riti es whi c h

Ru m i n a nt s di s pl a y .
29 8 . T h erest ru ct u res be l ongi ng to the s a me grou p

a re
w hi c h ca nn ot h ow ev er be a c c ou nt ed for i n thi s w a y
, , T h ey .
a re th e or g a n s th a t s ec r et e s p eci a l p ro d u ct s f a ci l it a ti ng dig es
ti on— the liv er p a n c rea s a n d v a ri ou s s m a ll er gl a nd s A l l

, , .
th es e a p pend a ges of th e a li m ent a ry c a na l l a rge a nd i n d e ,
p en d ent a s s om e of th em s ee m rea lly a ri s e by di fferenti a ,
ti ons f r om its c oa ts The p ri m ord i a l liver c ons i s ts of nothi ng

.
m ore th a n bil e cell s s ca tt ered a l on g a t ra ct of the i nt esti na l

-
su r f a c e
. A c cu m u l a ti on of th es e bil e c ell s i s a ccom p a n i ed by
-
i ncrea sed gr owt h of the s u r f a ce which be a r s th e m— a g rowt h

whi ch a t fi r s t t a k es the form of a cu l de s a c h a vi n g a n ou t s id e
- -
,
th a t p r oj ect s f rom th e i nt es ti ne i n t o th e peri vi s c era l c a vity -

.
A s th e m a ss o f bil e c ell s b ecom es g rea t er th ere a ris e s e

-
,
330 P H Y SI OL OGI CA L D E V E L OP ME NT .
c on da ry l a t era l ca viti es Op eni ng i n t o the p ri m a ry on e a nd ,
thr ou gh it i nt o the i nt es ti ne ; u ntil ev ent u a lly th es e c a viti es

, ,
w ith th ei r coa t i n g s of bil e cell s b ecom e r a m i fyi n g d u c t s d i s

-
,
t ribu t ed th rou gh the s olid m a ss we k now a s a liver H ow is .
thi s di fferent i a ti on ca u s ed ?
B efo re a t t em pti n g a ny a nsw er t o thi s qu es ti on it i s ,
r equ i s it e t o i nqu i r e th e n a t u r e o f bil e I s th a t which the

.
liv er th rows i nt o the i nt es ti nes a w a s t e p ro du ct O f the orga ni c

a c ti on s ? o r i s it a s ec r eti on a idi n g dig es ti on ? or i s i t a mix
t u re o f th es e ? Mod ern i n ves tig a ti ons imply th a t it i s m os t

l ik ely th e l a s t . Th e liv er i s f ou n d t o h a v e a c om p ou n d fu n c
ti on B erna rd h a s p rov ed t o th e s a ti s fa cti on of phy s i ol ogi s t s
.
,
h t
t a t e e h r g o es o n i n it a for m a ti o n o f gly c o g e n— a s u b s t a n ce
whi c h i s t ra ns form ed i nt o su g a r b ef ore it l ea v es th e liv er a nd

i s a ft erwa r d s ca rri ed a wa y by th e bl ood t o ev ent u a lly d i s
a pp ea r i n t he a c tiv e org a n s c hi e fly th e m u s c l es
,
I t i s a ls o
.
S h own exp erim en t a lly th a t th er e a r e g enera t e d i n t he liv er

, ,
c ert a i n bili a ry a c id s ; a n d by th e a i d e i th er o f th es e or o f
s om e oth er comp ou n d s it i s c l ea r th a t bil e render s c ert a i n

,
m a t er i a l s m ore a b s orb a bl e I ts effect on fa t i s d em ons t ra bl e

.
ou t o f t h e b o dy ; a nd th e g rea tly di m i ni s h e d a b s or pti on o f
fa t fro m t he food wh en the di sch a rg e o f bil e i nt o th e

i nt es ti ne i s p reven t ed i s p r ob a bly one of the ca u s es of th a t
,
pi ni ng a w a y w hich resu lts B u t whil e rec ogn i zi n g the f a c t

.
th a t the bile c onsi s t s i n p a rt of a s olv ent or s olv ents a i d

, ,
i ng diges ti on th ere is a b un d a n t evid ence th a t on

,
e e l em en t
o f it is a n effe t e p ro d u ct ; a nd p r ob a bly thi s is t h e p r i m a r y
el em en t . T he y ell ow green s u b s t a n c e c a ll ed biliv erdi ne i n

-
h erbiv ora a nd bili ru bi n i n m an a nd c a rniv ora which giv es ,
i ts c ol ou r t o bil e is a p rod u ct the grea t er p a rt of whi ch i s

,
n or m a lly c a st ou t f ro m th e syst em c onti nu a lly a s i s s h own ,
by th e c ont ra st b etw een th e nor m a l a nd a b norm a l c ol ou r s

of faec a l m a tt er s ,
a n d a s i s s till m or e st riki ngly s h own by
t h e effects on th e syst em wh en th ere i s a s t opp a ge of the

ex creti on ,
a nd a n a tt a c k o f j a u n dic e H en c e we a r e w a r
.
ra n t e d i n cl a s s i n g biliv er di ne a s a w a st e p rod u ct a nd w e ,
THE INNE R T ISS U E S OF A NI MA L S .
3 31
m ay fa i rly i nfer th a t the exc reti on of it i s the ori gi na l

f u nc ti on o f the li v er .
One fu rth er p reli m i n a r y i s req u i s it e We mu s t for a .
m om en t ret u rn t o th ose phy s i co ch em ic a l d a t a s et d own i n

-
t h e fi rs t c h a pt er of this w or k 7 We there sa w th a t
t he c o mpl ex a n d l a rge a t om ed c oll oid s whi c h m a i nly c omp os e
-
livi ng org a ni c m a t ter h a v e ext rem ely littl e m ol ecu l a r m o

,
bi l i ty ; a nd c onsequ ently ext rem ely littl e p ower o f diff u s i n g

, ,
them selv es Wh erea s w e s a w not only th a t th os e a b sorb ed

’
m a t t ers g a seou s a n d liqu id whi ch fu rth er the d ecom p os iti on

, ,
o f livi ng or g a n i c m a tt er h a v e v ery high diffu s ibiliti es bu t

, ,
a l s o th a t th e p ro d u c t s Of th e d eco mp os iti on a r e m u ch m o re
di ffu s ibl e th a n the com p onen t s of livi ng org a ni c m a tt er .
A nd w e s a w th a t a s a c on s eq u ence of thi s the t i s su es giv e

, ,
r ea dy ent ra nce t o th e s u b s t a n ces w hi c h d ec omp os e th em a nd ,
r ea dy e xit t o th e s u b s t a nce s i n t o w hi ch th ey a re d eco mp o s e d .
H en ce it f o ll ow s t h a t u n d er i ts i n iti a l f orm u n c om pli ca t ed

, ,
by nerv ou s a n d oth er a genci es the es c a p e of eff et e m a t ters

,
f rom th e org a ni s m i s a phy s i ca l a cti on p a ra ll el t o th a t which

,
goes on a m on g mix ed c oll oid s a n d c ry s t a ll oid s th a t a re dea d

or e v en i n org a nic E x creti on is a spec i a li zed f or m of this
.
s p ont a neou s a c ti on ; a nd w e h a v e t o i n qu i re h ow the sp ec i a l
i za ti on a ri s es .
T w o ca u s es c on s pi re to es t a blish it Th e fi rs t i s th a t th es e
.
p rod u ct s o f d ecomp os iti on a re di ffu s ibl e i n w id ely different

d egr ees Whil e the ca rb onic a cid a nd w a t er p er m ea t e the
.
ti s su es wi th ea s e i n a l l di recti ons a nd es c a p e m ore or l es s ,
f r om the exp osed s u r fa c es u rea a nd oth er w a s t e su b s t a nc es

, ,
i nca p a bl e O f b ei n g v a p ori zed c a nnot es ca p e th u s r ea dily

, .
T he sec on d i s th a t th e di fferent p a rt s of th e b ody b ei ng ,
su bj ec t t o di f fer en t phy s i c a l c on diti on s a re f r o m th e ou t set ,
s u r e sev er a lly t o f a v ou r th e exit of th es e v a r i ou s p ro d u cts of
d ec om p os iti on i n v a r i ou s d egrees H ow th es e ca u s es m u s t .
h a v e co op er a t ed i n l oca li zing th e e x cr eti on s w e s h a ll see on

-
,
rem emb eri n g h ow th ey n ow co op er a t e i n l oca li z i n g t h e Sep a

-
ra ti on of m orbid m a t er i a l s T h e ch a ra ct eri s tic s u b s t an ces of

.
3 32 P H Y S I OL OGICA L D E VE L OP ME NT .
g ou t a nd rheu m a ti s m h a v e thei r h a bi tu a l pl a ce s o f d ep os it . .
T u b ercu l ou s m a t ter t h ou gh it m a y be p res ent i n v a ri ou s

,
o rg a ns gr a vit a tes t owa r d s s om e m u ch m ore t h a n t owa r d s

,
o th er s Cer t a i n p r od u c t s of di s ea s e a re h a bit u a lly g ot ri d of

.
by the s kin i ns t ea d of c oll ecti ng i ntern a lly M os tly these

,
.
,
h a v e s p ec i a l p a rt s of the s ki n which th ey a ffect ra th er th a n

t he res t ; a nd th ere a re th ose w h i c h by b rea ki n g ou t sym ,
m et ri ca lly on the two s id es o f the b ody s h ow h ow d efi nit ely ,
t he pl a ce s of th ei r e x creti on a re d et erm i ned by c ert a i n

fa v ou ri n g c onditi ons which c orres p onding p a rt s m a y be pre
,
s u m e d t o fu rn i s h i n eq u a l d eg ree s Fu rt h er it is t o be
.
,
ob s er v e d o f th es e m o r bid s u b s t a nc es ci rcu l a t i n g i n the bl oo d ,
th a t h a vi ng once c om m enced segrega ti ng a t p a rti cu l a r pl a ces ,
th ey t en d t o c ontin u e s eg reg a ti ng a t th ose pl a ces A s s u m .
i ng th en a s w e m a y fa i rly d o th a t thi s l oca li za ti on o f

, , ,
'
e x c r eti on w hi c h we see con ti nu a lly c o mm enc i n g a f res h with

,
m orbid m a tt ers h a s a l wa y s g one on with the m a tt ers pro

,
du ced by th e w a s t e O f th e ti s su es l et u s t a k e a fu r th er s t ep
, ,
a nd a s k h ow l oc a li za ti on s b ec om e fixe d Oth er thi ngs eq u a l

.
,
th a t whi ch from i t s phy s i ca l c onditi ons i s a pl a c e of l ea s t

res i s t a n ce t o the exit o f a n effet e p r o d u ct w ill t en d t o b ec o m e ,
es t a bli s h ed a s t he pl a c e o f e x c r e ti on ; s i nce th e ra pid exit
o f a n eff e t e p ro d u c t w ill p rofit th e or g a n i s m Oth er thi ng s .
eq u a l a pl a c e a t w hi c h th e ex cr e t e d m a tt er p r od u c es l ea s t
,
d et ri m en t a l effec t will b ec om e the es t a bli s h ed pl a ce I f a t .
a ny p oi n t th e e xc rete d m a tt er p rod u c e s a benefi c i a l ef f ec t ,
then oth er thi ng s equ a l surviv a l Of the fitt es t will d et erm ine
, ,
it to thi s p oi nt A nd i f fa c ility of es ca p e a nywh ere g oes

.
a l ong with u tili z a ti on o f th e es c a pi ng s u b s t a n ce th en o th er , ,
thi ngs equ a l the exc reti on will be th ere l oc a l ized s till m ore
,
d eci s iv ely by su rviv a l of the fitt es t .
S u ch be i n g th e c on diti ons of th e p robl em l et u s a sk wh a t ,
will h a pp en with the li ni ng m emb ra ne of the a lim ent a ry

c a na l . T hi s phy s i ol o gi ca lly c onsid ered i s a n ext ern a l su r
, ,
f a c e ; a nd m a tt ers th rown off from it m a k e th ei r way ou t of

the b ody I t is a l s o a su rf a c e a l ong which i s m ovi ng the f ood
.
T HE INN E R T ISS U E S or A NIMA L S .
333
to be diges t ed N ow a m ong the v a ri ou s wa st e p ro du cts

.
,
c on ti nu a lly es ca pi ng fro m the livi ng ti s s u es s om e of the ,
m ore c ompl ex ones not v ery s ta bl e i n c omp os iti on a re lik ely

, , ,
if a dd ed t o t he f ood t o set u p c h a nges i n it S u c h c h a n g es

,
.
m a y eith er a i d or hi nd er th e p r ep a ra ti on o f t h e f oo d fo r
a b s orpti on I f a n effet e m a tt er m a ki n g i ts e xit th rou gh th e

.
,
w a ll of th e i nt es ti ne hi nd ers t he dig es tiv e p ro c es s t he en

, ,
feebl em ent a nd di s a pp e a r a nc e o f i ndivid u a l s i n which thi s

h a pp ens will p rev ent th e in tes ti ne from bec omi ng the es t a
,
bl i s h ed pl a c e for i t s e xit Whi l e i f it a id s the diges tive

.
p roces s the i nt es ti ne will for convers e rea s on s b ecom e m ore

, , ,
a n d m o re t h e pl a c e t o w hich i ts e xit i s li m it ed E qu a lly .
m a ni fes t i s it th a t i f th ere i s one p a rt of thi s a li m enta ry

ca n a l a t w hi c h m ore th a n a t a ny o th er p a rt t he fa v ou r a bl e
, ,
ef fec t resu lt s thi s wi ll b ecom e the pl a ce of ex c re ti on

,
.
T h u s th en rev er ti n g t o th e c a s e i n qu esti on w e m a y
, , ,
u n d ers t a n d h o w a p r o d u ct t o be c a s t ou t s u eh a s biliv er di n e
’
, ,
i f it eith er di rectly or i n di rectly s erv es a u s efu l p u rp os e ,
wh en p ou red i nt o a p a r ti cu l a r p a rt of the i nt es ti ne m ay l ea d ,
t o the f or m a ti on o f a p a t c h of ex c reti n g cell s on its wa ll ;

a nd o nc e thi s pl a ce Of ex c re ti on h a vi ng been e s t a bli s h ed th e ,
d ev el opm en t of a liv er is S imply a qu es ti on of ti m e a nd

n a tu ra l s el ecti on .
29 9 di fferen ti a ti on of a noth er ord er occu rri ng i n the

. A
a li m en t a r y c a n a l i s th a t by w hi c h a p a rt o f it i s d e v el o p e d
,
i nt o a l a t era l ch a m b er or ch a m b ers thr ou gh whi ch c a rb onic ,
a cid exh a l e s a n d o xyg en i s a b s or b e d C om p a r a tiv e a n a t om y

.
a nd em b r y o l o gy u n it e i n s h ow i n g t h a t a l u n g i s f o r m ed j u s t ,
a s a liv er or o th er a pp en d a ge Of th e a li m en t a ry ca n a l i s
form ed by th e gr owth of a h oll ow bu d i nt o t h e p er i vi s cera l

,
-
ca vity or s p a ce b et w een th e a li m en t a ry ca n a l a n d t h e w a ll o f
,
th e b ody T h e i nt er i or o f thi s bu d i s s i m ply a cu l de s a c o f

.
- -
th e a li m ent a ry ca na l with th e m u cou s li ni n g of w hi c h i t s

,
own m u c ou s li n i n g i s con ti nu ou s A n d the d ev el op m ent o f

.
this cu l de s a c i nt o an a i r ch a mber s impl e or eom p ou nd i s

- - -
, ,
.
3 34 P H YSI OL OGI CA L D E VE L OP ME NT .
m erely a g rea t ext ens i on of a rea i n the i nterna l s u rf a c e Of

t h e cu l de s a c a l ong w ith th a t s p eci a li za ti o n w hi ch fi ts it
- -
,
for exc reti ng a nd a b s orbi ng s u bst a nc es di fi erent f ro m th os e

w hich oth er p a rts o f the m u c ou s s u rf a ce e xc ret e a n d
a b s orb . T h es e l a t er a l a i r ch a mb ers u ni v ers a l -
,
a m o ng the high er Ver tebra ta a n d v ery g en era l a m on g th e
l ower a nd every wh ere a tt a ch ed t o the a lim ent a ry c a na l

,
b et ween the m ou th a nd the st om a ch h a v e not i n a l l c a s es the ,
res pi ra t ory f u n cti on I n m os t fi s h es th a t h a v e th em th ey

.
a re wh a t we k n ow a s s wim bl a dd ers -
I n s om e fi s h es t he .
c a viti es of th es e s wi m bl a dd ers a re c ompl et ely S hu t Off f rom

-
t he a li m en t a ry c a n a l : nev erth el ess s h ow i n g by th e c om m u ni ,
c a ti ons whi ch th ey h a ve with it du ri ng the em b ry oni c s t a ges ,
t h a t th ey a re origi n a lly di ver ti cu l a f ro m it I n oth er fi s h es .
th ere i s a p erm a nen t duc tus p neu m a ti ca e u nit i ng the c a vity ,
o f the s wim bl a dd er w i t h th a t o f t h e g ull et : t h e fu nc ti on

-
,
h owev er b ei ng s till not res pi ra t ory i n a n a pp rec i a bl e d egree,

‘
i f a t al l .B u t i n cer t a i n s till ext a nt r ep res ent a tiv es o f th e

s a u roid fi s h es a s t h e L epi dos teu s th e a i r bl a dd er i s
, , divid e d -
”
i nt o tw o s a cs th a t p os s ess a cell u l a r s t ru ctu re an d the ,
“
t ra ch ea which p roceed s f ro m it op ens high u p i n the th roa t -

,
”
a n d is s u rr oun d ed w ith a gl otti s I n th e A m p hi bi a th e
.
c orr e s p on di n g org a ns a re ch a mb ers o v er th e su rfa c es o f w hich
there a re s a ccu l a r d ep res s i ons i ndic a ti ng a t r a ns iti on t ow a rd s

,
th e a i r c ell s ch a ra c t er i zi ng l u ng s ; a nd a c com p a nyi ng thi s

-
a dv a n c e w e s ee a s i n t h e com m on T r i ton t h e h a bit o f c o m i ng

, ,
U p t o th e su r fa c e a nd t a ki ng d o wn a f resh s u pply of a i r i n
pl a ce of th a t di s ch arged .
H ow a r e th e i n t er na l a i r ch a m b ers res pi r a t ory or n on

-
,
r es pi ra t o r y d ev el o pe d ?
,
U p w a r d s from the a m phibi a n s t a g e ,
i n whi ch t h ey a r e p a rt i a lly r efill ed a t l ong i nterv a l s th ere is ,
n o d if fi cu lty i n u n d er s t a n di n g h ow by i nfi n it es i m a l s te p s, ,
th ey p a ss i n t o c om pl ex a n d ev er m ovi ng l u n g s B ut-
.
h ow i s th e di fferen ti a ti on th a t p rod u ces th em i n iti a t ed ?

H ow c om es a p orti on o f th e i n t ern a l s u r fa ce t o be s p ec i a li z ed
for c onv ers e with a m edi u m to which it is not n a t u ra lly
T HE INNE R T ISS U E S OF A NI MA L S .
exp os ed ? T he p robl em a pp ea rs a di ffi cu lt on e ; bu t th ere i s

a not un s a ti s f a ct ory s ol u ti on o f i t .
Wh en m a ny g old fi s h a re k ept i n a s m a ll a qu a ri u m a s
-
w i t h t h ou ghtl es s c ru elty th ey freq u ently a re th ey s wi m ,
c l os e t o t h e s u rf a c e s o a s t o b rea th e th a t w a t er which i s f ro m
,
i n s ta n t t o i ns t a n t a b s orbi n g f res h o xygen I n d oi ng thi s .
they o ft en pu t th ei r m ou ths p a rtly a b ov e th e s u r fa ce s o th a t ,
i n c l os i ng th em th ey t a k e i n b u bbl es of a i r ; a nd s om etim es
they m a y be s een t o c onti nu e d oi ng thi s— the reli ef du e t o
the s light e xt r a a era ti on of bl oo d s o s ec u red b ein g the ,
s ti m u l u s t o c onti n u e A i r thu s t a k en i n m a y be d et a in e d
. .
I f a fi sh th a t h a s t a k en i n a b u bbl e t u rn s i t s h ea d d own
w a r d s the b u bbl e w ill a s cen d t o th e b a ck of i ts m ou th a n d
, ,
there l odge ; a nd c om i ng withi n r ea ch of the c ont ra cti ons of

t he (E S Op h a g u s i t m a y be s wa ll ow e d
, I f th en a m ong fi sh
.
, ,
t hu s na tu ra lly l ed u p on oc ca s i on t o t a k e i n a i r b u bbl es t h ere -

,
a re a ny h a vi n g s light di ffe r ences i n t h e a li m en t a ry c a n a l th a t
fa c ilit a t e l odg m en t of th e a i r or s light ner v ou s differenc es

,
s u ch a s i n h u m a n b ei n gs c a u s e a n a ccid enta l a c ti on t o be
”
c Om e a t r ick , it m u s t h a pp en t h a t i f a n a dv a n ta ge a ccru es
f rom the h a bit u a l d et en ti on o f a i r bu bbl es th os e i n divid u a ls
-
,
m os t a pt to d et a i n th em w ill oth er thi ng s equ a l be m ore

, ,
lik ely th a n th e res t t o su rviv e ; a nd by the s u rviv a l O f

d es cend a nts in h eritin g th ei r p ecu li a riti es i n the gr ea t es t de
g rees a nd i nc rea s i ng th em a n es t a bli s h e d s t ru ct u re a nd a n
, ,
es t a bli s h ed h a bit m a
y a r i se A nd th a t th ey do i n s om e
.
wa y a r i s e we h a v e p roo f T he c om m on L oa ch sw a ll ows a i r
.
which it a ft erw a rd s di s ch a rges l oa d ed w ith ca rb oni c a c id .
F rom a i r thu s s w a ll ow ed th e a dva n t a ges th a t m a y be

d eriv e d a re Of two ki n d s I n the fi rs t pl a ce the fish i s m a d e
.
,
s p ec ific a lly ligh t er a n d th e m u s cu l a r e ff o rt n ee d e d t o k ee p it

,
from s i n ki n g i s di m i ni s h ed— o r i n d eed i f th e b u bbl e i s of

, ,
th e right s i ze i s a l t o get h er s a v e d
, T he cont r a s t be t ween th e
.
m ovem en t s o f a Goby whi ch a fter s wi m m i n g u p t ow a rd s the

, ,
su r fa ce f a ll s r a pidly t o th e b ott o m on cea s i ng i s

, t e x erti ons ,
a nd the m ov em ents of a T rou t which rem a i n s s u sp end ed j u st

,
336 P H YS IOL OGIC A L D E V E L OP ME NT .
b a l a nci ng it s el f by s light u n d u l a ti on s o f i ts fi ns s h ow s h ow ,
g rea t a n econom y res u lt s f rom a n in t ern a l fl oa t to fi sh es ,
w h i c h s ee k t h ei r foo d i n m i d w a t er or a t th e s u rfa c e
-
H en ce .
t h e h a bit o f s wa ll ow i n g a i r h a vi n g b een i ni t i at ed i n th e w a y
d es cr ibe d we see why na t u ra l s el ec ti on will i n cert a i n fi sh es
, , ,
a i d m odifi ca ti on s o f the a li m ent a ry ca n a l f a v ou ri ng i ts

l odgm en t— m odi fi ca ti ons c on s tit u ti n g a i r s a cs -
In.
t he s ec on d pl a ce w hil e f rom a i r thu s l odge d i n a i r s a cs t hu s

,
-
d evel op ed the a dv ant a ge w ill be th a t o f fl ot a ti on only if th e

,
a i r i s i n freq u ent ly ch a n g e d or nev er c h a n ge d the a dv a n t a g e

,
will be th a t o f s u ppl e m ent a ry r es pi r a ti on i f the a i r s a cs a re -
f ro m tim e to tim e p a rt i a lly empti ed a nd refill ed T he re .
qu i rem en ts of t h e a n i m a l w ill d et er mi n e w hi ch of t he tw o
f un cti ons p red om in a t es L et u s gl a n ce a t the di fferent s et s
.
o f con di t i o n s u n d er w hi c h th ese div er g en t m o difi ca ti ons m a y
be e xp ec t ed t o a r i s e .
T he res pi ra t or y d ev el op m en t i s n ot lik ely t o t a k e pl a ce i n

fi s h es th a t i nh a bit sea s or riv ers i n whi c h the su pply of
a e ra t e d w a t er nev er fa i l s : t h er e i s n o obvi ou s re a s on wh
y
th e est a bli s h ed b ra n chi a l r es pi ra ti on s h ou ld be repl a ced by a
p u l m oni c res pi ra ti on I nd eed i f a fi s h s b ra nchi a l res pi r a ti on
.
,
’
i s a d eq u a te t o i ts n ee d s a l os s w ou ld res u lt from th e effort of

,
c om i ng to the su r fa ce for a i r ; es p eci a lly d u ri ng th ose fi rs t

s t a ges o f p u l m on ic d e v el op m en t w h en t h e ext r a a e r a ti on
a chi ev ed w a s bu t s m a ll H ence i n fi s h es s o c i rcu m s t a nc ed

.
,
t h e a i r ch a m b ers a ri s i n g i n th e w a y d es c rib ed w ou ld na t u ra lly

-
b ec om e s pec i a li zed m a i nly or wh olly i n t o fl oa t s T h ei r c on .
t a i ned a i r b ein g i n frequ ently ch a nged no a dv a n t a ge w ou ld

,
a r i s e f rom th e d ev e l op m en t o f v a s cu l a r pl e xu s es o v er th ei r
s u rfa c es ; n othi n g w ou ld be g a i ne d by k eepi n g o p en th e c om
m u ni ca ti on b et w een th em a n d th e a lim en t a ry ca n a l ; and
th ere might thu s ev ent u a lly res u lt cl os ed ch a m b ers th e

g a s eou s c ont ent s of whi ch i n s t ea d of b eing obt a i n ed fr om
,
w ith ou t w ere s ec ret ed from th ei r wa ll s a s g a ses oft en a re

, ,
f rom mu cou s m em b ra nes . C ont ra ri w i s e a qu a tic ,
v er t eb ra t es i n which t he s wa ll ow i ng o f a i r b u bbl es b ec om i ng
“
-
,
TH E INNE R T IS S U E S OF A NIMAL S .
3 37
h a bit u a l h a d l ed t o the f orm a ti on o f s a c s t h a t l odged th e

,
b u bbl es ; a n d whi c h c on ti nu e d t o i n h a bit w a t ers n ot a l wa ys

s u pplyi ng t h em w i t h su f fici en t oxygen might be exp ec t ed t o ,
h a v e t he s a c s fu rt h e r d ev el op ed a n d t he p r a cti c e of c h a ng
,
i ng t h e cont a i n e d a i r m a d e regu l a r if eith e r of t wo a dva n ,
t a ges resu lt ed— e ith er t he a dv a nt a g e of be i n g a bl e t o liv e i n

Old h a bi ta t s th a t h a d b ec om e u nt en a bl e w ith ou t thi s m odifi
c a t i on or th e a dv a n t a ge o f b ei n g a bl e t o occu py n ew h a bit a t s
, .
N ow it i s j u st wh er e th es e a dv a n t a ges a re g a i n ed th a t we
see th e p u lm onic r es pi r a ti on c omi n g i n a i d o f the b ra n chi a l
res pi ra ti on a n d i n v a ri ou s d eg ree s repl a c i n g it

, S h a ll ow .
w a t ers a re li a bl e t o th ree ch a nges whi c h c ons pi re t o m a ke

’
thi s s u pplem en t a ry res pi r a ti on b enefi ci a l T he su m m er s s u n .
h ea ts th em a nd ra i s i n g the t em p er a tu res of th e a nim a l s th ey

,
c ont a in a cc el er a t es th e ci rcu l a t i on i n th es e a ni m a ls ex a lts

, ,
t h ei r f u nc ti on a l a c tiviti es i n c rea ses th e p r od u cti on of ca r

,
b oni e a cid a nd thu s m a k es a e ra ti on of the bl ood m ore need

,
fu l tha n u s u a l Mea nwhil e th e h ea t ed wa t er i n s t ea d o f

.
,
yi eldi ng t o the highly ca rb oni z ed bl oo d b r ou ght to the

b ra nchi ae the u su a l qu a n tity of o xyg en yi eld s l es s th a n
'
u su a l ; for a s t h e h ea t o f t h e w a t e r i n c r ea s es th e qu a n tity o f ,
a i r it c on t a i n s dimin i s h es A nd thi s g rea t er d em a n d for

.
o xyg en j o i ne d w ith s m a ll er s u pply p u s h ed to a n ext rem e ,
w h ere t h e w a t er i s n ea r ly a l l ev a p ora t ed i s a t l a s t s till m ore ,
i ntensely felt i n c ons eq u ence of th e exc es s o f ca rb onic a cid

di s ch a rged by th e nu m erou s c rea t u res c ong reg a t e d i n the
m u ddy p u ddl e s th a t r em a i n H ere th en it is th a t th e h a bit
.
, , ,
o f t a ki n g i n a i r b u bbl e s i s lik ely t o b eco m e e s t a bli s h e d

-
an d ,
th e org a n s for u tili zi n g th e m d e v el op e d ; a nd h er e it i s a c ,
co r di ngl y th a t w e fi nd a l l s t a g es of the t r a n s iti on t o a e r i a l

,
res pi ra ti on T he L oa ch b efore m en ti on e d w hi c h s wa ll ow s
.
-
,
a i r fr eq u en ts s m a ll w a t ers li a bl e t o be c o ns id er a bly w a r m ed
,
.
’
Th e A m p hi pn ou s Cu chi a a n a nom a l ou s ee l sh a p e d fi sh whi ch
,
-
,
h a s v a s cu l a r a i r sa c s op en i n g ou t a t t h e b a ck of the m ou th
-
,
i s g en er a lly fou n d l u rki n g i n h ol es a n d c revi ces on th e ,

”
mu ddy b anks of m a rsh es or sl ow m ovi ng riv ers ; a nd -
68
3 38 P H Y S I OL OGI CA L D E VE L OP ME NT .
t h ou gh i t s ph ol ogi ca l eq u iv a l ent s of th os e
a i r- sa cs a re n ot m or
a b ov e d es c r ib ed yet th ey e qu a lly w ell ill u s t r a t e t h e re l a ti on

,
b et wee n s u c h or ga n s a nd th e en vi r oni ng c on diti on S till

m ore S ign ifi c a n t i s the fa ct th a t the L ep idos i r en or m u d ,
”
fi sh a s it i s c a ll ed fr om i t s h a bi t s t h ou gh it i s a t ru e fi s h
‘
n ev erth el ess h a s l u n g s Bu t it i s a m on g the A m p hi bi a th a t

.
we s ee m os t c ons pi cu ou s ly thi s rel a ti on b et ween the d ev el o p

m ent of a i r b rea thi ng org a n s a nd the p ec u li a riti es of the
-
,
h a bit a t s. P ool s m ore or l es s di s s ip a t ed a nnu a lly a nd s o

, ,
ren d er e d u n i n h a bit a bl e by m os t fi s h es a re v ery g ener a lly,
pe opl ed by th es e t r a ns iti ona l type s Ju s t a s w e see t oo .

, ,
th a t i n v a ri ou s clim a t es a n d i n v a r i ou s ki nd s of s ha ll ow
w a t er s th e s u ppl em en t a ry a e ri a l res pi ra t i on i s nee d fu l i n dif
,
ferent d egrees ; s o do w e fin d a m on g th e A m p hi bi a m a ny
s t a g es i n th e su bstit u ti on o f the one r e s pi ra ti on for t h e o th er .
T h e fa ct s t h en a re s u ch a s giv e t o th e hyp oth es i s a vr a i

, ,
s em bl a n ce g rea t er th a n cou ld h a v e b ee n e xp ec t e d
'
.
T he rel a tiv e effec t s Of di rect a nd i n di rec t equ ilib r a ti on i n

es t a bli s hi ng thi s f u r th er h et ero g eneity m u s t a s i n m a ny , ,
o th er ca s es rem a i n u n d ec id e d
,
The h a bit o f t a ki ng i n bu b
.
bl es i s s c a rcely i nterp ret a bl e a s a resu lt of s p ont a neou s v a ri a

ti on : we m u s t rega rd it a s a ri s i n g a ccid ent a lly d u ri ng th e
ef fort t o obt a i n th e m os t a erat ed wa t er ; a s b ei n g p ers ev ere d
i n be ca u se o f the reli ef Ob t a i ned ; a nd a s grow i ng by r ep etiti on
i nt o a t end ency b equ ea th ed t o offs p ri ng a nd by th em or , ,
s om e o f th em i ncrea s ed a nd t ra ns m itt ed T he form a ti on of

, .
th e fi rs t s light m odifi ca ti ons of th e a li m ent a ry ca n a l f a v ou r

i ng th e l odgm en t of b u bbl es i s n ot t o be thu s expl a i ned S om e
, .
f a v ou ra bl e v a ri a ti on i n the s h a p e of th e p a ss a ge mu s t h ere
h a v e b een the i niti a l s t ep But the g ra d u a l i nc rea s e of thi s
.
st ru c tu ra l m odifi ca ti on by the s u rviv a l of i ndivid ua l s i n

which it i s ca rr i ed f u rth es t will I thi nk be a l l a l ong a id ed
, , ,
by i m m edi a t e a d a pt a ti on T he p a rt o f th e a liment a ry ca n a l
.
p revi ou s ly k ept f rom the a i r bu t now h a bit u a lly i n c ont a c t

,
with the a i r m u s t be i n s om e d egree m odifi ed by the a cti on

,
of the a i r ; a n d th e di rectly pr od u c ed m odific a ti on i nc rea s i n g

-
,
T HE I NNE R T ISS UE S OF A NI MA L S
"
.
3 39
i n t he i ndividu a l a nd i n s u c ce s s ive i ndi vidu a ls c a nnot cea s e ,
u ntil th ere i s a c o mpl et e b a l a nce be tween the a cti on s o f th e
c h a ng e d a gency a nd t h e c h a ng ed ti s s u e .
§ 3 00 . We c om e di fferenti a ti ons a mong the t ru ly

n ow to
i nner ti s s u es — the ti ssu es which h a v e di rec t c onv ers e neither
with th e envi ronm ent nor with the f oreign s u b s t a nc es t a k en
i nt o the orga ni s m f rom the envi i onm ent T hes e sp ea ki n g
‘
.
,
b roa dly a re the tis s u es whi ch l i e b et wee n the d ou bl e l a yer

,
f orm in g the i nt egu m en t w ith i ts a pp end a ges a nd the d ou bl e ,
l a yer forming the a lim ent a ry c a na l with its di ver ti c u l a We .
will t a k e fi rs t the differenti a ti on which p rod u ces the v a s cu l a r

sy s t em .
Cert a i n for c es p r od u cin g a nd a idi n g di s t rib u ti on of liqu ids

i n a n i m a l s c om e i n t o pl a y b e for e a ny v a s c u l a r s y s t e m e xi s t s ;
,
a n d con ti nu e t o fu rt h er c i rc u l a ti on a f ter t he d ev el op m ent of
a v a s cu l a r s y s t em T he fi rs t of th es e is os m ot ic e x c h a nge
.
,
a c ti n g l oc a lly a n d h a vi n g a n i ndi rect gener a l a c ti on ; the
s ec on d i s l o c a l v a r i a ti on Of p r ess u re whi c h m ov em en t Of th e
,
b ody t hr ow s on the ti s s u es a nd thei r c ont a i ned liqu id s A .
few w or d s a re need ed i n el u cid a ti on of ea ch I f in

.
a ny c r ea t u re h ow ev er S impl e different c h a nge s a re g oi n g on

, ,
i n p a r t s th a t a re di fferent l y c onditi one d— i f a s i n a H ydra , ,
o n e s u r f a c e i s e xp o s ed t o t he s u rr ou n di n g m ediu m whil e th e
o th er s u r fa ce i s e xp os ed t o di s s olv ed f o od ; th en be t w ee n the
u nlik e liq u id s which t he di s s imil a rly pl a c e d p a rt s c ont a i n

-
,
o s m otic c u rr en t s m u s t a ri se ; a n d a m ovem ent of liqu id

th rou gh the i nt erm edi a t e ti s su e mu s t go on a s l ong a s an
u nlik enes s be t w een t he liqui ds i s k ept u p T his p rim a ry
.
c a u s e of re di s t rib u ti on rem a i ns one of the c au s es of re di s

- -
t ri bu ti on i n ev ery m ore d ev el op ed org a ni s m : the p a s sa ge Of

-
m a tt ers i nt o a nd ou t of the c a pill a ri es i s ev eryw h ere thu s

s et u p . A n d Obvi ou s ly i n p rod u ci ng th es e l oc a l c u rr ent s ,
os m o s e m u s t a l s o i ndi r ectly p r od u c e g en era l c u rr ents or a i d ,
th em if oth erwi s e p rodu ced I n th e a b s enc e of a pu mpi n g

.
or g a n ,
thi s f orce is p rob a bly a n i m p ort a nt a i d to th a t
m ov ement of t he
t itiv e liqu id s w hich the f uncti ons s et
nu r
up . H ow the s ec on d ca u s e— the ch a nges o f i ntern a l

p res su re which a n a n i m a l s m ov em ent s p r odu ce— fu rthers
’
ci rcu l a t i on will be s u ffi ci ent ly m a nif es t

, T h a t p a rts wh i c h
.
a re b ent o r s t ra i n ed nece s s a rily h a v e th ei r c on t a i n ed v es s el s
s qu eezed h a s b een s h own

,
28 1 ) a n d w h eth er the b en d or
s tr a i n i s c a u s ed a s i n a pl a n t by a n ext ern a l force or a s

, , , ,
u s u a lly i n a n a n im a l by a n int ern a l force t h ere m u st be a

, ,
th ru s ti n g of liqu id s t owa rd s pl a ces of l ea s t res i s ta n ce — c om

m onl y pl a c es o f g rea t es t con s um pti on T hi s whi ch in a ni m a ls
.
w i th ou t h ea rt s is a m a i n a gent o f c i r cu l a ti on con ti nu es t o ,
fu rt h er it v ery c on s id era bly ev en a m on g th e high es t a n i m a l s .
I n th es e the effec t b ec om es a s it were s y s tem a ti zed T he v a lv es .
i n the v ei ns nec es s it a t e p erp et u a l p r op u l s i on s t ow a r d s t he

h ea rt .
E v en i n s u ch s impl e typ es a s th e H ydr ozoa ca viti es i n th e ,
ti s su es f a i n tly i ndi ca t e a s t ru ct u re whi ch f a cilit a t es th e t ra ns

fer o f nu t r itiv e m a tt er s T h es e c a viti es b ec o m e r es erv oi r s
. .
fill ed with th e pl a s m a th a t s l ow ly oozes th rou gh the s u b s ta n ce

o f the b ody ; a n d ev ery m ov em ent o f th e a n i m a l a ccomp a n i e d ,
a s it m u s t be by ch a n g e d p r es s u res a nd t en s i on s on th es e
reser v o i rs t end s h ere to fill th em a nd there t o s qu eeze ou t

,
thei r c ont en t s i n th a t or the oth er di recti on— p oss ibly a idi ng

t o p rod u ce by u ni on o f s ev era l c a viti es t h os e l a cu nae or
, ,
i rregu l a r c a na l s w hi c h the b ody i n s o m e c a ses p res ent s .
I rregu l a r c a na l s of t hi s ki nd n ot li ned w ith a ny m em,
b ra nes bu t b ei ng s imply ca viti es ru nn i ng th rou gh the fl es h ,
m a i n ly c on s tit u t e the v a s c u l a r s y s t em i n P ol yzoa a n d B ra ch

i op oda a nd s om e M ol l u s ca T h ou gh th e c en t ra l p a rt s o f a
.
v a s cu l a r sy s t em a re ru d ely d ev el op ed yet i ts p eriph er a l p a rts

,
c ons i s t of s i nu ses p erm ea tin g the ti s su es Th e high er ord ers .
o f M ol l us ca h a v e a m ore d ev el op ed s y s t em o f v e s s el s o r
a rt eri es which ru n i n t o th e su b s t a nc e o f th e b ody a n d end

,
i n l a c u nae or s imp l e fi ss u res T hi s endi ng i n l a c u nae t a k es

.
p l a ce a t v a ri ou s di s t a nces f rom the v a s cu l a r cent re I n s om e .
g enera th e a rt er i a l s t ru ctu re i s c a rri ed t o th e p eriph ery of

th e bl ood s y s t em -
whil e i n o th ers it s t op s s h ort mid w a y
,
.
T II E I NNE R T IS S U E S OF A NIMA L S .
3 41
T h rou gh ou t m os t d ers o f th e M ol l us ca the b a ck cu rren t o f

or
bl o od c onti nu es t o be c a rri ed by ch a nnels of the or igin a l

ki n d : th ere a re no t ru e v ei n s bu t the bl ood h a vi ng bee n
,
d eliv ered i nt o th e ti s s u es fi nd s its wa y b a ck t o the p eri

,
vi s cera l ca vity t h rou gh i n os cu l a ti n g s i nu ses A m on g th e .
C ep h a l op od s h owev er t he a ffer en t bl oo d c a n a ls a s we ll a s
, ,
-
,
th e effer en t on es a cqu i r e di s ti n ct w a ll s
, On p u tti n g
.
t ogeth er th es e f a ct s we m a y c onceiv e p retty c l ea rly the

,
s t a ges o f v a s cu l a r d ev el opm ent F rom the origi na l res erv oi r

.
o f nu t ritiv e liq u id b et ween th e a lim en t a ry ca n a l a nd th e
w a ll of the b o dy a p orti on p a rti a lly s hu t off b ecom es a c on

,
t ra ctil e v es sel ; a n d by i t s a cti on s th ere i s p rod u c ed a m ore

ra pid t r a n s fer o f t he nu t ritiv e liqu id t h a n w a s o r igi n a lly
p r od u ce d by th e m oti on s of th e a n i m a l C l ea rly the ext en .

,
s i on o f thi s con t r a c til e t u b e a n d th e d ev el op m en t f r om it o f
b ra n ch es ru nn i ng hith er a nd thit h er i nt o the ti s su es m u s t , ,
by d efi nin g the ch a nn el s Of bl ood th rou gh ou t a p a r t of i ts

c ou r se rend er i t s di s t rib u ti on m ore regu l a r a nd a ctiv e A s
, .
f a s t a s thi s c en t r i fu g a l g rowth a dv a nces s o f a s t a re the effer

,
en t cu rrent s of bl ood p rev ent ed f rom es ca p i ng l a t era lly

, ,
oblig ed t o m ov e f r om th e cen t r e t owa r d s th e ci r c u mferen c e ;
a nd s o f a s t a l s o d oes th e l es s d ev el op e d set Of c h a nnel s be c om e

-
of neces s ity occu pi e d by a f

,
feren t cu rren t s Wh en by a p a ra l
.
,
l el i n cr ea se of defi ni t en es s th e l a cu nae a nd i rre gu l a r s i nu s es

,
th r ou gh whi ch th e a fferen t cu r ren t s p a ss bec ome t ra ns ,
f orm ed i nt o v ei ns th e a ccom p a n yi n g di s a pp ea r a nc e of a l l
,
st a gna nt or s l ow m ovi n g coll ecti on s of bl ood impli es a fu r

-
,
th er imp r ov em en t i n th e c i rcu l a ti on .
By w h a t a gen c y i s effec t ed thi s di fferen ti a ti on Of a d efi n it e

v a s cu l a r s y s t em ? N O s u ffi ci ent reply i s o bvi ou s T he .
genes i s of th e p ri m ordi a l h ea rt i s not c om p reh en s ibl e a s a

re s u lt of di r ec t eq u ilib ra ti on a n d w e c a nn o t rea dily s ee ou r
,
wa y t o i t a s a r esu l t o f i n di rect equ ilib ra ti on ; for it is diffi

c u lt t o i m a gi ne w h a t f a v ou r a bl e v a r i a ti on n a t u ra l s el ec ti on
c ou ld h a v e s ei zed h old of to p rodu ce s u ch a s t ru ct u re A .
c ont ra c til e t u b e th a t a id ed the di s t rib u ti on of nu t ritiv e

3 42 P H Y SI OL OGI CAL D E VE L OP ME NT .
liqu id ha vi ng b een once es t a bli s h ed su rviv a l of the fitt es t

, ,
w ou ld s u ffi ce for i ts g ra d u a l e xt ens i on a n d i t s s u c c es s iv e
m odifi ca ti on s Bu t wh a t were the ea rly st a ges of the c on
.
t ra ct il e t u b e while it wa s yet n ot s u ffici ently form ed t o h elp

,
ci rc u l a ti on a nd whil e it m u s t nev erth el es s h a v e h a d s om e

,
a dv a nt a g e w ith ou t which n o s e l ectiv e p r oc es s cou ld g o on ?
Th e qu es ti on s eem s i ns ol u bl e T o a n oth er p a rt of
.
the q u es ti on h owev er a n a n s w er m a y be v ent u red

, , I f we .
a s k the origin of th es e ra mi fyi n g ch a nnel s whi ch fi rs t ,
a pp ea ri n g as simpl e l a cu nae ev en t u a lly be com e v es sel s ,
h a vi ng d efin it e w a lls a r eply a d m itti ng of c ons id er a bl e

,
j u s tific a ti on is th a t the cu rrent s of nu t ritive liqu id f orced

, ,
a
a nd d r a wn hith er a nd thith er th rou gh t h e ti s s u es th em s elv es ,
i niti a t e th es e c h a nnel s We k now th a t s t rea m s ru nn i ng

.
o v er a nd th r ou gh s olid a n d q u a s i Solid i n org a nic m a tt er -

,
t end to ex ca v a t e d efinit e c ou rs es We s a w rea s on for con .
cl uding th a t the d evel opm ent o f s a p c h a nnel s i n pl a nt s -
c on f orm s t o this genera l p r i nciple Ma y we not then .
s u sp ect th a t the nu t ritiv e liqu id c on t a ined i n the ti ssu e

of a s i m p l e a n i m a l m a d e to ooze n ow i n thi s di recti on a nd
'
n ow i n th a t by th e ch a nges o f p r es s u r e whi c h the a n im a l s

’
m ovem ent s c a u s e com es t o h a ve cert a i n li nes a l ong which it

,
i s th ru s t b a ckw a r d s a nd f orw a rd s m ore th a n a l ong oth er

li nes ; a n d m u s t by repe a t ed p a s s i ngs m a k e th es e m ore a nd
m ore p erm ea bl e u n til th ey b ec om e l a cu nae ? S uch a cti on s
w ill i n evit a bly g o on ; a nd s u ch a cti on s a pp ear c om pet en t to
p rodu ce s om e a t l ea st of th e ob s erve d effects The l ea ding
, ,
.
fa c ts whi ch i n di ca t e th a t this is a p a rt c au se of v a sc u l a r
d ev el opm en t a re th es e .
Gro wths n orm a lly rec u rri ng i n c ert a i n pl a c es a t c ert a in

i nt erv a l s a re a cc omp an i ed by l oca l form a ti ons of bl ood ves
,
-
s els . T he p eri odic m a t u ra ti on o f ova a m ong th e Ma m m a l i a

s u ppli e s a n i n s t a nc e T h r ou gh th e s t rom a of a n o v a ri u m a re
.
di s t ribu t ed i nnu m era bl e m i nu t e v es i cl es whi ch i n th ei r ea rly , ,
st a ges a re mic rosc opic Of th es e s ev era lly c ont a i ned i n thei r

,
.
,
mi nu t e ovi s a cs a ny one m a y d ev el op : the d et ermi ni ng

-
,
T HE INNE R TISS U E S OF A NI MA L S .
3 43
ca u s e b ei ng p rob a bly s om e s light exces s of nu t riti on When .
t he dev e l opm en t i s b ecom i ng ra pid th e c a pill a r i es o f t he

,
n eigh bo u ri n g s t r om a i n c rea s e a n d f o r m a pl ex u s on t h e w a ll s
of th e ovi s a c -
N ow s i n ce th ere i s n o typi ca l di s t rib u t i on of
. .
th e d ev el opi ng ova ; a n d s i nc e th e i nc rea s e of a n ov u m t o

a c erta i n s i ze p r ec ed es t h e i n c rea se o f v a s cu l a r ity r ou n d i t ;
we c a n s c a rcely h elp con c l u di ng th a t th e s etti ng u p of cu r

rent s t ow a r d s t he p o i n t of g r ow t h d et er mi n es th e for m a
ti on of the bl ood v es s el s I t m a y be th a t h a vi ng once c om

-
.
m en c ed thi s l oc a l v a s c u l a r St ru c t u r e c om pl et es it s elf i n a
,
typic a l m a nner ; bu t it s eem s c l ea r th a t thi s g rea t er d ev el op

m en t of bl ood v es s el s a rou nd the g rowi n g ov u m i s i n iti a t ed
-
by the d ra u ght t owa rd s it A b nor m a l g row ths s h ow

.
s till be tt er thi s rel a ti on o f ca u s e a nd effe c t T he fa l se m e m .
b ra nes s om eti m es f ou n d i n th e b ronch i a l t u b es i n i n fl a m

m a t ory di sea ses m a y pe rh a p s fa i rly be h eld a b norm a l i n bu t
,
a p a rti a l sen se : it m a y be s a id th a t t h ei r v a sc u l a r s y s t em s
a re f or m ed a ft er th e typ e o f the m em b ra nes t o w hi c h th ey a re
a ki n. Bu t t hi s ca n sca rc ely be s a id o f t he m orbid g rowt h s

c l a ss ed a s m a lign a n t T h e bl ood v es s el s i n a n enceph a l o id
.
-
c a n c er a re l ed to enl a rg e a n d r a m i fy Oft en t o a n i m m ens e

, ,
e xt ent , by the u nfoldi ng o f t he m orbid m a s s t o whi ch th ey

ca rry bl oo d A li en a s i s th e s t ru ct u re a s a wh ol e t o th e typ e
.
of th e or g a ni s m ; a nd a li en i n g rea t m ea su r e a s i s i t s ti s su e
t o the ti ssu e on whi c h it i s s ea t ed ; it n ev ert h el es s h a pp ens

t h a t the gr owt h of t h e a li en ti s s u e a n d a c c omp a nyi ng a h
s t ra cti on o f m a t eri a l s f rom the bl oo d v es s el s d et erm in e a-
,
c or res p ondi n g g r owt h o f th ese bl o od v es sel s Unl es s th en

-
.
, ,
w e s a y th a t th ere i s a p rovid enti a lly c rea t ed typ e of v a s cu l a r

-
s t ru c t u re fo r ea ch ki n d o f m or bid g r owt h v thi

( a n d e en s
w ou ld not m u ch h elp u s si nce th e v a scu l a r s t ru c t u re ha s

,
n o cons t a n cy w i thi n th e li m it s o f ea ch ki n d
) we a re com ,
p el l ed t o a dmit th a t i n s om e w a y or oth er the cu rren t s o f

bl oo d a re h ere di rectly i n s t ru m ent a l i n form i n g th ei r ow n
c h a nnel s . One m ore pi ec e o f e vid en ce b efore c i ted ,
a s ex emplifyi ng a d a pt a ti on m a y be c a ll ed t o mi nd ;
3 44 P H YSI OL OGI CA L D E V E L OP ME NT .
When a ny m a i n ch a nn el for bl ood l ea di n g t o o r f rom a ,
c ert a i n p a rt o f t h e b o dy h a s b een r en d e red imp ervi ou s

, ,
o th er s a m on g th e c h a nn e l s l ea di n g t o or fr om thi s sa m e p a rt ,
enl a r g e t o t h e e x t ent requ i s i t e fo r f u lfilli ng t h e e x t r a fu n c
ti on t ha t fa ll s u p on th em : the en l a r gem en t b ei ng c a u sed a s ,
w e m u s t i n fer by t he i n cr ea se of the c u rren t s c a rri ed

, .
H ere t h en a re fa ct s w a rra nti ng i n d u ctiv ely th e d ed u ct i on

, ,
a b ov e d r a wn I t i s t ru e th a t we a re l eft i n th e d a rk res p ect i n g

.
t h e c om pl exiti es o f th e p r oc es s H ow th e ch a nn els for bl ood

.
c om e t o h a v e li m iti ng m em b r a n es a n d m a ny of th em m u s
,
c u l a r c oa t s th e h y p ot h es i s d oes n ot h elp u s t o s a y
,
B ut th e
.
e vid ence a s s ig ned g oes fa r t o w a r ra n t th e b eli ef t h a t v a s cu l a r
d ev el opm ent i s i n i ti a t ed by di rec t equ ilib ra t i on ; t h ou gh i n

di r ect equ ilib ra ti on m a y h a v e h a d the l a rger s h a re i n es t a bli s h
i ng th e s t ru c t u res w hi c h di s ti n gu i s h fi ni s h e d v a s c u l a r sy s t em s .
3 01 . O f th e
i nner ti s s u es whi ch r em a i n l et u s next t a k e
b one I n wh a t m a nner i s di fferenti a ted thi s d ens e s u b s t ance
.
s e rvi n g i n m ost ca s es for i nt ern a l s u pp ort ?

Wh en c ons id er i ng th e v ert eb ra t e sk el et on u nd er i ts
m orph ol ogi ca l a s p ec t it wa s p oi nt ed ou t th a t the
f orm a t i on o f d en s e ti s su es i nt erna l a s w ell a s ext erna l i s i n
, , ,
s om e c a s es a t l ea s t b rou ght a b ou t by th e m ec h a ni ca l forc e s

,
t o be res i s t ed T h r ou gh w h a t p r oc ess i t i s b r ou gh t a b ou t we
.
c ou ld n ot th en s t a y t o i nq u i r e : thi s qu es ti on b ei n g not
m orph ol ogic a l bu t phy s i ol ogi ca l A n sw ers t o s om e ki nd red

.
qu es ti on s h a v e s i nce b een a t t em pt ed C e rt a i n a c ti on s t o
.
w hich th e i nt ern a l d en s e ti s su e s of pl a n ts m a y be a s c rib ed ,
h a v e b een i ndic a t ed ; a n d m or e rec ently a na l og ou s a cti ons ,
h a v e b een a s s ign ed a s ca u s es of s om e ext erna l d en s e ti s su es

o f a n im a ls We h a v e now t o a s k wh eth er a cti on s of th e
.
s a m e n a t u re h a v e p r od u c e d th es e i nt erna l d ense ti s s u es o f
a n i m a ls .
T h e p robl e m i s a n i nv olv ed on e B ones h a v e m ore th a n

.
o ne s t a ge T h ey a r e m em b r a nou s or ca rtil a gi nou s b efore th ey

.
b ecom e os s eou s ; a nd th ei r s ucc es siv e c omp onent s u b s t a nces

T HE INNE R T ISS U E S OF A NIMA L S .
345
so fa r di ffer t h a t the effect s of m ech a n i ca l a cti ons u p on the m

di ffer A n d ha vi n g t o d ea l w ith t ra ns i ti ona l s t a t es i n whi ch
.
b on e i s form ed of m ix ed ti s s u es h a vi ng u nlike phys ic a l

,
p r op ert i es a nd u n lik e m i nu t e s t ru ct u res th e effect s o f

,
s t ra i n s b ec om e t oo com pli ca t ed t o fo ll ow w ith p r ec i s i on .
A nyt hi ng i n t he w a y o f i nt erp ret a ti on m u s t th ere f ore be

reg a r d ed a s t en t a tiv e I f a n a ly s i s a n d c o mp a ri s on S h ow th a t
.
t he ph enom en a a re n ot i nc o ns i s t ent with th e hyp oth es i s of

m ech a ni ca l genes i s it i s a s mu ch a s c a n be exp ect ed L et u s
, .
fi rs t ob serv e m ore nea rly th e m ec h a nic a l c on diti on s to which

b ones a re su bj ec t .
T he en d o s k el et on o f a m a m m a l w ith th e m u s cl es a n d
-
lig a m en t s h oldi ng i t t ogeth er m a y be ru d ely c omp a red to a

,
s t ru c tu re b u ilt u p o f s t r u t s a nd ti es ; o f w hich s p ea ki ng,
genera lly th e s t ru t s b ea r th e p r es s u res a nd th e t i es b ea r th e

,
t en s i on s T h e fra m ew ork o f a n ordi na ry i r on r oo f w ill giv e

.
a n id ea o f th e fu nc t i on s o f t h ese t w o el em en t s a n d o f th e
,
m ech a ni ca l ch a ra c t ers r eq u i red by th em S u ch a fra m ew or k

.
c on s i s t s p a rtly Of pi ec e s w hi ch h a v e ea ch t o b ea r a th ru s t i n
th e di rec ti on of i t s l en gth a n d p a rtly of pi ec es whi c h h a v e
,
e a c h t o b ea r a p u ll i n t h e di r ec ti on o f i t s l ength ; a nd t h es e
st ru t s a n d t i es a re di fferently form ed t o a d a pt th em to th es e
di fferent s t ra i n s F u rth er it s h ou ld he rem a rk ed th a t th ou gh
.
,
th e rigidity of th e f r a m ew ork d ep en d s on th e ti es whi ch a re

fl exibl e a s m u ch a s on the s t ru t s whi ch a r e s ti ff yet th e ti es
, ,
h elp t o giv e th e r igidi t y s i m ply by s o h oldin g the s t r u t s i n

p o s iti on th a t th ey ca nn ot es ca p e fr om th e thru sts whi ch fa ll
on th em N ow th e lik e rel a t i on h old s with a di fferen c e
.
a m on g th e b on es a n d m u s c l es : t h e diff eren ce b ei n g th a t h er e
th e t ies a d m it of b e i n g l en gth ened o r s h ort ene d a n d th e

s t ru t s of b e i n g m ov e d a b ou t u p on th e i r j oi nt s T h e m ech a
.
n i ca l r el a ti on s a r e n ot a lt ere d by thi s h ow ev er
,
T he a c ti ons
.
a r e o f e s s en ti a lly th e s a m e ki n d i n a n a ni m a l th a t i s s t a n d
i n g or k eepi n g it sel f i n a s t ra i ne d a ttit u d e a s i n one th a t

, ,
i s c h a n gi ng i ts a ttit u d e —th e s a m e i n s o fa r th a t we h a v e
i n ea ch a set of fl exibl e p a r t s th a t a re p u l l i ng a nd a set of
3 46 P H YS IOL OGICA L D E V E L OP ME NT .
r igid p a rts th a t a re res i s ti ng I t nee ds bu t t o rem ember the

.
s u dd en c o ll a ps e a nd fa ll which t a k e pl a ce wh en th e mu s cl es
‘
a re p a ra ly z e d or t o rem emb er th e i n a bility o f a b a re s k e l et on

,
t o s u pp ort its el f t o s ee th a t th e s t ru t s w i th ou t t he ti es ca n
,
not s u fli c e A nd we h a v e bu t t o thi nk of the f orml es s m a s s

.
i n t o whi c h a m a n wou ld s i nk w h en d ep riv ed of h i s b ones to ,
s ee th a t t h e ti es w ith ou t t h e st ru ts c a nno t s u f fi ce T o t ra ce .
the wa y i n w hich a p a rt ic u l a r b one h a s i ts p a rt i cu l a r th ru s t

t h r own u p on it m a y n ot a l wa y s be p ra cti ca bl e
,
T h ou gh it .
i s ea sy t o p erceiv e h ow a fl exor or ext ens or of the a rm ca u ses

by i ts t en s i on a rea c t iv e p res s u re a l on g the li ne o f th e
.
_
hu m eru s a nd i s ena bl ed t o p rod u ce its effect only by t he

,
r igidi ty of t he h u m eru s ; ye t it i s not s o ea s y t o p erce iv e h ow

’
s uch b on es a s th ose of a h or s e s p elvi s a re si mil a rly a cted
u po n S till a s th e weight o f the hi nd qu a rt ers h a s t o be
.
,
t ra n s ferred f rom the b a ck to t he f eet a n d m u s t be s o t ra ns ,
f err ed th rou gh the b ones it is m a ni fes t th a t th ou gh th es e

,
b ones f orm a v ery c rook ed li ne the weight mu s t p r odu ce a ,
p res s u re a l on g th e a xi s of ea ch : the mu s c l es a nd liga m ents

c oncerne d s er vi ng h ere a s i n oth er ca ses s o t o h old the
, ,
b ones th a t th ey b ea r the p ressu re i ns t ea d of b ei ng di s pl a ced

by it N ot f org etti ng th a t m any p roc es s es of th e b ones h a ve
.
t o be a r t en s i ons w e m a y th en s a y th a t genera lly th ou gh by

, ,
n o m ea ns u n iv ersa lly b on es a re in t ern a l d en s e m a s s es t h a t

,
h a v e t o be a r p res s u res— p r es s u res whi ch i n th e cyli nd ric a l

b on es bec om e l ongit u di na l th ru s t s L ea vi ng ou t e xc epti on a l
.
c a ses l et u s c ons id er b on es a s m a s s es th u s c i r cu m s t a n c e d
, .
Wh en givi ng r ea s on s for th e b eli e f th a t the v ert ebra t e

s k el et on i s m e ch a ni ca lly or igi n a t ed o ne of th e fa ct s u t i n
p,
e vid enc e w a s th a t i n th e v ert eb ra t e s e ri es th e t r a n siti on f ro m

,
th e c a rtil a gi n ou s t o th e os s eou s s pi ne b egi n s p eriph era lly

ea c h v ert eb r a b ei n g a t fi rs t a r i ng o f b one su r
r ou n di n g a m a ss O f c a rtil a g e A nd it w a s p oi n t ed ou t th a t
.
thi s p eriph era l os s ifi ca ti on i s o s s ifi ca ti on a t the regi on o f

grea t es t p res su res N ow it i s not v ert eb rae only th a t foll ow
.
this c ou rs e of d evel opm ent I n a cyli nd ric a l b one th ough

.
,
T HE INNE R T ISS UE S OF A NIMA L S .
347
it diff erently ci rcu m s t a nced the pl a ce s of c om menci ng oss i

is ,
fi ca ti on a re s till the pl a c es on which the s ev eres t s t res s fa ll s .
L e t u s c ons id er h ow s u ch a b one th a t h a s t o b ea r a l ong i tu

di n a l p res su re i s m ech a ni ca lly a ffect ed I f the end .
o f a w a lkin g c a ne be th r u s t w ith f orc e a g a i n s t th e g r ou n d th e

-
,
c a ne b en d s ; a nd p a rti a lly res u mi ng its s t ra ight n es s wh en re

l i eved a g a i n b end s u s u a lly t owa rd s th e s a m e sid e wh en th e
, , ,
th ru s t is renewed A b end s o c a u sed a ct s on the fib res of the

.
c a ne i n nea rly the s a m e wa y a s d oes a b en d c a u s ed by su p

p or ti ng the ca n e h ori zon t a lly a t i ts tw o end s a nd su s p en di ng
a w eight f ro m its middl e I n eith er c a s e th e fib res on th e
.
c onv ex s id e a re ext end ed a nd th e fib res on t he c onca ve sid e

c omp res s ed K ind red a c ti ons oc cu r i n a rod th a t i s s o thick
.
a s n ot t o yi e ld vi s ibly u nd er t he for c e a ppli ed I n the a b s enc e .
of co mpl et e h om og enei t y o f i t s s u b s t a nc e compl et e s ymm e t ry ,
i n i ts f orm a nd a n a pplic a ti on of a force ex a c tly a l on g i ts

,
a xi s th ere mu s t be s om e l a t era l d efl ec ti on ; a nd theref ore

,
s om e di s t ribu ti on of t ens i ons a nd p res su res of the ki n d i ndi

c a ted A n d th en a s th e fa ct whi ch h ere s p eci a lly c oncern s u s
.
, ,
weh a v e t o n ot e th a t th e s t r ong es t t ens i on s a nd p r es s u res a re

b orne by the ou t er l a yers o f fib res N ow the sh a ft of a l ong .
b on e s u bj ect t o m ech a n i c a l a c ti ons of thi s kind s i m il a rly h a s

, ,
its ou t er l a yer m os t s t ra i n ed I n this l a yer th eref or e on the

.
, ,
m ech a ni ca l hyp oth es i s os s ific a ti on sh ou ld c om m enc e a nd h ere

, ,
i t does com m ence — c om m en c es t oo m id w a y be t w een the end s

, , ,
w h ere t h e b end s p rod u c e on the s u p er fici a l p a rt s th ei r m os t

i nt en s e effect s Bu t w e h a v e n ot i n thi s pl a c e s imply
.
t o o b s erv e th a t os s ific a ti on c om m ences a t th e pl a ces o f g rea t es t

s t res s bu t t o a s k wh a t ca u ses it t o d o thi s
, C a n w e t r a c e the .
phys i ca l a cti on s w hi ch s et u p thi s d ep os it Of d en se ti ssu e ? I t

i s I t hi nk p os s ibl e to i n di ca t e a t ru e ca u se th a t i s a t w or k
“
, ,
th ou gh wh eth er it i s a su ffic i en t ca u se m a y be qu es ti one d .
We c oncl u d ed th a t i n cert a i n oth er c a s es th e f orm a ti on o f ,
d ens e tis s u e i ndi rectly resu lt s from th e a lt ern a t e squ eezi ng

a nd rel a x a ti on o f th e v es s el s ru nn i n g th r ou gh the p a rt ; a n d
the i nqu i ry n ow t o be m a d e is w h eth er i n d ev el opi ng b one , , ,

the s a m e a cti ons g o on i n s u ch wa y s a s t o p rod u c e th e ob

s erv e d ef fect s A t th e ou t set we a re m et by wh a t s eem s a
.
fa t a l diffic u l ty— c a rtil a ge i s a n on v a s cu l a r ti s s u e : thi s su b

-
s t a n ce O f whi c h u n os s i fi ed b ones cons i st i s not p erm ea t ed

by mi nu t e c ana l s c a rryi n g nu t ritiv e liqu id a nd c a nn ot , ,
th erefore be a sea t of a cti ons s u ch a s th ose a s s ign ed

,
.
T hi s a pp a rent diffic u lty h owev er fu rni s h es a c on fi rm a ti on

, ,
.
For c a rtil a ge th a t i s wh olly wi th ou t p er m ea ti ng c a na l s d oes

not oss i fy : o s sifi ca ti on t a k es pl a c e on ly a t th ose p a rt s o f
it i nt o w hi ch the ca n a l s p enet ra t e . H en ce w e get a d ,
di ti ona l rea s on for s u s p ec ti ng th a t b one for m a t i on is du e

-
t o th e a ll eged c a u se ; s i n ce it oc c u rs wh ere m ech a ni ca l

s t ra i ns c a n p r od u c e t h e a c ti on s d escr ib ed bu t d oes n ot o ccu r
,
w h ere m ech a n i c a l s t ra i ns c a nn o t p rodu ce th em L et u s .
con s id er m or e c l osely w h a t th e s ev er a l f a c t or s a r e I t will .
s u ffic e for th e a rg u m ent if we c om m ence with th e ext erna l

v a s cu l a r l a y er a s a l rea dy exi s tin g a nd c ons id er wh a t
,
w ill t a k e pl a c e i n it . C a rt il a g e i s el a s tic— i s s om e
wh a t ext en s ibl e a n d s p rea d s ou t l a t era lly u n d er p res su re
, ,
bu t resu m es i t s f or m wh en reli ev ed . H ow th en w ill th e

, ,
m i nu t e c h a nn el s t ra v er s i n g it i n a l l di r ecti on s be a ffec t ed a t
t h e pl a c es wh ere it i s s t ra i ned by a b en d ? T h os e o n t h e
c on v e x s id e w ill be l a t era lly s q u eez ed i n th e s a m e w a y th a t
,
w e s a w th e s a p v es s el s on th e c on v e x S id e o f a b en t b ra n ch
-
a re s q u eez ed ; a n d a s ex u d a ti on o f th e s a p i n t o th e a dj a c en t
p ros en chy m a w ill be c a u sed i n the one ca s e s o i n the oth er , , ,
th er e will be ca u sed exu d a ti on of s eru m i nt o the a dj a cent

c a rt il a ge : ext ra nu t riti on a n d i n crea s e o f s t ren gth res u lti n g
i n b oth c a ses T h e p a r a ll el cea s es h ere h ow ev er
.
,
I n th e .
s h oot o f a p l a nt b ent i n v a ri ou s di rec ti on s by th e w i n d t h e

, ,
S id e whi c h w a s l a t ely c om p r es s ed i s n e w ext en d ed ; a nd
h ence th a t s q u eezin g of th e s a p v es s els whi ch resu l ts from

-
e xt en s i on s u ffi ce s t o feed a n d h a r d en t h e ti s s u e on a l l s id es
,
o f th e s h oot . B ut it i s n ot s o wi t h a b one H a vi ng yi eld ed

.
o n one s id e u n d er l ong it u di n a l p ress u re a n d r esu m ed a s

,
nea rly a s m a y be its p revi ou s sh a p e w h en t he p res s u re is

T HE INNE R T ISS U E S OF A NIMA L S .
3 49
t a k en off th e b one yi eld s a ga i n t owa rds the s a m e sid e wh en

,
a g a i n l ongit u di na lly p r es s ed H ence th e s u b s t a n ce of i t s

.
conc a v e s id e nev er rend er e d c on v e x by a b en d i n th e o pp o

,
s it e di recti on wo u ld not r ec e iv e a ny ex t r a nu t r i t i on did n o

,
o t h er a cti on c o m e i n to pl a y Bu t if we c on s id er h ow i nt er
.
m itt ent p r ess u r es m u s t a ct on ca rtil a g e w e S h a ll s ee t h a t ,
t h ere w ill r es u lt ext r a nu t r iti on of t he conca v e sid e a l s o .
S qu eeze b et ween tw o pi eces o f gl a ss a t hi n bit o f c a ou tch ou c

whi ch ha s a h ol e th rou gh it Whil e the ca ou t ch ou c s p rea d s
.
ou t a w a y fr o m th e c en t re it a l s o s p rea d s i nw a rd s s o a s
, ,
p a rti a lly t o cl os e the h ol e E verywh ere i t s m ol ecu l es m ov e

.
a w a y i n di rec ti on s o f l ea s t res i s t a n c e ; a n d for th os e n ea r
the h ol e t he di rec ti on of l ea s t res ist a nc e i s t ow a r d s the h ol e

, .
L et t hi s h ol e s t a nd for th e t ra n s v er se s ec t i on o f one o f th e
mi nu t e ca na l s or ch a nnel s p a s s ing th r ou gh c a rtil a ge a n d it ,
w ill be m a n if es t th a t on the s id e of th e u n os s i fi ed b on e m a d e
c onca v e i n the wa y d es cribed the com p res s ed c a r til a ge w ill
,
s qu eeze t h e c a na l s t r a v er s i n g it ; a nd i n th e a b senc e o f,
p er fec t h om ogen ei ty i n th e ca rtil a ge th e s qu eeze wi ll ca u s e

,
ex t r a e x u d a ti on from th e c a na l s i nt o th e ca rtil a ge T hu s .
e v ery a dditi on a l s t r a i n w ill giv e t o th e c a rt il a g e it f a ll s u p on ,
a n a dditi on a l s u pply of th e m a t er i a l s for g row th S o th a t .
p resently the s id e w hi ch by yi eldi ng m ore th a n a ny oth er

, ,
p rov es it sel f t o be th e wea k es t will c ea s e t o be th e w ea k es t

, .
Wh a t fu rth er w ill h a pp en ? S om e oth er s id e will yi eld a

littl e— the b end s will t a k e pl a ce i n s om e oth er pl a ne ; a nd
the p or ti ons of ca r til a ge on whi c h rep ea t e d t en s i on s a nd
p res s u res n ow fa ll will be s t rength ened T hu s the r a t e o f .
n u t riti on g r ea t es t a t th e pl a c e w h ere th e b en di n g i s g rea t es t

, ,
a n d ch a ngi ng a s th e i n c id e nc e o f f o rces c h a n g es will b ri ng ,
a b ou t a t e v ery p oi n t a b a l a n ce b et ween th e res i s t a n c es a n d
th e s t r a i ns T hu s t oo t h ere will be d et er mi ned th a t p er i

.
, ,
h
p era l i nd u ra ti on w hi c h w e s ee i n b on es s o c i rcu m st a n c ed .
A s i n a s h oot w e s a w th a t th e w oody d ep os it t a k es pl a c e
t ow a rd s the ou t s id e of the cyli nd er wh ere a cc ordi ng t o the, ,
hyp oth es i s it ou ght t o t a k e pl a ce ; s o h ere we s ee th a t the

, , ,
3 50 P H YSI OL OGICAL D E V E L OP ME NT .
e xc ess of exu d a ti on a nd h ard eni ng occu rri ng wh ere the ,
s t r a i n s a re m os t i n t e n s e w ill f or m a c yli n d er h a vi n g a d en s e
,
ou t s id e a n d a p or ou s o r h oll o w i n s id e T h ese pr o
.
c es ses will be es s en ti a lly the s a m e i n b ones su bj ect t o m ore

c om pl ex m ech a n i ca l a c ti ons s u c h a s s u n d ry of th e fl a t b on es
,
a nd o th er s th a t s erv e a s i n t ern a l f u lc ra B e t h e s t ra i n s.
t ra n s v ers e or l ongitu di na l be th ey t or si on st ra i ns or mixed

,
s t r a i n s t h e ou t er p a rt s of th e b one will be m ore a f fec ted by

'
th em th a n i ts i nner p a rt s T h ey will theref ore t en d ev ery

.
wh ere t o p r od u ce res i s ti ng m a sses h a vi ng ou t er p a r ts m ore

d ense th a n thei r i nner p a rt s A nd by c a u s i n g m os t g row th
.
w h ere th ey a re m os t i nt ens e th ey will c a ll ou t rea ctiv e forc es

,
a d eq u a t e t o b a l a nce th em T h er e a re d ou btl es s
.
o b s t a cl es i n th e wa y o f thi s i n t er p ret a ti on I t m a y be sa id .
th a t th e f orces a cti ng on th e ou t er l a y ers i n the m a nn er

d es crib ed wou ld c om p res s the c a na ls t oo littl e t o p rod u ce
,
th e a ll eged effect s ; a n d if ev enly dist rib u t ed a l ong th e wh ol e

'
l engths of the l a yers they wou ld p rob a bly do s o Bu t i t

, .
n eed s only t o be nd a fl exibl e m a s s a nd ob s er v e t h e t end ency
t o f or m c rea s es on th e c onc a v e s u r fa ce t o f eel a s s u red th a t

,
a l on g t he s u r f a c e of a n o ssifyi n g b one th e yi eldi n g o f t he ,
ti s su e wh en b ent w ill not be u nif orm I n the a b sen ce of .
c ompl et e h om ogeneity th e i n t ers titi a l yi eldi ng will t a k e

,
pl a ce a t s om e p oi nt s m or e th a n oth ers a n d a t one p o int ,
a b ov e a l l o th e r s .Wh en a t the wea k es t p oi nt— the cent re

,
o f comm en c i n g o ss ifi ca t i on— a n e xt ra a m ou nt o f d ep o s it h a s
b een c a u s ed it w ill c ea s e t o be th e wea k es t ; a nd a dj a cent

,
p oi nt s now the wea k es t will b ecom e th e pl a ces of yi eldi ng

, ,
a nd i nd u ra ti on . I t m a y be fu rt h er obj ec t ed th a t the hyp o

th esis i s i nc omp a tibl e with th e p ers i s t ence O f c a rtil a ge for s o
l ong a tim e b et ween th e epiphy s i s of b ones a nd th e b ony
m a s s es whi ch th ey t ermi na t e B ut th ere is the reply th a t .
th e pl a c es o cc u pi e d by thi s c a rtil a ge b ei ng pl a c es a t which

t he b one l en gth en s th e n on oss i fi ca ti on i s i n p a rt a pp a rent
,
-
o nly— i t i s ra th er th a t n ew c a rtil a g e is f orm ed a s f a s t a s t he
e—existi ng c a r til a g e os s i fies a n d th ere is th e f u rt h er reply

p r
T HE INNE R T ISS U E S OF A NI MA L S .
351
th a t th e l
s own es s Of lti m a t e os s ific a ti on of thi s p a rt i s
the u ,
du e t o i t s non v a scu l a ri ty a n d t o m ech a ni ca l c on diti ons

-
,
whi c h a re u n fa v ou ra bl e t o i ts a cqu i rem e n t of v a s c u l a rity .
Once m ore th ere i s th e d em u rrer th a t i n th e epiphy s es os s i fi

,
c a ti on d oes n ot b e gin a t t he s u r fa ce bu t w i t hin th e m a ss o f
the c a rtil a ge E xpl a n a ti on of thi s i m pli es a bility t o f o ll ow

.
ou t th e m ech a n i c a l a cti on s i n a r es ili en t s u b s t a n c e w hi ch lik e ,
i ndi a ru bbe r a d m it s of bei n g di s t or t ed i n a l l w a y s by p ress u re

-
,
a nd recov e r i ng i t s f o r m a nd it s ee m s imp oss ibl e t o s a y h ow

,
t he m ore s u p erfi c i a l a nd m ore d ee p s ea t e d c a n a l s t ra v ers i n g

-
it will be res pectiv ely a ffect ed .
Of c ou rs e it i s n ot m ea nt th a t thi s osseou s d ev el op m ent

by di rect eq u ilib ra ti on t a k es pl a ce i n th e i n divid u a l T h ou gh .
it i s a c or oll a ry f rom the a rgu m ent th a t i n ea ch i ndivid u a l

t he p r oc ess m u s t be furth ered a nd m odifi e d by the p a r tic ul a r
a c ti on s t o w hi ch th e p a rt i cu l a r b one s a r e e xp os ed ; et t h e
y
l ea di ng t ra it s of s t ru ct u re a s su m ed by the b ones a re a s s u m ed
i n c on f or mity w ith t he i nh eri ted typ e T hi s h owev er i s n o.
, ,
di ffi cu lty T he type it sel f i s t o be reg a rd ed a s the a ccu m u

.
l ated res u lt of s u ch m odifi ca ti on s t ra ns m itt ed a n d i ncrea s ed

,
f rom g enera ti on t o genera ti on T he a c ti on s a b ov e d es cribe d

.
a s t a ki n g pl a c e i n th e b on e o f a n i n divid u a l m u s t be u n d er ,
s t oo d a s p r o d u ci ng th ei r t ot a l e f fec t li tt l e by li t tl e i n th e
c orres p onding b ones O f a l ong s er i es of in divid u a l s E v en i f .
bu t a s m a ll m odifi ca ti on ca n be s o wrou ght i n the i ndividu a l ,
yet i f s u c h m o difi ca ti on or a p a rt o f it i s i nh er it a bl e w e
, , ,
m a y r ea dily un d er st a n d h ow i n the cou rse of geol o gic ep och s

, ,
th e ob s erv ed s tr u ct u res m a y a ri s e i n th e a s s ign e d w a y .
H ere m a y fit l y be a dd ed a s t r ong c onfi rm a ti on I f we fi nd .
c a ses wh ere i ndivid u a l b ones s u bj ect i n exc epti ona l d egr ees
,
t o th e a cti on s d e s crib ed p resen t i n exc epti ona l a m ou nts the

,
m odifi ca ti on s a tt rib u t e d t o th em w e a r e gr ea tly h elp e d i n

,
u n d er s t a n di ng h ow th ere m a y be p rod u c e d i n th e ra c e th a t
a gg re g a t e of m odific a ti on s whi ch the hyp o th es i s impli es .
S u ch c a ses occ u r i n r ick etty child ren I a m i n d ebt ed t o Mr

. .
B u s k for p oi nti n g ou t th es e a b nor m a l f orm a ti ons of d ens e

ti s s u e th a t a re n ot a pp a rently e xplic a bl e a s res u lt s o f
,
3 52 P H YSI OL OGI CA L D E V E L OP ME NT .
m ec h a ni ca l a cti on s a n d re—a cti on s I t wa s only on t r a ci n g

.
ou t t h e p r oces s e s h er e a t w or k t h a t th er e su gg es t ed it sel f t h e
,
s p ec ific i nt er p r et a ti on of t h e no r m a l p r oc ess a s a b ov e set ,
fort h . Wh en from c on s ti tu ti on a l d efect b ones do

, ,
n ot os s ify w ith du e ra pidity a n d a re m ea nwhil e s u bj ec t t o

,
the o rdi n a ry s t r a i ns th ey b ec om e di s t ort ed

,
Rem em b er i ng .
h ow a m a s s wh i c h h a s b een m a d e t o yi eld i n a ny di r ecti on

by a force it c a nnot w ith s t a nd i s s om e littl e ti m e be fore i t
,
r ec ov er s c ompl etely i t s p revi ou s f orm a nd u s u a lly i n d eed

, , ,
u n d er g oes w h a t i s c a ll e d a p erm a n en t set ; it i s i n fera bl e

th a t w h en a b one i s r ep ea t e dly ben t a t th e s a m e ti m e th a t
t he liqu id c on t a i ned i n i t s c a n a l s i s p oor i n th e m a t er i a l s
for f orm i n g d en se t i s su e th ere w ill not t a k e pl a ce a p r op or
,
t i ona te s t ren gth en i n g Of th e p a rt s m os t s t ra i n ed ; a n d th es e

p a rt s w ill give wa y . T hi s h a pp ens i n r i ck et s B u t thi s .
h a vi ng h a pp ened th ere g oes on w h a t i n t el eol ogi ca l l a ng ua ge

, , ,
we c a ll a rem edi a l p r oc es s S u pp osi n g th e b one t o be one

.
com m on ly a f fect ed— a fem u r ; a nd s u pp os i n g a p erm a nen t

b en d t o h a v e b een c a u s ed i n it by th e weigh t o f th e b ody ;
the s u b sequ ent resu lt i s a n u nu su a l d ep os iti on o f c a rtil a gi n
ou s a nd o s s e ou s m a tt er on th e con ca v e S id e o f th e b o ne If .
t he b one i s rep res en t ed by a s t ru n g bow t h en th e d ep os it ,
o cc u rs a t th e p a rt r ep re s en t ed by th e s p a ce b etw een th e bow
a n d t he s t r i n g . A nd t hu s occu rri n g wh ere i t s re s i s ta n ce i s

m os t effect iv e it i ncrea ses u ntil the a pp r oxi m a t ely s t ra ight
,
-
pi ece of b one form ed withi n the a rc h a s b ec om e s t rong enou gh

,
t o b ea r t he p res su re w ith ou t a pp r eci a bly yi eldi ng N ow .
thi s di rect a d a pt a ti on s ee m i n g s o lik e a s p ec i a l p rovi s i on

, ,
a n d f u rn i s hi n g s o rem a rk a bl e a n i n s t a nc e of w h a t i n m edi ca l ,
'
bu t u n s ci en tific l a n gu a g e i s c a ll e d th e vi s m edi ca tri a: n a tu rae

, ,
i s s i m ply a resu lt of th e a b ov e d es c rib e d m ech a ni ca l a cti on s

-
a n d re a cti o n s ,
goi n g on u n d er th e ex cepti ona l conditi on s .
E a ch ti m e su ch a b en t b on e i s s u bj ec t t o a f orc e whi c h a g a in
b end s it the sev eres t c omp r es s i on fa lls on th e s u b s t a nce of
,
i ts c onc a v e s id e .E a ch tim e th en the c a n a ls ru nn i ng

, ,
th r ou gh this p a rt of i ts s u b s t anc e a re vi ol ently s qu eezed

T HE IN N E R T ISS U E S OF A NIMA L S .
3 53
fa r m ore s qu eez ed th a n th ey or a ny oth er of the c a na l s

w ou ld h a v e b ee n h a d t h e b on e rem a i ne d s t ra ight
,
H en ce .
,
o n e v ery r ep eti t i o n o f th e s t r a i n th ese c a na l s ne a r t h e c on ,
c a v e s u r fa ce h a v e th e i r cont en t s f orc e d ou t i n m or e th a n
n or m a l a b u nd a nc e T he m a t er i a l s for the f o rm a ti on o f ti ss u e
.
a r e su ppli e d i n q u a nt ity g rea t er t h a n ca n be a ss imil a t e d by
t he ti ssu e a l rea dy f or m ed ; a nd from th e ex ces s o f exu d ed

pl a s m a new ti s su e a ri s es
,
*
A l a y er of org a ni z a bl e m a t eri a l
.
a c c u m u l a te s b etween the c onca v e s u rf a c e a nd the p eri

os t eu m ; i n t hi s a c c or di n g t o the or din a r y c ou rs e of ti s s u e
,
gr ow th new v es s el s a pp ea r ; a n d th e a dd ed l a y er p resently
,
a s s u m es t h e hi s t o l o gi c a l ch a ra ct er o f t h e l a y er f r om whi c h
it ha s g r own Wh a t next h a pp ens ? T hi s a dde d l a yer
.
,
f u rth er f ro m th e neu t ra l a xi s th a n th a t whi c h ha s th rown

it ou t i s now th e m os t sev erely c om p res sed a n d i ts v es s els
, ,
a r e t h e m os t s e v erely s qu eezed The pl a c e of gre a t es t exu da.
ti on a nd m os t r a pid d ep os it of m a tt er i s th eref ore t ra n s ferred ,
t o t hi s new l a y er ; a nd a t the s a m e ti m e th a t a c tiv e n u t riti on

i n crea s es its d ens ity th e ex ces s of org a ni za bl e m a t eri a l
,
f orm s a noth er l a yer ext ern a l t o it : the s u cces s iv e l a yers s o

a dd ed enc r o a chi ng on t h e s p a c e b et ween t h e c on c a v e s u r fa ce
,
o f t h e b on e a n d t h e ch or d o f i t s a re Wh a t li m its .
t he encroa ch m ent on thi s s p a ce — wh a t st op s th e p r o c es s of

fi lli n g it u p ? T h e a ns wer t o thi s qu es ti on w ill be m a ni fes t
wh en Ob servi n g th a t th ere c om es i n t o pl a y a c a u s e w hi c h
g ra d u a lly di m i ni s h es the f orces fa lling on ea ch n ew l a y er .
F or the t ra ns v ers e s ec ti on a l a rea i s s t ep by s t ep i nc rea se d ;

a n d a n i n crea s e of t h e a rea ov er w hi c h th e weight b orn e is
di s t r ib u t ed impli es a rel a tiv ely s m a ll er p res su re u p on ea ch

,
p a rt of it F u rth er a s the t ra ns vers e di m en si ons of the b one

.
,
i nc rea s e the m a t eri a ls c om p os i n g i ts c onvex a n d con ca v e

,
l a y ers b ec om i n g fu rther f rom the neu t ra l a xi s b ec ome b ett er

, ,
To th is i m p l i ed i nfere nce i t i s j
ob ected th at e x c es s of nut ri t i ve m a te
ri a l does not nec e s s a r i l y l e a d to corr e spon di ngl y i nc rea s ed

growth .My re pl y
”
i s th a t a conco m i ta nt fa ct or i s a c ti v i ty of th e ti ss ue , a nd th a t i n i ts a b s ence
gro wth i s not t o be e x pec ted .
69
3 54 P H Y SIOL OGI CAL D E VE L OP ME NT .
pl a c ed for res i s ti ng the st ra in s t o be b orne S o th a t b oth by .
t he i nc rea s ed qu a ntity of d ens e m a tt er a nd by its m ech a ni ca lly

m ore a dv a n t a geou s p os iti on th e bending s of the b one a re
-
,
p r og res s iv ely d ecrea s ed Bu t a s th ey a re d ec rea sed ea c h

.
,
n ew l a y er f or m e d on t h e conc a v e s u rfa c e h a s i t s s u b s t a n ce
a nd i t s v es s el s l es s c o m p res s e d ; a n d th e res u l t i ng g rowt h
a n d i n d u r a ti on a re ren d ere d l e s s ra pid E vid ently then the .

, ,
a dditi on s sl ow ly dimi ni s hi n g w ill ev en t u a lly c ea se ; a nd t hi s

, ,
will h a pp en w h en t he b one n o l ong er be nd s T h a t i s to s a y .

,
t h e thi ck eni ng O f th e b one will rea ch i t s li m it wh en th ere is

equ ilib r i u m b et w een th e i nc id en t f orc es a nd t h e f or c es whi ch
r es i s t th em H ere in d eed w e m a y t ra c e wi t h grea t cl ea rnes s

.
, ,
th e p r oc es s Of di rect equ ilib ra ti on— m a y see h ow a n u nu su a l

f orce f a lli ng on the m ovi ng equ ilib ri u m of a n org a nism a nd
,
n ot o v erth row i n g it g oes on w o rkin g m odific a ti ons un til t he

,
r e a cti on b a l a nc es th e a cti on .
T h a t h owev er whi c h n ow chi efly c onc ern s u s is t o n ot e

, , ,
h ow thi s m a rk ed a d a pt a ti on su pp ort s the gener a l a rgu m en t .
Un qu es ti on a bly b one i s i n this c a s e f or m ed u n d er th e i nfl u

enc e o f m ech a n ic a l st re ss a nd f or m ed j u s t w h ere it m os t
,
eff ect u a lly m eets t he s t r ess T his resu lt n ot oth erwi s e .

,
expl a i ned i s expl a i ne d by the hyp oth esi s a b ov e s et f orth

,
.
.
A n d w h en we see th a t this s p eci a l d ep os it Of b one i s s e

c ou nt ed for by a cti ons lik e th os e t o which b one f orm a ti on in -
g enera l i s a sc r ib ed th e p r ob a bility th a t th es e a re the a cti ons

,
*
a t w ork b ec om e s v ery grea t .
In r e cen t
yea rs (s ince 1 8 9 0) P rof . Wil hel m Rou x , in e ss ay s on func
ti onal ada pt a ti on, h a s set forth s om e v i ews aki n t o th e foregoi ng i n re s pec t
t o th e g ene ra l b el i ef th ey i m pl y, th ou gh di fi eri ng i n re s pe ct of the physi o

l ogi ca l p roce s s e s he i ndi c a t e s The fol l owi ng rcl ev a nt p ass age h a s b een
.
t ra ns l a te d for m from a n a rti c

e l e of hi s i n the Rea l E nc yc l op ed i c der g e
s um m ten Hei l ku de
n — “
A m ore com pl ete theory of fun cti onal a da p ta t i on
b v th e a ut hor is founded on th e pt i on th a t th e functi onal
a s su m
‘ ’
s t i m ul u s ,
or
‘
th e a ct of e x erci s i ng th e functi on ’ (i n m us cl es a nd gl a nds ) , a nd es pe
c i al l y, i n th e c a s e of bones th e c oncus s i on a nd tens i on ca u s e d by s t re s s a nd
s tra i n, e x ert a
‘
t rop i c s ti m ul u s on th e c el l s , i n cons eq uence of wh i ch, a nd
h ’
a l ong wi th an i ncrea s e d a bs orpt i on of n ut ri m ent , th ey grow a nd e v entu a l l y

i ncrea se ( or t he os te obl a s t s at the p oi nt of
grea ter s ti m ul us form m ore bone) ;
wh il e , c onv ersel y, w i th conti nued i na cti v i ty, by a bs ence of th es e s ti m ul i th e
THE IN N E R T IS S U E S OF A NIM A L S .
355
Of c ou r s e it is not a ll eged t h a t
st ru ctu res a ris e in s
os eou s
this wa y a l one The b ones of the

. a nd v a ri ou s d erm a l
s ku ll
b ones c a nnot be thu s i nt erp ret ed H ere the na tu ra l s el ec .
ti on of f a v ou ra bl e v a r i a ti ons a pp ea r s the onl y a s s ign a ble

c a u s e— the eq u ilib ra ti on i s indi rect We kn ow th a t os si fi c .
d ep os it s now a nd th en occu r i n ti ssu es wh ere th ey a re not

u s u a lly fou n d ; a n d s u c h d ep os it s origi n a lly a b nor m a l if
, ,
th ey occu rr ed i n pl a ces wh ere a dv a nt a ges a ros e f rom th em ,
might rea dily be es t a bli s h e d a nd i ncrea se d by s u rviv a l o f the

fi tt es t E s p eci a lly m ight w e exp ec t thi s t o h a pp en wh en a
.
c on s tit u ti ona l t end ency to f orm b one h a d b een es t a bli s h ed by

a c t i ons o f t h e kin d d es c ribe d ; for it i s a f a mili a r f a ct th a t
di fi erent i a ted typ es of ti s su e h a vi ng o nc e b ec o m e el em ent s

,
o f a n or g a n i s m a re a pt occ a s i on a lly t o a r i s e i n u nu s u a l
,
pl a c es a nd there to r ep ea t a l l thei r p ec u li a r hist ol ogic a l

,
ch a ra ct er s A nd thi s m a y p os sibly be the rea s on why the

.
b on es o f the s ku ll th ou gh n ot exp osed t o force s s u ch a s

,
th os e which p ro d u c e i n oth er b ones d ens e ou t er l a yer s i n

, ,
c l u di n g l es s d en s e i n t eri or s n e v erth el es s rep ea t thi s g ener a l

,
t rai t of b ony s t ru ctu re Whil e h owev er it i s b ey on d d ou bt

.
, ,
t h a t s om e b on es a re not du e t o th e di rec t i n fl u enc e o f

m ec h a ni ca l s t res s w e m a y I t hi nk c on cl u d e th a t m ech a nic a l
, , ,
st ress i niti a t es b one f orm a ti on -

.
§ 3 02 Wh a t i s th e origi n of nerv e ? I n wh a t w ay do its

.
p rop erti es s t a nd r el a t ed t o th e p r op ert i es of th a t p rot opl a sm

w h en ce th e ti s su es i n g ener a l a ri se ? a n d i n wh a t w a y i s it
di fferenti a ted fr om p rot opl a s m s i m u lt a neou s ly wi th the oth er
ti s su es ? T h ese a r e p r ofou n dly i nt eres ti ng q u es ti ons ; but
qu es ti on s t o whi ch p os itiv e a n s wers ca nn ot be e xp ect ed .
A l l th a t ca n be d one is t o i n di ca t e a ns wer s which s ee m

f ea s ibl e .
T h a t th e p rop erty s p eci a lly d i s pl a y ed by nerv e i s a p ro ,
n ou ri s h m e nt o f th e cel l decl i ne s so th a t th e w a s te i s i ns u ffi ci e ntl y rep l a ce d ( or

o th er wi s e th a t th e b o ne -s ub s ta nc e g ra d ua l l y l o s e s i t s power of res i s ta nce to
t h e os teobl a s t s form ed a s a re s ul t of i na cti i ty v
356 P HYS I OL OGI CAL D E V E L OP ME NT .
p erty whi ch p r ot opl a s m p os ses s es i n a l ow er deg ree i s m ani ,

-
w
fes t . T h e s a rco d e o f a Rh i z op o d a n d th e s u b st a n ce of a n
u n i m p r eg n a t e d ov u m exhibit m ov em en t s th a t i m ply a p ro
,
a a t i on o f s ti m u l u s fr o m on e p a r t o f th e m a s s t o a n ot h er
p g .
We h a v e n ot fa r t o s eek for a p rob a bl e origi n of thi s p h en o

m en on T h ere i s go o d r ea son for a scribi n g it t o the ext reme
.
i ns t a bility of the org a n i c c oll oid s of whi ch p rot opl a s m con

s i st s . T h es e i n c omm on w ith c oll oid s i n genera l a s s u m e dif
, ,
ferent i s om eric form s w i th g rea t f a cility ; a nd th ey di s pl a y n ot

s i m ply i s om eri s m bu t p olym eri s m F u rth er thi s rea di nes s
.
,
t o u nd erg o m ol ecu l a r re a rra ng em ent h a bit u a lly s h ow s it s elf

-
,
i n c oll oid s by th e r a pid p rop a ga ti on of t he r e a rr a ng em ent -
from p a rt t o p a rt A s P rof Gr a h a m h a s s h own m a tt er i n

. .
,
thi s s t a t e oft en p ec ti zes a lm os t i n s t a nt a neou s ly— a t ou c h

w ill t ra n s f orm a n en ti re m a s s T h a t i s t o s a y th e ch a nge of
.
,
m ol ecu l a r s t a t e once s et u p a t on e end s p rea d s t o th e oth er ,
end— th er e i s a p r og res s o f a s ti m u l u s t o c h a n g e ; a n d thi s i s
wh a t we see i n a n erv e S o m u ch b ei ng u n d ers t ood l et u s

.
,
re s t a t e th e c a s e m or e c ompl e t ely
-
.
Mol ecu l a r ch a n ge implyi n g a s it d oes m oti on o f m ol ecu l es

, ,
c om mu ni ca t es m oti on t o a dj a cent m ol ecu l es ; be th ey of the

s a m e ki n d or of a differen t ki n d I f th e a dj a cen t m ol ec u l es
.
,
e i t h er o f th e s a m e ki n d or of a di ffer en t ki n d be s t a bl e i n ,
c om p os iti on a t em p or a ry i ncrea se o f o s c ill a ti on i n th em a s

,
w h ol es or i n th ei r p a r t s m a y be th e only resu lt ; bu t if th ey
, ,
a re u n s t a bl e th ere a r e a pt t o a ri se c h a n g e s o f a rr a ngem en t
a m on g th em or a m ong th ei r p a r ts o f m ore or l ess p erm a n ent

, ,
ki nd s . E s p eci a lly i s thi s s o with t he c ompl ex m ol ec u l es

whi ch f or m c oll oid a l m a tt er a n d w ith the org a nic c oll oid s
,
a b ov e a l l H e n ce it is t o be i n ferred t h a t a m ol ecu l a r di s
.
t u rba n ce i n a ny p a rt o f a livi n g a n i m a l s et u p by eith er a n ,
e xt ern a l o r i nt ern a l a g en c y w ill a lm os t cer t a i nly di s t u rb a nd

,
c h a n ge som e o f th e s u r rou ndi n g c oll oids n ot origi na lly i m
plic a t ed— w ill di ffu s e a w a v e o f ch a nge t ow a rd s o th er p a rt s

o f th e or g a n i s m : a w a v e whi c h will i n th e a b s en c e o f er
p ,
f eet h om ogeneity t ra v el fu rth er i n s om e di rec ti on s th a n i n

,
T HE I NNE R TISS U E S OF A NIMA L S .
3 57
others . L et us ask ne xt wh a t
will d et erm i ne the
di fferenc es of di s t a n ce t ra v ell e d i n diff eren t di rec ti on s Ob .
vi ou s l y a ny m ol ec u l a r a git a ti on s p rea di n g f ro m a c en t re w ill ,
g o f u r th es t a l on g ro u t es t h a t o ffer l ea s t res i s t a nce W h at .
rou t e s w ill th es e be ? T h ose a l ong whi ch t h ere l i e m os t m ol e

cu l es th a t a re ea s ily c h a ng e d by t h e diff u s ed m ol ec u l a r m oti on ,
a nd w hich yet d o n ot t a k e u p m u ch m o l ecu l ar m oti on i n a s
s u m i n g th ei r new s t a t es M ol ecu l es whi ch a r e t ol era bly s t a bl e

.
wi ll not rea dily p rop a g a t e the a git a ti on ; for th ey will a b s orb

it i n the i nc rea s e of th ei r own os cill a t i on s i n s t ea d of p a s s i n g ,
it on Mol ecu l es w hich a r e u ns t a bl e bu t whi ch i n a s s u mi n g

.
,
is om eric f orms a b sorb m ot i on w ill not rea dily p rop a g a t e it ;

, ,
si nce it wi ll di s a pp ea r i n w orki ng the ch a ng es i n th em Bu t .
un s t a bl e m ol ec u l es w hi c h i n b e i ng i s o m er i c a lly t ra n s f or m e d
, ,
d o not a b s or b m oti on a nd s till m ore t h ose whi c h i n b ei ng

, ,
s o t ra n s form e d giv e ou t m oti on will r ea dily p r op a g a t e a ny

, ,
m ol ecu l a r a git a ti on ; s i n ce th ey w ill p a s s on the i m p u l s e eith er

u n dimi ni s h e d or i nc rea s e d t o a dj a c ent m ol ec u l es
, , If .
th en we a s su m e a s we a re n ot only wa rra nt ed i n d oi ng bu t
,
a r e o blig e d t o d o th a t p rot opl a s m c ont a i ns tw o or m ore

,
c o ll oid s either mi ngl ed or fee bly c om bi n ed ( s i nce it can not

,
c ons i s t of si m pl e a lb u m en or fib ri n or c a s ei n or a ny a lli e d ,
p roxim a t e p ri n cipl e ) ; it m a y be concl u d ed t h a t a ny m ol e

cu l a r a git a ti on s et u p by wh a t w e c a ll a s tim u l u s w ill diffu s e ,
it s el f further a l on g s om e li nes th a n a l ong oth er s i f the com ,
p o n e n t s o f t h e p r ot opl a s m a r e not q u it e h om o g en eou s ly di s
p ers ed a nd if s om e o f th e m a r e i s om eric a lly t r a n sform e d

,
m ore ea s ily or with l es s exp en di ture of m oti on th a n

, ,
o th er s ; a n d it w ill es p eci a lly t r a v e l a l on g s p a c es occu pi ed
chi efly by th os e m ol ecu l es whi ch giv e ou t m ol ecu l a r m o

ti on d uri ng th ei r m et a m orph oses if th ere s h ou ld be a ny ,
s u ch . Bu t n ow l et u s a s k wh a t s t ru ct u r a l effec t s
will be w rou ght a l on g a t ra ct t ra v ers e d by thi s w a v e of
m ol ecu l ar di s t u rb a nc e A s is sh own by th os e t ra ns for m a
.
ti on s whi ch so ra pidly p rop a ga t e th em s elv es thr ou gh c oll oid s ,
m ol ec u l es th a t h a ve un d erg one a c ert a in c h a nge of f orm ,

a re a pt to c om m u ni ca t e a like ch a nge of f orm to a d

j a c ent m ol ecu les of the s a m e ki nd— th e i m p a ct of ea ch
ov er th row i s p a s s ed o n a nd p ro d u ce s a n oth er o v erth row .
P rob a bly th e p r onenes s t owa rd s i soch roni s m of m ol ecu l a r

m ov em ents nec ess it a t es thi s . I f a ny m ol ecu l e h a s h a d
i ts c omp onent s re—a rr a n ged a nd thei r os cill a ti on s cons e
,
qu ently a lt ered there resu lt m ov em ent s not c oncord a nt with

,
t h e m ov em ents i n a dj a c ent unt ra n s form ed m ol ecu l es bu t ,
which imp ressi ng them s elv es on the p a rts of s u ch untra ns

,
f or m ed m ol ecu l es t end t o genera t e i n th em c oncor d a n t m ov e

,
m ents— t end th a t is t o p rod u c e the re a rra ngem ent s i nv olved

, ,
-
by th es e c onc or d a nt m ov em ent s I s t hi s a cti on li m it ed t o

.
s t rictly i so m er ic s u b s t a n ces ? or m a y it ext en d t o s u b s t a nc e s
th a t a re cl os ely a lli ed ? I f a l on g w ith the m ol ecu l es of a

c om p ou nd c oll oid th ere a re mi ngle d th os e of s om e ki nd red
c oll oid ; or if with the m ol ecu l es of thi s c om p ou nd c oll oid
th ere a re mi ngl ed the c omp onent s ou t of whi ch oth er su ch
m ol ecu l es m a y be f orm ed ; th en th ere a ri s es th e qu es ti on
d oes the s a m e i nfl uenc e which t end s t o p rop a ga t e the i so
m eri c t ra ns f orm a ti ons t end a l s o t o f orm new m ol ecu l es of
,
the s a m e ki n d ou t of the a dj a c en t c om p onent s ? T h er e is

r ea s on t o s u s p ect th a t it d oes .A l rea dy wh en t rea ti ng of th e
nu t r iti on o f p a rt s it wa s p oint ed ou t th a t w e a re
o blig ed t o rec o gn i z e a p ow er p os s ess e d by ea ch ti ss u e t o b u ild
u p ou t of the m a t er i a l s b r ou ght t o it m o l ec u l es o f th e s a m e
, ,
typ e a s th os e of whi c h it i s f orm ed T hi s bu ildi n g u p of like

.
m ol ecu l es s eem s expli ca bl e a s ca u s ed by the t end ency of the

n ew c om p onen ts which th e bl oo d s u ppli es t o a cq u i re m ov e
,
m ent s i s och ronou s with th os e of the lik e c omp onen t s i n the

ti s su e ; whi ch th ey c a n do only by u niti ng i nt o lik e com
p ou n d m ol ecu l es N ec es s a rily th ey mu s t g ra vit a t e t owa rds
.
t t q ilib i m ch t t f q ilib i — vi g
a s a e of e u r u ; s u s a e o e u r u m m o n
q
e u ilib ri u m o f c ou r se — m u s t be one i n whi ch th ey os cill a t e
i n th e s a m e tim es with n eighb ou ri n g m ol ecu l es ; a nd s o
t o os cill a t e th ey m u s t f a l l i nt o g r ou ps id entic a l with the
g rou ps a rou nd th em . I f thi s be a genera l p rincipl e of
T HE I NNE R TISS U E S OF A NI MA L S .
3 59
ti s s ue gr owt h a n d rep a i r we m a y c oncl u de t h a t i t w ill a pply

-
,
i n the ca se b efore u s . A w a v e o f m ol ecu l a r di s t u rb a nce

p a ss i ng a l ong a t ra ct of mi n gl ed c oll oid s c l os ely a lli ed i n c om
p os i t i on a nd is om eri ca lly t ra ns formi n g the m ol ecu l es o f one
,
o f th em will be a pt a t the s a me tim e t o form s om e new m ol e

,
c u l es of th e s a m e typ e a t a ny pl a c e w h ere th er e e xi s t t h e
,
p r oxim a t e c omp onent s either u ncom bi ned or f eebly c om bi n ed

,
i n s om e not v ery different w a y A nd thi s wi ll be m ost lik ely

.
t o occu r wh ere t he m o l ec u l es of th e coll o id th a t a re u n d er

g oi ng the is om eri c ch a nge p red omi na t e bu t h a v e sca tt ered
, ,
th rou gh them the oth er m ol ecu l es ou t o f whi ch th ey m a y be

form ed eith er by c om p os i ti on or m o difi ca ti on
,
T h a t i s to .
s a y, a w a v e o f m ol ec u l a r di s t u r b a nc e diffu s e d fr o m a c en t r e ,
a nd t r a v e lli n g f u rth es t a l ong a li ne w h ere l i e m o s t m ol ecu l es
th a t can be i s om er i ca lly t ra ns for m ed with f a cility w ill be ,
lik ely a t th e s a m e ti m e to fu rther di fferenti a t e th i s li ne a n d ,
m a k e it m ore c h a ra c t eri ze d th a n b efore by th e ea s y t ra n s - ~
f or m a bility of i ts m ol ecu l es . One a dditi on a l s t ep ,
a nd t he i nt erp r et a ti on i s rea ch e d A n a l ogy s h ows it t o be

.
not i m p rob a bl e t h a t th ese org a n i c c o ll oid s i s om eric a lly t ra ns

,
f orm ed by slight m ol ecu l a r i m p a ct or i nc rea s e o f m ol ecu l a r

m oti on w ill s om e of th em res u m e th ei r p revi ou s m ol ecu l a r
,
s t ru c t u r es a ft er th e di s t u r b a n c e h a s p a s s ed We k n ow th a t
.
wh a t a re s t a bl e m ol ec u l a r a rra ngem ent s u n d er one d eg ree of

m ol ecu l a r a gi t a ti on a re not s t a bl e u nd er a n oth er d e gree ; a nd
,
th ere i s evid ence th a t re—a rra ng em ents of a n i ncon s pi cu ou s

ki nd a re occa s i ona lly b rou ght a b ou t by v ery s light ch a nges
o f m ol ec u l a r a git a ti on Wa t er su ppli e s a cl e a r c a s e P r of
.
. .
Gra h a m i n fer s th a t w a t er u nd er g oe s a m o l ec u l a r r e a rr a n ge -
°
m ent a t a b ou t 3 2 — th a t i ce h a s a c o ll oid f or m a s well a s a
c ry s t a ll oid form ,d ep end en t on t em p era t u re S en d th rou gh .
i t a n ext ra wa v e of the m ol ecu l a r a gi ta ti on w e ca ll h ea t a n d ,
i ts m ol ec u l es a ggreg a t e i n one w a y L et the w a v e di e a wa y

.
,
a nd i t s m ol ecu l es r esu m e th e i r p r evi ou s m o d e o f a g r e g a ti on

g .
A n d obvi ou s ly s u ch t ra ns form a ti on s m a y be rep ea t e d b a c k

w a r ds and f orw a rd s withi n na rrow limit s of t emp era tu re .
3 60 P H Y SIOL OGI CA L D E V E L OP ME NT .
N ow a m ong the ext rem ely u ns t a bl e org a n i c c oll oids s u ch a ,
ph enom en on i s fa r m ore likely t o h a pp en S u pp ose then th a t .

, ,
t he ner v e c oll oid i s one of w hi ch t he m ol ecu l es a re c h a nged in

-
f or m by a p a s s i ng wa v e of ext ra a git a ti on bu t r esu me th ei r ,
p revi ou s form wh en the wa v e h a s p a s s ed : the p revi ou s f or m

b ei n g t he m os t s t a bl e u n d er the c on diti on s which th en recu r .
Wh a t f oll ows ? I t foll ows th a t thes e m ol ecu l es will be r ea dy

a g a i n t o u n d e r g o i s om er i c t ra n s f or m a ti on w h en th ere a g a i n
occ u r s th e s t im u l u s ; w ill a s b efor e p r op a g a t e t he t ra ns form a

, ,
ti on m os t a l on g t he t ra ct where s u c h m ol ecu l es a r e m os t
a b u nd a n t ; w ill a s b efore t en d t o f or m new m ol e c u l es o f th ei r
, ,
o wn typ e ; w ill a s b efor e m a k e t he li ne a l on g wh i c h t h ey l i e

, ,
on e of ea s i e r t ra n s f er for th e m ol ec u l a r a git a ti on E v ery .
rep etiti on will h elp t o i n c rea s e t o i nt eg r a t e t o d efi ne m ore

, ,
c om pl et ely the cou rs e o f th e es ca pi ng m ol ecu l a r m oti on

,
e xt e n di n g i ts rem o t er p a r t w hil e it m a k es i ts nea rer p a rt
m ore p erm ea bl e— w ill h elp th a t i s t o f or m a li ne of di s

, ,
ch a rge a li ne for c on d u cti ng imp res s i on s a nerv e

, , .
S u ch s ee ms t o m e a n ot u n fa i r s eri e s o f d ed u c ti on s f ro m
th e kn own h a bit u d es of c oll oids i n gener a l a n d th e org a nic
c oll oids i n p a rti cu l a r A nd I thi nk th a t th e impli ed na t u re
.
a n d p r op erti es o f ner v e c orres p on d b ett er w ith th e ob s er v e d
ph enom ena th a n d o the na tu re a nd p rop erti es impli ed by

.
o th e r hyp o th es e s Of cou rs e th e sp ecu l a ti on a s it h ere st a nds

.
i s bu t t en t a tiv e a nd l ea v es mu c h u nexpl a i n ed
,
I t giv es n o .
o bvi ou s reply t o the q u es ti on s— wh a t c a u ses t he f orm a ti on of
n er v e s i n di r ec ti on s a d a pt ed t o th e need s ? w h a t d et er m i nes
th ei r a pp rop ri a t e conn exi ons — q u e s t i ons h owever to whi ch , , ,
w h en we c om e t o d ea l w ith phy s i ol ogi ca l i nt eg ra ti on we m a y ,
fi n d n ot u n sa ti s fa c t ory a ns wer s M or eov er it s a y s n ot hi ng

.
a b ou t t h e g enes i s o f g a ngli a A g a ngli on it i s c l ea r m u s t

.
, ,
c ons i s t of a coll oid a l m a tt er equ a lly u ns t a bl e or s till m ore ,
u n s t a bl e
,
whi ch w h en di s tu rb ed fa ll s i n t o s om e di fferent
, ,
m ol ecu l a r a rra ngem ent p erh a p s ch emi c a lly s impl er a nd giv es

, ,
ou t i n s o d oi n g a l a rg e a m ou nt o f m ol ec u l a r m o ti on— s e r v e s
a s a res erv oi r o f m o l ecu l a r m oti on which m a y be s u dd enly

T HE I NNE R T IS S UE S OF A NI MA L S .
3 61
di sch a rge d a onl g

ff erent nerv e or nerv es wh en ex c it e
an e ,
men t of a n a fferent n erv e ha s di s eng a ged it H ow s u ch a .
s t ru ct u re a s thi s r es u lts th e hyp oth es i s d oes not s h ow B u t

,
.
a dm i t ti ng th es e s h ort co mi n gs it m a y still be h eld th a t we
a re i n th e w a y p oin t ed ou t ena bl e d t o f orm s om e id ea of the

, ,
a cti on s by whi c h n er v ou s tiss u e is di fferen ti a t ed .
3 03 .p ec u l a ti on a ki n t o a nd c onti nu ou s with the l a st

A s , , ,
i s s u gges t e d by a n i nqu i ry i nt o the origi n o f m u scu l a r ti s s u e .
C ont ra c tility a s well a s i rrit a bility i s a p rop erty o f p ro t opl a s m

o r s a r c od e ; a n d a s b efor e s u gg es t ed
,
is not i m p rob a bly
du e t o i som eri c c h a n ge i n one or m ore of i t s c om p onen t c ol
l oid s I t i s a fea s ibl e s u pp ositi on th a t of th e s ev era l i s om eri c
.
ch a n ges s imu l ta neou s ly set u p a m on g t h es e c om p onen t col

l oid s s om e m a y be a cc om p a ni ed by ch a nge of b u lk a n d s om e
,
n ot . C l ea rly the i so m eri c ch a nge u n d erg one by th e c oll oid

whi c h we s u pp os e t o f or m nerv e m u s t be one n ot a cc omp a n i e d,
by a pp rec i a bl e ch a nge of b u lk ; s i n c e ch a ng e of b u lk impli es

”
“
i nt ern a l work a s phy s i c i s ts t er m it a nd th eref ore ex
, ,
pen di tu re o f f orce C on v ersely th e c oll oid ou t o f which

.
,
mu s cl e origi n a t es m a y be one th a t rea dily p a ss es i nt o a n

,
i s om eri c s ta t e i n whi c h it oc cu pi es l es s s p a c e : th e m ol ecu l a r

distu rb an ce c a u s i ng thi s c ont r a cti on be i n g c ommu nic a t ed t o
it f rom a dj a cent p orti on s of nerv e s u b s t a nce th a t a re m ol e -
cu l a r l y di s t u r b ed ; or b ei n g oth erw i s e c o m m u n i c a t ed t o it
by di rect m ech a nic a l or ch em i ca l s ti m u li : a s h a pp ens wh ere

n erv es do n ot exi s t or wh e r e th ei r i n fl u en c e h a s b een cu t
,
of f
. T hi s i nt er p ret a ti on s eem s i nd ee d t o be di rec tly a t , ,
v a ri a nce w i th t he fa ct th a t m u s cl e d oes n ot di m i ni sh i n bu lk
d u ri n g c on t ra cti on bu t m erely ch a n ges i ts s h a p e T h a t w hi ch .
we see t a k e pl a ce w it h th e m u s cl e a s a wh ol e is s a id a l s o t o ,
t a k e pl a ce w ith ea ch fi bre — whil e it s h ort en s it a l s o b roa d en s .
T h er e i s h ow ev er a p os s ibl e s ol u ti on of thi s di ffi cu l t y

, A .
c ont r a cti ng coll oid yi eld s u p i t s w a t er ; a n d the contra ct ed

c o ll oid
p l u s t h e free w a t er m a y h a v e th e s a m e b u lk a s b efor e
,
th ou gh the c oll oid ha s l ess I f it he r epli ed th a t i n thi s

.
3 62 P H Y S I OL OGICAL D E VE L OP ME NT .
c a se th e
wa t er s hou ld b ecom e visibl e b et ween th e s u b s t a nce
o f th e fib re a n d its s a r c ol em m a or s h ea th it m a b e re j oi n e d
y,
th a t thi s i s not neces s a ry— i t m a y be d ep os it ed i nters titi a lly .
P os sibly the st ri a t ed s t ru ct u re is on e th a t f a c ilit a t es i ts

e x u d a ti on a n d s u b s equ en t r e—a b s or p t i on ; a nd t o this m a
y be
du e th e s u p er i ority of s t r i a t ed mu s c l e i n ra pidity of c ont ra e
ti on . Gran ti n g th e sp ecu l a t iv e ch a ra c t er of t hi s
i n t erp ret a ti on l et u s s ee h ow fa r it a g rees with th e fa ct s I f
, .
t he a cti on s a re a s h ere s u pp os ed th e c ont ra ct e d or m ore i nt e

,
gr a t ed s t a t e of the mu s cu l a r c oll oid w ill be th a t whi ch it

t en ds c onti nu a lly t o a s s u m e —th a t i n t o which it h a s a n i n
c rea s i n g a ptit u d e t o p a ss wh en a r tifici a l p a r a ly s i s h a s b e en
p rodu c ed a s s h ow n by D r N orris— th a t i nt o which it l a p s es

, .
c ompl et ely I n ri g or m or ti s T he sens ibl e m oti on gen era t ed

.
by the c on t ra cti on ca n a ri s e on ly f ro m th e t ra ns form a ti on

o f i nsen sibl e m oti on T hi s i ns ens ibl e m oti on su dd enly
.
yi eld ed u p by a cont ra cti ng m a s s impli es the fa ll of i ts

,
c omp onent m ol ecu l es i nt o m ore s t a bl e a rra ngem ent s A nd .
there ca n be no s u ch fa ll u n l es s th e p revi ou s a rra ngement i s

-
u n s t a bl e . F rom thi s p oi nt of Vi ew t oo it i s p os s ibl e

, ,
t o s ee h ow th e hyd ro c a rb ons a nd c a rb o hyd ra tes con su m ed

- -
i n mu scu l a r a c ti on m a y p r od u c e th e i r e ffect s
,
F or th e s e
.
no n n it r og en ou s el em ent s o f food
-
w h en con s u m ed i n the
,
ti s su es giv e ou t l a rge a m ou nt s of m ol ecu l a r m oti on T h ey

,
.
do this i n p resenc e o f th e m u s cu l a r c oll oid s whi ch h a v e l os t

m ol ecu l a r m oti on d u r i ng th ei r f a ll i n the s t a bl e or c on t ra ct ed
st a t e F rom the m ol ecu l a r m oti on th ey giv e ou t m a y be
.
,
r es t ored th e m ol ecu l a r m oti on l os t by th e c on t r a ct e d c oll oid s ;
a n d th e se c ont ra c t ed coll o ids m a y thu s h a v e th ei r m ol ecu l es
ra i s e d t o th a t u n s t a bl e s t a t e f ro m which a g a i n f a lli ng th ey
, ,
c a n a g a i n g enera t e m ec h a n ic a l m oti on .
T his c onc epti on of th e na t u re a n d m od e o f a cti on o f m u s

cl e w hil e it is s u gg es t e d by k nown p rop er ti es of c oll oid a l
,
m a t t er a nd c onform s to the r ec ent c oncl u s i ons of org a nic

ch emi s t ry a nd m ol ecul a r physics es t a bli s h es a com p reh en s ibl e
,
rel a ti on b et ween the vit a l a cti ons of the l ower a nd th e high er

T HE I NNE R T IS S U E S OF A NIMA L S .
3 63
an im a ls I f w e c ont em pl a t e th e m ovem ents of cili a of a

.
,
’
Rhi zop od s p seu d op o di a of a P olyp e s b ody or o f th e l ong
’
, ,
p end a nt t ent a c l es of a M edus a we s h a ll s ee g rea t con gru ity ,
be t wee n th em a nd thi s hyp othes i s Bea r i n g i n mi n d th a t th e .
c ont ra c til e s u b s t a nce of d evel op e d m u s cl e i s a ff ect e d not by

nerv ou s in fl u en c e only bu t wh ere nerv ou s i n fl u ence i s de
, ,
s troy e d i s m a d e t o c ont r a ct by m ec h a n i ca l di s t u r b a nc e a nd
,
ch emi ca l a cti on we m a y i n fer t h a t it d oes not di fl er i ntri n

,
s i c a l l y f rom th e p r i m ordi a l c ont r a c til e su b s t a n c e w hi ch in ,
th e l owes t a ni m a l s ch a nges i ts b u lk u n d er o th er s ti m u li th a n
,
th e nerv ou s We sh a ll s ee s ign ific a nce i n th e fa ct a s oer

.
t a i ned by D r Ra ns om th a t v a ri ou s a gent s which e xcit e

.
,
a n d a r res t ner v e m u s c u l a r m ov em ent s i n d e v el op ed a nim a l s

-
,
e xcit e a n d a rres t t h e p rot opl a s m i c m o v e m en t s i n ova We .
s h a ll u n d er s t a n d h ow tis su es n ot yet di fferent i a t ed i n t o m u s
c l e a nd n e r v e h a v e this j oi n t i rri t a bili ty a n d con t r a ctility ;

,
how m u sc l e a n d ner v e m a y a ri se by th e s eg reg a ti on of th ei r

mi ngl ed c oll oid s the one of whi ch not a pp reci a bly a lt eri ng
, ,
i ts b u lk d u ri ng i s om eric ch a nge re a dily p rop a g a t es m ol ecu l a r

,
di s t u rb a nc e whil e the o th er c on t ra cti ng w h en i s om eri ca lly

, ,
ch a nged l es s r ea dily p a s s es on the m ol ec u l a r di s t u rb a nce ;

,
a n d h ow by thi s di f
,
feren ti a ti on a nd i nt eg r a ti on of the con
d u cti ng a nd the c on t ra cti ng c oll oid s th e one ra m ifyi ng ,
th r ou gh the oth er it b ec om es p os s ibl e for a w h ol e m a ss t o

,
c ontra ct su dd enly i n s t ea d of c ont ra cti ng g ra du a lly a s it d oes

, ,
wh en u ndi fferenti a t ed .
The qu es ti on r em a i ni n g t o be a s k e d i s— Wh a t c a u s e s the
s p eci a li z a t i on o f c on t ra c til e s u b s t a n c e ? —Wh a t ca u ses th e
g row th of c oll oid m a s s es which m onop oli ze thi s c on t ra c tility ,
a n d l ea v e ki n d red coll oid s t o m on op oli z e o th er p ro p e rti e s ?
H a s n a t u r a l sel e cti on g ra d u a lly l oc a li ze d a n d i nc r ea s ed th e

p ri m ordi a l mu s c u l a r s u b s t a n ce ? or ha s th e frequ ent rec u r
ren ce of i r rit a ti on s a nd con s eq u en t c on t r a c ti on s a t p a r tic u l a r
p a rt s d on e it ? We h a v e I thi nk r ea s on t o c oncl u d e th a t
, ,
di rect eq u ilib r a ti on ra th er th a n i ndi rect equ ilib r a ti on h a s b ee n

chi efly op er a tiv e The rea s oni ng th a t wa s u s ed i n th e c a s e
.
3 64 P H Y SIOL OGI CAL D E VE L OP ME NT .
o f ner e a v ppli es eq u a lly i n th e c a se of mu s cl e A p orti on .
o f u ndi fferen t i a t e d ti ss u e c ont a i ni ng a p r e d o mi na n c e of th e
c oll oid th a t c on t ra ct s i n ch a n gi ng will d u ri ng ea ch ch ange

, , ,
t en d t o f orm new m ol ec u l es of its own typ e from the oth er

c o ll o id s di ffu s e d th r ou gh it : th e t en d ency of th es e ent a n gl e d
c oll oids t o fa ll i nt o u n ity with th os e a r ou n d th em will be ,
a id e d by ev ery s h ock o f i s om eri c t r a n s form a ti on H en c e r e

.
,
e a t ed con t r a c ti on s w ill fu rth er th e g r ow th o f t h e c ont ra c ti ng

p
m a ss a nd a dv a nce i ts di fferenti a ti on a nd i nt eg ra ti on
, If .
,
t oo we r em em b er th a t t h e m u s c u l a r c oll oid i s m a d e t o con

,
t ra ct by m ech a nic a l di s tu rb a n ce a nd t h a t a m on g m ech a nic a l

,
di s t u rb a nces one whi ch w ill m os t rea dily a ffect it s i m u l ta

n eou s l y t h r ou gh ou t i t s m a ss i s ca u s ed by s t ret chi n g w e s h a l l ,
be con sid er a bly h elp ed t owa r d s u nd er s t a ndi n g h ow th e c on

t ra ctil e ti ss u es a re d ev el op ed I f ext en s i on of a m u s cu l a r
.
c oll oid p revi ou s ly a t res t p r od u ces i n it th a t m ol ecu l a r di s

,
tu rba n ce which l ea d s t o i s om eri c c h a ng e a n d d ecrea s e of

b u lk th en th ere i s no di ffi cu lty i n expl a i ning the m ov em ent s
,
of c ili a ; t h e for m a ti on o f a con t ra ct il e l a y er i n t h e v a s cu l a r
s y s t em b ec om es c omp reh en s ibl e ; ea ch dil a t a ti on o f a bl oo d
v es s el c au s ed by a gu s h o f bl oo d w ill be foll owed by a con

,
s t r i c ti on ; th e h ea rt w ill p u l s a t e vi ol ently i n p r op or ti on a s
it i s vi ol ently di s t end ed ; a rt eri es w ill d ev el op i n p ower a s

t h e s t res s u p on th e m b ec om es gr ea t er ; a n d we sh a ll s i m i
l a rl y h a v e a n e xpl a na t i on o f th e i nc rea sed m u scu l a rity o f
th e a li m ent a r y ca n a l which is b rou ght a b ou t by i nc rea s e d
di s t en s i on of it .
T h a t th e p rod u c ti on o f c ont ra c til e ti ssu e i n cert a i n l oca li

ti es i s du e t o the m ore f req uen t excit em ent i n th ose l oca liti es
,
o f th e c on t r a c tility p o s s es s e d by u n di f feren ti a t ed ti s s u e i n
g enera l is a vi ew h a rm on i zi n g w ith t ra it s w hi ch th e di ffe
,
r en ti a t ed c on t r a ctil e ti s s u e e xhibits T h ese a re th e rel a

.
ti on s b et w een m u s cu l a r ex erci se m u scu l a r p ower a n d m u s

, ,
c u l a r st ru c t u re ; a n d it i s t h e m or e needf u l for u s h ere t o
n oti c e th em b eca u s e of c er t a i n a n om a li es th ey p resent ,
whi ch a t fi rs t sight s eem i nc onsi s t ent with the beli ef th a t

, ,
T HE INN E R T IS S UE S OF A NIM A L S .
3 65
t he f u ncti on a lly d eterm i ned m odifi ca t i on s of mu s c l e a re i n

-
h eri t a bl e .
M u s c l es di sa g ree g rea tly i n t h ei r ti n t s : a l l gra d a ti on s

'
between w h i te a n d d eep red bei ng o b serv a bl e C on t ra s t s a re .
vi s ibl e b etween th e m u scl es of di fferent a ni m a l s b etw een the ,
m u sc l es o f t h e s a m e a n im a l a t di ffe ren t a g es a n d bet wee n ,
di fferent mu s cl es o f th e s a m e a nim a l a t the s a m e a ge .
We w ill gl a nce a t the fa ct s u nd er th es e h ea d s : n oti ng u n d er

e a c h o f t h e m t he conn exi o n w hi c h h ere c hi efly con c ern s
u s— t h a t b et ween t he a ctivity of m u scl e a nd i ts d epth

o f c o l ou r . The c old bl oo d ed Ver tebr a ta a re t a k en
-
,
a s a gr ou p di s ti ng ui s h ed f ro m th e w a r m bl ood ed by the
,
-
w hi ten es s o f th ei r fl es h ; a nd th ey a re a l s o di s t i n gu i s h ed by
thei r c om p a ra tiv e i nert nes s T h ou gh a fi sh or a rep til e ca n
.
e x ert con s id era bl e force for a s h o rt tim e it i s not ca p a bl e ,
o f p r ol o ng ed ex er ti on Bi r ds a n d m a m m a l s s h ow grea t er
.
e nd u ra n c e a l ong w i t h th e d a r k er c o l ou red m u s c l es I f a m on g
-
.
bi r d s them s elv es or m a mm a l s th em s elv es we m a k e com p a ri

s on s ,
w e m eet with ki nd red con t ra s t s— e s p ec i a lly b et ween
Wild a n d d om es tic crea t u res o f a lli ed ki n d s Ba rn d oor f owls .
-
a re light er fl esh ed th a n m o s t u nt a m e d g a lli n a c eou s bi r d s ;

-
a n d a m on g th es e l a s t th e ph ea s a n t m o vi ng a b ou t bu t littl e , ,
i s light er fl esh ed th a n th e p a r t ridge a n d th e gr ou s e w hich a re

-
m ore nom a dic T he m u s cl es o f th e s h ee p a re n ot on t he

.
a v er a g e s o d a r k a s th os e o f th e d eer ; a nd it i s s a id th a t
the fl es h of th e w ild b oa r i s d a r k er th a n th a t o f the pig

-
.
P erh a p s h ow ev er t h e c on t ra s t b et w ee n the h a r e a nd the

, ,
r a bbit a f fo r d s a m on g fa m ili a r a nim a l s th e be s t ex a mpl e of

, ,
the a ll eg ed r el a ti on : th e d a r k fl esh ed h a r e h a vi ng n o ret rea t

-
a n d m a ki n g w id e e x cu r s i on s whil e t h e w hit e fl es h ed ra bbit

,
-
,
p a s s i ng a gr ea t p a rt of i ts ti m e i n i ts b u rrow r a rely wa nd er s ,
fa r fr om h om e The p a r a ll el con t r a s t b et w een y ou ng

.
a n d ol d a ni m a l s h a s a p a ra ll el m ea ni ng Vea l i s m u ch .
w h it er th a n b ee f a n d l a m b is of light er c o l ou r th a n m u tt on
, .
T h ou gh a t fi rs t s ight th ese f a c t s m a y n ot s eem t o fu rni s h

confi rm a t ory e vid en c e s i nc e l a mb s i n th ei r pl a y a pp ea r t o
,
3 66 P H YS IOL OGI CA L D E VE L OP ME NT .
e xp end m or e m u scu l a r f orce th a n th ei r s ed a t e d a ms ; yet the

m ea n i ng of th e con t r a s t i s rea lly a s a ll eg ed F or i n con s e .
qu ence of t he l a w t h a t t he s t ra i ns whi c h a n i m a l s h a v e t o
.
o v er c om e increa se a s th e cu b es of the di m ens i ons w hi l e

, ,
t h ei r p ow ers of ov erc om i n g th em i n crea s e only a s t h e s q u a res
(g t h e m ov em en t s o f a n a d u lt a ni m a l cos t m u c h m ore
i n m u s cu l a r effort th a n do th ose o f a y ou n g a ni m a l : th e
r es u lt b ei n g th a t t h e s h ee p a nd t h e c ow ex erc i se t h e i r m u s cl es
m ore vig orou s ly i n th ei r qu i et m ov em en ts th a n the l a mb ,
a n d t h e ca l f i n th e i r liv ely m ov em ent s I t m a y be a dd ed a s .
s ign ifi c a nt th a t th e d om es ti c a ni m a l i n whi c h n o v ery m a rk ed

,
d a rk en i ng o f the fl es h t a k es pl a ce a l ong w ith i ncrea s i n g

a g e n a m ely the pig i s one w hi c h or di n a rily k ept i n a sty
, , , ,
l ea d s s o qu i escen t a li fe th a t the a s s igned c a u se o f d a rkenin g

d oes not com e i nt o a cti on B u t p erh a p s the m os t
.
c onc l u s iv e e vid en ce s a r e t h e c on t r a s t s w hi c h e xi s t b etw een
th e a ctiv e a n d i na c tiv e m u s c l e s of th e s a m e a n i m a l Bet w ee n .
t he l eg m u s c l es o f f ow l s a n d th ei r p ec t or a l mu sc l es t h e di ffer
-
,
enc e of c o l ou r i s f a m ili a r ; a n d w e k n ow th a t fowl s ex er ci se
th ei r l eg m u s cl es m u ch m ore th a n the mu scl es which m ov e

-
th ei r wi ng s S i m il a rly i n th e t u rk ey i n th e gu i nea f owl i n

.
, ,
t h e ph ea sa nt A nd t h en a ddi ng m u c h t o th e f or c e of thi s
.
,
e vid en c e w e see th a t i n p a r t ridges a n d g rou se whi c h be l on g

, ,
t o th e s a m e or d er a s ou r d om es ti c f ow l s bu t u s e th ei r wi n g s
a s c on s t a n tly a s th ei r l eg s littl e or no diff erence is visibl e

,
bet ween th e col ou r o f th es e two g rou p s o f m u scl e s S p eci a l .
con t r a s t s lik e th es e do n ot h ow ev er exh a u s t th e p roo f s ; for

, ,
th ere i s a s till m ore signific a n t genera l c on t ra s t T he .
m u s cl e of the h ea rt whi c h is the m os t a ctiv e of a l l mu scl es

, ,
i s th e d a rk est of a l l m u s cl es .
T h e c onnexi on of ph en om ena thu s sh own i n s o m a ny w a y s ,
impli es th a t the bu lk of a m u s cl e i s by no m ea ns th e s ol e
m ea s u re o f th e q u a ntity o f f orc e it ca n ev olv e I t w ou ld seem .
th a t oth er thi ng s eq u a l th e d epth of c ol ou r v a ri es w ith the

, ,
c ons t a ncy of a cti on ; whil e o th er thi ng s equ a l the b u lk v a ri es

, ,
with the a m ou nt of f orc e th a t h a s t o be pu t f orth u pon oc ca

TH E INNE R T IS S UE S OF A NIMA L S .
3 67
si on. T h es e of c ou rs e a re a pp roxi m a t e rel a ti ons More .
c orrectly we m a y s a y th a t the a c ti ons of p a le mu s cl es a re

e ith er r el a tiv ely feebl e th ou gh f requ en t ( a s i n th e m a s s iv e
fl a nk s of a fi s h ) or rel a tively i nfrequ en t th ou gh s t rong ( a s i n

,
t he p ec t ora l m u s c l es of a com m on fow l ) whi l e th e a c ti on s of

d a rk m u s cl es a re b oth f requ ent a nd s t rong S om e su ch dif .
ferent i a ti on m a y be a n ti c ip a t e d by i nf er en c e f rom th e res pec

tiv e phy s i ol o gi ca l req u i rem ent s A mu s cl e whi c h ha s u p on
.
occ a s i on t o e v olv e c ons id era bl e f or c e bu t which h a s th erea ft er

,
a l o ng p er i o d o f res t d u r i ng w hi c h r ep a i r m a y r es t ore it t o
ef fic i ency req u i res ne ith er a l a r ge res erv e o f t he con t ra ctil e

,
s u b s t a nce th a t i s i n s om e w a y d e t eri ora t ed by a cti on nor ,
highly d ev el op e d a ppli a nc es for b ri ngin g it nu t r itiv e m a t e

-
ri a l s a n d r em ovi n g e ffet e p ro d u ct s Wh ere c ont ra r i wis e a n

.
, ,
ex er t ed m u s cl e whi c h h a s u n d er g one m u c h m ol ecu l a r ch a n ge
i n ev olvi ng m u c h m ec h a n i c a l force h a s s oon a ga i n t o ev olv e

,
m u ch m ech a ni c a l f orc e a n d s o on c onti nu a lly ; it i s cl ea r th a t

,
e ith er t he qu a ntity of c on t r a c til e su b s t a nce p r es en t m u s t be
grea t or th e a pp a ra t u s for nu t riti on a nd d ep u ra ti on m u s t

,
be v ery effici en t or b oth ,

H ence w e m a y l ook for m ar k e d
.
u nl ik eness es o f mi nu t e s t ru ct u r e b et ween m u scl es which a re
m a rk edly cont ra s t ed i n a c tivity A nd w e m a y s u sp ect th a t

.
th es e c ons picu ou s c ont ra s t s of c ol our b et ween a ctiv e a nd

i na ctiv e mu scl es a re du e t o thes e impli ed di fferenc es of
' '
mi nu t e s t ru ct u re : p a rtly di fferenc es be tween th e nu mb er s

o f bl ood v es s els a nd p a rtly di f
-
ferenc es b et ween th e qu a ntiti es
or qu a liti es of s a r c ou s m a tt er .
H ere th en we h a v e a key t o th e a pp a ren t a n om a ly a b ov e

, ,
hi n t ed a t— th e m a i nt ena nce of bu lk by cert a i n mu s cl es which

h a ve b een rend ered c om p ara tiv ely i na c tive by ch a ng ed h a bit s
o f li fe . T h a t th e p ec t or a l m u s cl es of th os e d om es tic bi r d s
.
w hi ch fly bu t littl e h a v e n ot d wi n dl e d t o a ny gr ea t ext ent

, ,
h a s b een th ou ght a f a c t a t v a r i a nc e with the c oncl u s i on th a t

fu ncti ona lly p rod u c ed a d a pt a ti on s a re i nh erit a bl e
-
I t ha s .
b een a r gu ed th a t if p a rt s whi ch a re ex ercis ed i nc rea s e n ot ,
only i n th e i n divid u a l bu t i n th e ra c e whil e p a rts which

‘
,
3 68 P H Y SIOL OGI CA L D E V E L OP ME NT .
b ec om e l ess a ct iv e d ecrea s e ; t hen a not a bl e di fferenc e of si ze

s h o u ld exi s t be t ween t h e m u sc l es u s e d for fligh t i n bi rd s th a t
fly m u ch a nd th os e i n bi rd s of a n a lli ed ki nd t h a t fly li ttl e
,
.
B u t a s we h ere s ee thi s is n ot t h e t ru e i m pli ca t i on

, ,
T he .
c h a ng e i n s u ch c a ses m u s t be chi efly i n v a s c u l a r ity a nd a b u n
d a n ce of con tra c til e s u b s t a nce ; a nd ca nnot be t o a ny g rea t ,
e x t en t i n b u lk
,
F or a bi r d t o fly a t a l l i t s p ect ora l m u s c l es
.
, ,
b ones of a tt a chm en t a n d a l l a cc om p a nyi ng a ppli a nces mu s t

, ,
be k ept u p t o a c ert a i n l ev e l o f p ow er I f th e p a rt s d wi n dl e
.
m u ch the c rea t u re w ill be u na bl e t o li ft it s el f from the

,
gr ou nd Bea ri n g i n mi n d th a t the f orce whi ch a bi rd ex

.
p ends t o su s t a i n it s elf i n the a i r du ri ng ea ch s u cces siv e i ns t a nt

o f a s h ort flight i s o th er thi ng s eq u a l
,
the s a m e a s it ex
,
p en d s i n ea c h s u c ces siv e i ns t a nt of a l ong flight w e s h a ll s ee ,
t h a t th e m u s cl es em pl oy ed i n th e tw o ca s es m u s t h a v e s om e
thi ng lik e eq u a l i n t en s iti es of cont ra ctil e p ower ; a nd t h a t the
s t ru ct u ra l di f
ference s b et w een th em m u s t h a v e rel a ti on m a i nly
t o th e l en gth s of ti m e d u ri n g whi ch th ey ca n c on ti nu e t o re
p ea t c ont ra cti ons of like i nt ens ity T h a t i s t o s a y w hil e the
.
,
p ow er of fligh t i s ret a i ned a t a l l th e m u s cl es a n d b ones c a n

,
n ot g rea t ly d wi n dl e ; bu t th e d w i n dli n g i n bi rd s w h ose flights

,
a re s h ort o r i nfrequ ent o r b oth w ill be i n t he r es er v e st oc k

,
o f th e s u b s t a n ce th a t i s i n ca p a c it a t e d by a cti on or i n th e ,
a ppli a nc es t h a t k eep t h e a pp a ra t u s i n r ep a i r or i n b oth , .
Only w h ere a s i n the s t ru thi ou s bi r d s th e h a bit o f flight i s

, ,
l os t c a n we exp ect a t r ophy of a l l th e p a rts c oncerned i n

,
flight ; a n d h ere we fi nd it .
A re s u ch di fferenti a t i on s a m on g th e m u s cl es f u nc t i on a lly
p rod u ced ? or a re th ey p rod u ced by th e na t u ra l s el ecti on of
v a ri a ti ons di s ti n gu i sh ed a s s p ont a n eou s ? We h a v e I thi nk , ,
g ood g rou nd s for c oncl u di ng th a t th ey a re f u ncti on a lly pro

d u c ed . We k n ow th a t i n i ndivid u a l m en a n d a n i m a l s the ,
p ower of s u s t a i ned a cti on i n m u s cl es i s ra pidly a d a p ta bl e t o

t he a m ou nt of s u s t a i ned a cti on re q u i red We k n ow th a t
.
”
bei ng ou t o f c on diti on i s u su a lly l es s s h own by th e i na bili ty
“
,
to pu t ou t a vi ol ent effort th a n by the i na bility t o conti nue

TH E INNE R T ISS U E S OF A NIMA L S .
3 69
m a king vi ol ent eff ort s ; a n d we kn ow th a t the res u lt of t ra i n

i n g for p ri z e fi ghts a nd r a ces is m ore s h own i n th e p ro l ong a
-
,
ti on of en ergy t h a n i n th e i n t ens ifi ca ti on of en ergy A t t he .
s a m e ti m e e xp eri en ce h a s t a u ght u s th a t t h e st r u c t u ra l c h a n ge
,
w h i ch a ccom p a ni es thi s fu ncti ona l ch a n g e i s n ot s o m u c h a ,
c h a n ge i n t h e b u lk o f t h e m u s cl es a s a ch a ng e in th ei r i n t er
n a l s ta t e : i n s t ea d of be i ng s o ft a n d fl a bby th ey be c om e h a rd .
We h a v e i ndu ctiv e p roo f th en tha t e xerci s e of a m u s cl e c a u ses

, ,
s o m e i n t er s ti t i a l gr ow th a l ong w ith th e p ower o f m ore s u s
t a i n ed a c t i o n ; a n d th ere c a n be n o d ou bt th a t th e one i s a
con di t i on t o t he oth er Wh a t i s this i nt ers ti ti a l growth ?
.
T h ere i s r ea s on t o s u s p e ct t h a t it i s i n p a rt a n i nc rea s ed
d ep os i t of t he s a rcou s s u b s t a nc e a nd i n p a r t a d ev el op m ent of
bl ood v es sel s Mi cros COpi c ob serv a ti on t end s to c onfi rm the
-
.
conc l u s i on s b efor e d ra w n th a t r ep et i t i on o f c ont r a c ti on s fu r

,
th er s th e for m a ti on o f the m a tt er whi ch con t ra ct s a nd th a t ,
g rea t er d ra u ght s of bl oo d d et er mi ne g rea t er v a s cu l a r i ty .
A nd if th e cont ra s t s of m ol ecu l a r s t ru ct u re a nd th e con t r a s t s

of v a s c u l a r ity di rec tly c a u s ed i n m u s cl es by c ont r a s t s i n th ei r
,
a ctiviti es ,
a r e t o a ny d e g ree i n h erit a bl e ; th ere resu lt s a n
e xpl a n a ti on of th os e c ons tit u ti on a l differ enc es i n th e co l ou r s
a n d t e xt u r es o f m u s cl e s w hi c h a c c om p an y con stit u t i ona l

,
differen ce s i n th ei r d eg rees of a ctivity .
I t m a y be a dd ed th a t if we a re w a rra nt ed i n s o a s c ribi n g
the di fferenti a ti on s of m u s cl es f ro m one a noth er t o di r ec t
eq u ilib r a ti on then we h a v e th e m ore rea son for thi nkin g
,
th a t th e di fferenti a ti on of m u s cl es i n genera l f rom oth er

s t r u c t u r es i s a l s o due t o di rect equ ilib ra ti on T h a t u nlik e
.
n esses be t w een p a rt s o f the c on t ra c til e ti s s u es h a vi n g u n lik e
fu n cti on s a re c a u s e d by th e u nlik enes s es of th ei r fu n cti on s

, ,
r en d er s it th e m ore p r ob a bl e th a t th e u nlik en es s es b et w een
con t r a c til e ti s s u e a n d o th er ti s su es h a v e b een c a u s ed by a na

,
l og on s u n lik enes s es .
3 04 . T h es e i nt erp ret a ti ons whi c h h a v e a l rea dy occu pi ed

,
t oo l a rge a s p a ce,
mu s t h ere be cl o s ed Of c ou rs e ou t of
.
70
3 70 P H Y SI OL OGICA L D E V E L OP ME NT .
ph enom ena s o mu ltitu di nou s a n d v a r i ed it h a s b een i m pra c ,
t i ca bl e t o d ea l w ith a ny bu t the m os t i m p ort a n t ; a n d i t h a s

b een p ra cti ca bl e to d ea l with th es e only i n a gener a l w a y .
Mu ch h owev er a s rem a i n s t o be expl a i n ed I thi nk th e p os s i

, , ,
bi l i ty o f t ra ci ng in s o m a ny c a ses t he a ct i ons t o whi ch th es e

, ,
i ntern a l di fferent i a ti ons m a y ra ti on a lly be a scrib ed m a k es it ,
lik ely th a t t he rem a i ni ng i nt erna l di fferenti a ti ons a re d u e t o

k ind red a cti on s We fin d evid ence th a t i n m ore c a s es
.
,
t h a n s eem ed p rob a bl e th es e a cti on s p r od u ce th ei r effec t s

,
di rec tly on th e i ndivid u a l ; a n d th a t t he u nlik en es s es a re

p r od u ced by a ccu m u l a ti on of s u ch effect s f rom genera ti on t o
g enera t i on Whil e for a l l th e oth er un lik enes s es w e h a v e
.
, ,
a s a n a d equ a t e c a u s e th e i n di rect effects w r ou ght by th e su r

,
~
vi va l genera ti on a ft er g enera ti on o f th e i n divid u a l s i n w hi c h

, ,
f a v ou ra bl e v a ri a ti on s h a v e oc cu rred— v a ri a ti on s s u ch a s t h os e
of w hi c h h u m a n a n a t o my fu rni s h es en dl ess i n s t a n ces T hu s .
a c c ou n ti ng fo r s o m u ch we m a y n ot u n rea s on a bly p res u m e

,
th a t th ese co op era t iv e p roces s es of di rect a n d i ndi rect equ il i

-
b ra ti on will a cc ou n t for wh a t rem a i ns .
N
[ OTE — A fter
h a vi n g di s mi s sed thi s revi sed ch a pt er a s
d one wi th a nd s ent it to th e p ri nter fu rther th ou ght c on
, ,
c erni ng th os e diff erent i a ti on s w hi ch p ro d u ce b on e h a s re ,
mi nd ed m e of a f a ct o f ext rem e a n d v ari ed s ign ifi ca nce '
n a m e d i n th e fi rs t v o l u m e I refer t o th e form a ti on o f
.
a d a ptiv e s t ru c tu res r oun d th e en d s of di s l oc a t ed b ones a n d ,

”
t o th e f or m a t i on of
“
f a l s e j oi nts .
T h es e a re ont ogenetic ch a n ges o f whi ch phyl ogeny yi eld s

n o e xpl a na ti on T h ey do n ot repe a t the t ra it s o f a nc es t ra l
.
org a nis m s , a nd th ey c a nn o t be a s c r ibe d t o eith er o f t h e
rec og ni z ed ev o l u ti on a ry fa ct ors I f a h u m er us be b rok en

.
a cros s a n d fa ili ng t o s et p res ently c om es t o h a v e i ts two

, ,
.
l oos e en d s s o m odifi ed a s i n a m ea s u re t o s imu l a t e the p a rt s

o f a nor m a l j oi nt — th e end s be c om i n g s m o oth c ov ered w ith ,
p eri os t eu m a nd s u ppli ed with fib rou s ti ssu e a nd a tt a ch ed ,
by liga m ent s i n s u ch w a y s a s to a ll ow of res t ra i ned m ov e

TH E I NNE R T ISS U E S OF A NI MA L S .
m ents — it i si m p oss ibl e t o t hi nk th a t na t u ra l s el ecti on h a s

h a d a nythi n g t o do with th e p ower o f a dj u stm ent thu s
s h own . N o s u r viv a l o f i n divid u a l s i n whi ch a d a pt a ti on s
o f t hi s ki n d n ow i n one pl a c e a n d n ow i n a n o t h er were
, ,
b et ter a nd b etter eff ected c ou ld a cc ou n t for a cqu i rem ent of

,
the a bility . N or c a n i t be s u pp os ed th a t the a bility might

r es u l t f r om a fu ncti ona lly p r od u c ed h a bit ; s i nce it i s
-
s ca rc ely c on ceiv a bl e th a t th e n u mb er o f c a ses i n whi c h indi
v i du a l s p r ofit ed by it ( a t fi r s t a littl e a n d gr a d u a lly m ore )
c ou ld be s u ch ( ev en did th ey s u rviv e ) a s t o a ffect the

c on s t i t u ti on o f t h e s p ec i es B oth of t he a ll eged c a u s es o f
.
s t ru c t u ra l m o difi c a ti on s a r e ou t of c ou r t I t is m a nifes t
.
,
t oo th a t t h e foreg oi ng hyp oth es i s r es pe cti n g b one f orm a ti on

,
-
yi eld s u s not th e s light es t h elp .
Bu t on c a re fu lly c ons id eri n g t he f a ct s c ert a i n ph en om en a

,
o f p r ofou n d m ea n i n g m a y s t r ik e u s H e r e i n a p a rt o f t h e
.
,
b ody w h ere no s u ch ti s s u es ordi n a rily exi s t a n d t o w hi ch no

s u ch s t ru c t u r e s a r e o r di n a r ily a pp r op r i a t e th er e a r i s e tiss u e s
,
a n d s t ru c t u r es a d a pt ed t o th e phy s i c a l ci rcu m s t a n c e s imp os ed
on th a t p a r t Ou t of w h a t do th ese a bn orm a l bu t a pp rop r i a t e

.
ti ss u es a r i s e ? T he s u b s t a nces a rou nd— o s s eou s c a rtil a gi n ,
ou s m emb ra n ou s— c on s i s t o f diff er en ti a t ed e l em ents t oo fa r

,
sp ec i a li zed t o a ll ow o f t ra ns form a ti on T hes e new ti s su es .

,
th en m u s t origi na t e from th e u n differen ti a t ed p rot opl as m

,
p erv a di ng the p a r t T he u nit s of thi s p rot opl a s m s u bj ect t o

.
,
t he a cti on s p rop er t o a n a rti cu l a ti on b egi n t o a s su m e th e a p

,
O i t e hist o l o gic a l t r a its — a re d et erm in ed by l oca l sti m u li

p pr r a
t o f or m ti ssu e s or di na rily a s s oci a t ed with s u ch stimu li Wh a t .
i s the i nevit a bl e impli c a ti on ? T h es e u nit s— phy s i ol ogi ca l or

c on s tit u ti on a l a s w e m a y ca ll t h em — m u s t h a v e p os s es se d
,
l a t en t p ot enti a liti es of fa lli ng i nt o th ese s p eci a l a rra n gem en t s

u n d er st res s o f s u c h c on diti on s A t one p oi n t th ere a r i s es
.
p eri os t eu m an d a t a noth er lig a m en t ou s ti ssu e whil e for th e ,
s h a pi n g o f th e en d s o f t h e b ones— h er e i nt o a r u d e hi nge d
for m a n d th ere i nt o a r u d e b a ll a n d s ock et f orm a c cor di n g t o

- -
,
th e h a bit u a l m ov em ents— th ere g oes on s om e a pp rop r i a t e de

p os it of b one H ence we m u s t c onclu d e th a t in th e u nits of

.
p rot opl a s m whi ch h a ve n ot yet b ee n org a ni zed i nto s p ec i a l

ti ss u es th ere res id es th e a bility t o t a ke on one or oth er typ e
,
o f hi s t ol ogi ca l s t ru ct u re a c cor di n g t o c i rcu m s t a n c e s ; a n d ,
fu rth er th a t t h ere res id es i n ea c h of th e m the s till m ore

,
m a rv ell ou s a bili ty t o c oop er a t e w ith ki n d re d u n its di s p ers e d
a rou n d i n d ev e l opi n g t h a t a r ra ng em en t o f th e p a rt s r eq u i re d
”
S o th a t w hil e th es e u nit s
“
t o con s t i tu te a f a l s e j oi n t .
h a v e a g en era l p rocli vity t ow a r d s th e s t ru ct u re of th e

o r g a n i s m a s a wh ol e t h ey h a v e a l s o p r ocliviti es t owa rd s
,
s t ru ct u r es p r op er t o th e l o c a l c on di t i ons i n t o whi c h t h ey
fa ll . T h ere is l a t ent i n ea ch u nit t he c ons tit u t i on o f t he

en t i re o r g a ni s m a n d by i m pli ca t i on th e con s t it u ti on of ev ery
org a n ; a n d ea ch u ni t whil e c oo p er a ti n g wi t h t h e a gg r eg a te i s
r ea dy t o t a k e p a r t i n th a t p a rt ic u l a r a r r a n ge m ent p rop er t o
t h e p os i ti on it h a s f a ll en i nt o I f the r ea der will refer b a ck

.
to 9 7 d 9 7 6 i n whi c h it is s h ow n th a t ea ch m ember of a
, ,
h u m a n s oci ety p os s es s es a c om bi n a ti on of p ot enti a liti es lik e

th ese he will be the be tt er ena bl ed t o beli ev e th a t this thi ng
,
m a y be s o whil e h e i s u n a bl e t o c on c eiv e h ow it i s s o .
A n d h ere i nd eed l et it be p oi n t ed ou t h ow c ompl et ely

, ,
i rrel ev a n t i s th e t es t of c onc eiv a bl enes s a s a ppli ed t o th ese

u ltim a t e phy s i ol ogi ca l a cti on s F or a s h ere f ro m the un
.
,
u n it ed end s o f th e b r ok en b one th ere p r es en tly a ri ses a ru d e

,
j o i nt with fi t m em b ra nes liga m ents a nd ev en syn ovi a l fl u id

, , ,
th ou gh w e a re a b s ol u tely u na bl e t o i m a gi ne th e p roc es s by
which th e a dj a cen t ti s s u es p ro d u c e thi s s t ru ct u re ; s o th ere
m a y be f r o m a n or g a n en l a rge d by fu n c ti on s u ch rea ctiv e ,
e ffec t u p on t h e s y s t e m a t l a rge a s ev en t u a lly t o i nfl u ence t h e
re p r od u ct iv e c ells th ou gh w e m a y be a b s ol u t ely u na bl e to
,
i m a gi ne how thi s c an be d one ]

CH A P TE R I X .
PH Y S IO L O GI CA L I N T E GRA T IO N IN AN I M AL S .
§
3 05 . P H Y S IO L O GI CA L
di ff e renti a ti on a nd phy s i ol o gic a l
i nt egra ti on a re c orrel a tiv es th a t v a ry t og eth er We h a v e but

,
.
t o rec oll ect the f a mili a r p a ra ll el b etw een the divi s i on of l a b ou r

i n a s oci ety a nd th e phy s i ol ogi c a l di vi s i on o f l a b our, to s ee
tha t a s f a s t a s the ki nds of w ork p er f orm ed by the com
p onent p a rt s of a n org a ni s m b ec om e m ore nu m erou s a nd a s ,
f a s t a s ea ch p a rt b ecom es m ore res t ricte d t o i ts own w ork s o ,
f a s t mu st th e p a rt s h a v e th ei r a cti ons c ombi ned i n s uch wa ys

th a t no one c a n go on with ou t the rest a nd the res t c annot
g o on w ith ou t ea ch one .
H ere o u r i nqu i ry m u s t be h ow the rel a ti ons hip of th es e

,
two p r oc es ses is es t a bli s h e d— wh a t c a u s es the i nt egra ti on t o

a dv a nce
p a ri p a ss u with the differenti a ti on T h ou gh it i s
.
m a ni fe s t d p ri or i th a t the m u tu a l d ep end ence of f u ncti ons

, ,
m u s t be p r op orti on a t e t o the sp eci a li z a ti on of fu nc ti ons ;

yet
it rem a i ns t o fi nd the m od e i n whi ch the i nc rea s i ng co ordi -
na ti on i s d et er m i ned .
A l rea dy a m ong t he I n d u cti ons of Bi ol ogy thi s rel a t i on

, ,
b etween differenti a ti on a nd i nt egr a ti on h a s been sp ecifi ed

a n d ill u s t ra t ed Bef ore d ea li ng with it d ed u ctiv ely ,
a few fu r th er ex a mpl es
grou p ed so a s t o exhibit its s ev era l
,
a s p ect s will be a dv a nt a g e ou s
, .
3 06 .
I f the l owly g ni zed P l a na ri a ha s i ts b ody b rok en
-
or a ,
up a nd its gu ll e t d et a c h ed this wi ll, for a whil e c ontinue

, ,
3 73
3 74 P H YS I OL OGI CA L D E VE L OP ME NT .
to p erf orm when ca ll ed u p on j u s t a s th ou gh it

i ts fu n cti on ,
w ere i n i t s pl a ce : a fr a g m ent of th e c r ea t u re s own b ody

’
pl a ced i n th e g u ll et will be p r op ell ed th r ou gh i t or sw a l

, ,
l owed by i t B u t a s th e s eem i ng s t ra n genes s of t h i s fa c t

.
,
i m pli es we fi n d no su c h i n d ep end ent a cti on s of a na l og ou s

,
p a rt s i n th e hi gh er a n i m a l s A ga i n a pi ece cu t ou t o f th e
.
,
di s c of a M edus a c onti nu es with g rea t p ers i s t ence r ep ea ti n g

t h ose rhyt h m i ca l c ont ra c ti ons w hich w e s ee i n th e di s c a s a
w h ol e ; a nd thu s p rov es t o u s th a t th e c ont ra c til e f u n c t i on
i n ea ch p orti on Of t he di s c i s i n g rea t m ea su re i nd ep en d en t
, .
Bu t it i s n ot s o w ith the l ocom ot iv e or g a n s of m ore di fferen

t i a ted typ es Wh en s ep a r a t e d from the res t th ese l ose th ei r
.
p ow ers of m ov em en t T he only m emb er o f a v ert eb r a t e

.
a ni m a l which c on ti nu es t o a ct a f t er d e t a c h m en t i s th e h ea rt ;
,
a nd the h ea r t h a s m ot or p ow er s compl e t e w ithi n it sel f .
Wh ere th ere i s thi s s m a ll d ep en d ence of ea ch p a r t u p on

th e w h ol e th ere is bu t s m a ll d ep en d en ce of th e wh ol e u p on
,
e a c h p a rt . T he l onger tim e whi c h it t a k es for the a rres t of

a f u n c ti on t o p rod u c e d ea th i n a l es s —di fferenti a t ed a n i m a l
th a n i n a m ore—di fferenti a t ed a n im a l m a y be illu s t ra t ed by

,
t h e c a s e of res pi ra ti on S u ff oc a ti on i n a m a n s p eedily
.
c a u ses res i s t a nce t o th e p a s sa ge of the bl ood th rou gh the

c a pill a ri es f oll owed by conges ti on a nd s t opp a ge of the h ea rt
,
g rea t di s tu rb a nce th r ou gh ou t the s y s t em resu lt s i n a few

s ec on d s,
a nd i n a mi nu t e or tw o a l l t h e fu n c ti ons c ea s e .
Bu t i n a f rog with i ts u n d ev el op e d r es pi ra t or y or g a n a nd a
, ,
s ki n th rou gh whi ch a c ons id era bl e a e ra ti on o f th e bl oo d i s
c a rri ed on b rea thi ng m a y be s u s p end ed for a l on g tim e with

,
ou t i n j u ry D ou btl ess this di fference is p roxim a t ely du e t o

.
the g rea t er fu ncti on a l a c tivity i n th e one ca se th a n i n th e

o th e r a n d th e m ore p res s i n g ne ed for di s ch a r gi n g th e
, p ro
du ced c a rb on di oxid e ; bu t th e grea t er fu n cti on a l a ctivity b ei ng

it s elf m a d e p os s ibl e by th e high er sp eci a li za ti on of fu nct i ons ,
thi s rem a i ns th e p rim a ry c a u s e of th e g rea t er d ep en d en ce of

t he oth er f uncti on s on res pi ra ti on wh ere th e res pi ra t ory
,
a pp a ra t u s h a s b ec o m e highly s p eci a li zed .H er e

P H YSI OL OGICA L I NT E GRA TI ON I N A NIMA L S .
i nd eed we s ee the rel a ti on u n d er a noth er a s p ec t T hi s m ore

,
.
ra pid r hyth m of th e fu n c ti on s w hi c h i nc rea s ed h e t er o g eneity
o f s t ru ct u re m a k es p os s ibl e i s it se lf a m e a n s o f i n t eg ra t i n g
,
th e fu nc ti on s . Wa t ch wh en it i s ru nni ng d own a c om pli

, ,
c a t ed m a chi ne of whi c h th e p a r ts a re n ot a cc u ra t e ly a dj u s t ed ,
o r a r e so w o rn a s t o be s om ew h a t l o o s e T h ere w ill be
.
Ob served c ert a i n i rregu l a r iti es of m ov em ent j u s t b ef or e it

c o m es t o res t— c e r t a i n of th e p a rt s w hi c h s t op fi rs t a re ,
a g a in m a d e t o m ov e a littl e by t h e con ti nu e d m o v e m en t o f
th e res t a n d th en b ecom e th em s elv es i n t u rn th e c a u s es o f

, , ,
renewe d m oti on i n o th er p a r t s whi c h h a v e c ea se d t o m ov e .
T h a t i s t o s a y whil e t he conn ect ed rhyth m i ca l ch a n ges o f

,
t he m a chi ne a r e qu i c k th ei r a c ti on s a n d rea cti on s on one

,
a n oth er a re r e gu l a r— a l l th e m o ti on s a re we ll i n t eg ra t e d ; bu t
a s the v el oci t y di m i n i s h es i rregu l a r i t i es a ri s e—th e m o ti on s
b ecom e s omew h a t di s i nt eg ra t ed S i m il a rly with or g a nic

.
fu nc ti ons : i ncrea se of th ei r ra pidity i nv olv es i nc rea s e Of a

j oi nt m om ent u m whi ch cont rol s ea c h a n d c o ordi n a t es a l l -
.
T h u s if we comp a re a s na k e with a m a m m a l w e s ee th a t ,
i t s fu n cti on s a r e not ti ed t ogeth er s o c l os ely T he m a m m a l

. .
,
a nd e s p eci a lly th e s u p eri or m a m m a l requ i res f ood w ith c o n

,
s i dera bl e re gu l a rity ; k ee p s u p a r es pi ra ti on whi c h v a r i es
w ithi n bu t m o d era t e li m i t s ; a nd h a s p eri od s of a c tivity a n d

res t th a t a lt ern a t e ev enly a n d fr equ ently B u t th e s n a k e
.
,
t a ki n g food a t l on g i nt erva l s m a y h a v e th es e i nt erv a ls

,
g rea tly ext end ed with ou t fa t a l resu lt s ; i ts d or m a n t a nd i ts

a c t iv e s t a t es recu r l es s u n i for mly ; a n d i t s r a t e of r espi r a ti o n
v a ri es withi n m u ch w id er li m it s— n ow b ei n g s ca r c ely per

c ept i bl e a nd n o w a s you m a y p r ov e by e x c i t i ng it b ecomi n g
, ,
c on s pi cu ou s . S O th a t h ere w h ere th e rhyth m s a r e v ery s l ow

, ,
th ey a re i ndivid u a lly l ess regul a r a nd a re u nit ed in t o a l es s

,
r eg u l a r c om p ou n d rhyt h m m
a r e l es s i nt eg r a t e d .
P erh a p s the c l ea res t gener a l id ea of th e co ordi n a ti on o f -
f uncti ons th a t a cc omp a ni es th ei r s p eci a li za ti on i s Obt a ined ,
by ob s ervi ng th e sl ownes s with whi ch a littl e diff erenti a t ed -
a ni m a l res p on d s t o a s tim u l u s a ppli e d t o on e o f i t s p a rt s ,

a nd the pidity with w hi ch s u ch a l oca l s ti m u l u s is re

ra
s p onded t o by a m ore diff erenti a t ed a nim a l-

A s ea a nem one .
-
a nd a fly will s erv e for t h e c omp a r i s on A t en t a c l e of a s ea .
a n em on e w h en t ou c h e d
,
s l ow ly c on t ra c t s ; a n d if t h e t ou c h
,
h a s be en ru d e th e cont ra c ti on p res ently ext en d s t o the oth er

,
t ent a c l es a nd even tu a lly t o t he enti re b ody : the s ti m u l u s to

m ov em en t i s g r a d u a lly diff u sed th rou gh ou t t he o rg a n i s m .
Bu t i f you t ou c h a fly or r a th er i f you c om e nea r enou gh

,
t o th rea t en a t ou ch the enti re a pp a ra t u s of flight i s i n st a nt ly

,
b rou gh t i nt o c om bi ned a cti on Wh ence a ri s es this c ont ra s t ?.
Th e one crea t u r e h a s bu t f a i ntly s p eci a li zed c on t ra ctil e

o r g a n s a nd fib r es for c on v eyi ng imp res s i on s
,
T he oth er h a s .
d efinit e m u s cl es a nd nerv es a nd a c o or di na ti ng cent r e T he -

.
p a rt s Of t he li tt l e differenti a t ed s ea a nem one h a v e th ei r

- -
f unc t i ons s o feebly c o ordi na t ed th a t one m a y be s t rongly

-
,
af fect ed for s om e tim e b ef or e a ny eff ect i s f elt by a noth er a t

a di s t a n ce fro m it ; bu t i n t he m u c h di ffer en ti a t ed fly v a ri ou s -
,
re m ot e p a r t s i n st a ntly h a v e c h a n g e s p r op a g a t e d t o th e m f r om
t he a ffec t e d p a r t a n d by th ei r u nit ed a c ti ons th u s s et u p th e

, ,
wh ol e organi s m a dj u s ts it s elf s o a s t o a v oid the d a nger .
T h es e few a dd ed il l u s t ra ti on s will m a k e th e n a t u re o f thi s

g enera l rel a ti on s uffi ci ently cl ea r L et u s now p a s s to the .
i nt erp ret a t i on of it .
§ 3 07 . If a H ydra i s t itive liq u ids

cu t i n tw o, the nu r
di ffu s ed th rou gh i ts s u b s t a nc e c a nnot esca p e ra pidly s i nce ,
th ere a re no op en ch a nnels for th em ; a nd h enc e the c on

d i ti ons o f the p a rts a t a di s t a nc e from th e cu t i s bu t li ttl e
a ffect ed Bu t wh ere a s i n the m ore differenti a t ed a n im a l s
.
,
-
,
t he nu t ritiv e liq u id is c ont a i ned i n v es s el s which h a v e con

t i nu ou s c om m u n ic a ti ons c u tti ng the b ody i n two or c u tti ng
, ,
o ff a n
y c ons id era bl e p orti on of it i s f oll ow ed by esc a p e of ,
th e liqu id f rom th es e v ess el s t o a l a rge ext ent ; a nd thi s

a ff ec ts the n u t riti on a nd ef fici ency of orga ns rem ot e f ro m
the pl a c e of i n j u ry T h en wh ere a s in fu rth er d ev el op ed
.
,
-
c rea tu res th ere exists an a pp a ra t us fo r p rop el li ng the bl ood

,
P H YS IOL OGICA L I NT E GRA TI ON I N A NI MA L S .
3 77
thr ou gh th es e ra m ifyi ng ch a nnel s i n j u ry of a s ingl e one ,
will c a u s e a l oss of bl ood th a t q u i c kly p ros t ra t e s the enti re

o rg a ni s m H enc e th e ri s e of a c ompl et ely difl erenti a ted va s
.
-
cu l a r s y s t em is the r i s e of a sys te m w hich i nt egra t es a l l

,
m embe rs o f th e b ody by m a ki ng ea ch d epend ent on the i n

,
t egri ty of t he v a scu l a r sy s t em a n d th eref ore on the i nt e grity

,
o f ea ch m em be r th rou gh which it ra m i fi es In .
a noth er m o d e t o o t he es t a bli s hm en t o f a di s t ribu t i ng a pp a

, ,
ra t u s p r o d u c es a phy s i ol o gi c a l u n i on th a t is gr ea t i n p rop or
ti on a s this di s t rib u ti ng a pp a ra t u s i s effici ent A s f a s t a s it .
a ss u m es a f u nc ti on u nlik e t h e res t ea ch p a r t Of a n a ni m a l
,
m odifi es the bl o od i n a wa y m ore or l es s u nlik e the res t b oth ,
by the m a t eri a l s it a b s t ra ct s a n d by the p r od u ct s it a dd s ;

a n d h en c e th e m o re diff er en ti a t e d the v a s cu l a r s y s t e m be
c om es the m ore d oes it int eg ra t e a l l p a rt s by m a ki ng ea ch
,
o f th em f ee l th e qu a lit a tiv e m o difi c a ti on of t h e bl oo d whi c h
ev ery oth er ha s p rod u c ed T hi s i s s imply a n d c ons pi cu ou s ly

.
ex em plifi ed by the l u ng s I n the a b s enc e of a v a scu l a r

.
s y s t em or i n the a b s ence o f on e th a t i s w ell m a rk e d Of

,
f
from the imbe ddi n g tis s u es th e nu t ritiv e pl a s m a or the c ru d e
,
bl ood gets wh a t s m a ll a e ra ti on it ca n only by comi ng nea r

, ,
’
th e c rea t u r e s ou t er s u rfa ce or th o se i nner s u r f a c es whi ch a re
,
b a th ed by w a t er B u t w h ere th ere h a v e b ee n for m ed d efi nit e

.
c h a n ne l s b ra nchi ng th r ou gh ou t th e b ody a n d p a r tic u l a rly

,
wh ere th ere e xi s t s p eci a li zed org a ns for p u mpi ng th e bl oo d

th rou gh th ese ch a nnel s it m a ni fes tly bec om es p os s ibl e for
,
the a era ti on t o be c a rr i e d on i n one p a rt p e cu li a r ly m odifi e d

t o fu rth er it whil e a l l oth er p a rt s h a v e th e a e ra t ed bl oo d
,
b rou ght to th em A nd how grea tly the differenti a ti on of th e

.
v a scu l a r sy s t em thu s b ecom es a m ea ns of i nt egra ti ng th e

v a ri ou s org ans i s sh own bv the f a t a l res u lt th a t f oll ows wh en
,
th e c u rr ent of a era t ed bl ood is i nt erru pt ed .
H ere i nd eed it b ec om es obvi ou s b oth th a t c ert a i n physi o

, ,
l ogic a l di fferenti a ti ons m a k e p ossibl e c ert a in phy s i ol ogic a l

i nt eg ra ti ons ; a nd th a t c onv ers ely th es e i nt eg ra ti ons m a k e
, ,
p os s ibl e oth er differen ti a ti on s Besid es th e w a st e p ro d u cts

.
3 78 P H YS I OL OGI CA L D E VE L OP ME NT .
w hi c h p e th r ou gh th e l ung s th ere a re w a s t e p rodu cts

es ca ,
w hi c h es c a p e th r ou gh t h e s ki n th e ki d ney s th e liv er
, ,
Th e .
bl ood h a s s ep a ra t ed f rom it i n ea ch of thes e st ru ctu res the ,
p a rticu l a r p rod u ct w hi ch thi s s t ru ct u re h a s b ecom e a d a pt ed

t o s ep a r a t e ; l ea vi n g th e oth er p rod u c t s t o be sep a r a t ed by the
oth er a d a pt e d s t r u ct u re s .H ow h a v e th es e s p ec i a l a d a pt a
ti ons b een m a de p os s ibl e ? By un i on of the orga n s a s
rec ipi ents o f one c i r cu l a ti n g m a s s O f bl ood Whil e th ere i s no
.
eff i ci ent a pp a ra t u s for t ra n s f er of m a t eri a ls th r ou gh th e b ody ,
th e w a st e p ro d u c ts of ea ch p a rt h a v e t o m a k e th ei r es c a p e
l oc a lly ; a nd the l oc a l ch a nn el s of es c a p e m u s t be comp et ent
t o t a k e Off i nd ifferently a l l th e w a s t e p rod u cts B u t it .
b ec om es p ra ctic a bl e a n d a dv a nt a geou s for the differently

l oc a li zed ex cretin g s t ru ctu res t o b ec om e fitt ed to s ep a ra t e
differen t w a s t e p r od u ct s a s s oon a s the c om m on ci rcu l a ti on
,
th rou gh th em grows s o effi ci ent th a t th e p rod u c t l eft u nex

ers t ed by one is q u ickly c a r ri ed t o a n oth er b ett er fitt e d t o
exc ret e it . S o th a t the i n t egra ti on Of th em th r ou gh a

c omm on v a s cu l a r sys t em i s th e c on diti on u nd er w hi ch only
,
th ey ca n b ec om e differen ti a t ed . P erh a p s th e cl ea res t

id ea of the wa y i n whi ch di ff erenti a ti on l ea d s t o i nt eg ra ti on ,
a n d h ow , a ga in i ncrea sed i n t egra ti on m a k es p os s ib l e s till

,
f u rth er differenti a ti on w i l l be Obt a i n ed by cont em pl a ti ng the

,
a na l og ou s d ep en d en c e i n th e s oc i a l org a ni s m Wh il e it h a s
.
no r oa ds a cou nt ry c a nn o t h a v e i ts i nd u st r i es m u ch s p ec i a l
,
i zed : ea c h l oca lity m u s t p ro d u c e a s b es t it ca n the v a ri ou s

, ,
c om m oditi es it c onsu m es s o l ong a s it h a s n o fa ciliti es for

,
b a rt er with oth er l oca liti es Bu t th e l oca l iti es b ei ng u nl ik e i n

.
th ei r na t u ra l fi tnes ses for the v a ri ou s i n d u s t ri es th ere t ends ,
ev er t o a ri s e s om e e x ch a n ge Of the c o m m oditi es th ey ca n
res p ec tiv ely p ro d u c e w ith l ea s t l a b ou r T hi s ex ch a ng e l ea ds

.
t o th e f orm a ti on o f ch a nnel s of c om m u n i ca ti on The cu r .
r ents o f c om m oditi es on c e s et u p m a k e th ei r f o ot p a th s a n d
,
-
h ors e t ra cks m ore p erm ea bl e ; a nd a s f a st a s the res ist a nce

-
t o exch a nge b ec om es l ess the c u rrents of c omm oditi es

,
b ec om e grea t er . E a ch l oc a lity t a k es m ore of th e p rod u ct s

P H YS I OL OGICA L INT E GRA T I ON I N A NIMA L S .
of a dj a cent ones a n d ea c h l oca lity d ev ot es it s el f m ore t o the

,
p a rticu l a r i ndu s t ry for w hi ch it i s na t u ra lly b es t fit ted : the

fu n cti ona l i nt egra ti on m a k es p o s s i bl e a f u rth er fu ncti ona l
di fferenti a t i on T hi s fu rth er fu ncti ona l di fferen ti a ti on rea ct s
. .
T h e g rea t er d em a n d for th e s p ec i a l p rod u c t Of ea ch l oca lity ,
ex c i t e s i m p r ov e m en t s i n p r o d u c ti on— l ea d s t o th e u se o f
m e th od s whi ch b oth ch ea p en a nd p erf ec t the c omm odity

'
H ence res u lt s a s till m ore a ctiv e exch a nge ; a s till c l ea rer

o pe ni ng o f th e c h a nnel s of c o mm u n ic a ti on ; a still cl os er
mu t u a l d ep en d ence Y et a n ot h er i nfl u ence com es i n t o pl a y

. .
A s fa s t a s t he i nt er cou rs e a t fi rs t o n ly b et ween n eighb ou ri ng

,
l oca liti es m a k es for it s el f be t ter roa d s— a s f a s t a s r iv ers a re

,
b ridged a nd m a rsh es m a d e ea s ily p a ss a bl e the r es i s t a nce t o

,
di s t rib u ti on b ec om es s o fa r dimi ni s hed th a t the thi n g s g rown

,
or m a d e i n ea c h di s t rict ca n be p r ofit a bly c a r r i ed t o a g r ea te r
di s t a nce ; a nd a s the econ omi ca l i ntegr a ti on i s thu s e xt end ed

o v er a w id er a rea th e economic a l di fferen ti a ti on i s a ga i n
,
i nc rea sed ; s i nc e ea ch di s t ri c t h a vi ng a l a rger m a r k et for its

,
c om m odity ,
i s l ed t o d ev ote it self m ore excl u s iv ely t o pro
du c i ng thi s comm odi ty T h es e a cti on s a nd rea c ti on s c on
.
ti nne u ntil the v a ri ou s l oca li ti es b ec om i ng g rea tly d ev el op ed

,
a n d highly s p ec i a li z ed i n th ei r i n d u s t r i es a r e a t th e s a m e
,
ti m e fu nct i on a lly i nt egra t ed by a net w or k o f roa d s a nd ,
fi na lly ra il wa y s a l ong which ra pidly c i rcu l a t e th e cu rrent s

,
s ev er a lly s en t ou t a n d r eceiv e d by th e l oc a liti es A n d it will

.
be m a n ifes t th a t i n i n divid u a l o rga n i s m s a lik e c or rel a tiv e

p r ogres s mu s t h a v e b een cau s ed i n a n a na l og ou s wa y .
3 08 . A noth er an d
high er f orm of phy s i ol ogi ca l i nt egra
ti on i n a ni m a l s i s th a t w hi ch the nerv ou s s y s t em effec t s
, .
E a ch p a rt a s it b ec om es s p eci a li zed b egi ns t o a ct u on th e

'
, p
res t n ot o nly i ndi re c tly th r ou gh t h e m a t t er s it t a k es fro m
a n d a dd s t o t h e bl o o d bu t a l s o di rec tly t h r ou h th e m o l e u l a r
, g c
di s t u rb a nc es it s et s u p a n d di ffu s es Wh eth er n er v es th em
.
s elv es a r e di ffer enti a t e d by th e m ol ecu l a r di s t u r b a n ces th u s
p r op a ga t ed i n cert a i n di recti on s or wh eth er th ey a re oth er

,
3 80 P H Y S I OL OGICA L D E V E L OP ME NT .
wi s e di fferenti a t ed it mu s t eq u a lly h a ppen th a t a s f a s t a s

,
th ey bec om e ch a nnel s a l on g whi ch m ol ecu l a r di s t u rb a nce s

t ra v el t he p a rt s th ey c onne ct b ecom e phy s i ol ogi ca lly i nte
,
g ra t ed i n s o fa r tha t a c h a nge i n one i nit i a tes a ch a nge i n

,
t he oth er We m a y di m ly p erceiv e t h a t if p orti ons of wh a t

.
w a s origi na lly a u ni form m a s s h a vi ng a com m on f u ncti on ,
u n d er t a k e s u bdivi s i ons o f the fu nct i on the m ol ec u l a r

,
c h a ng es g oi ng on i n th em w ill be i n s om e w a y co m p l em en
t a ry t o one a n ot h er : th a t p ecu li a r form of m ol ec u l a r m oti on

whi ch th e on e h a s l os t i n b ecomi ng s p eci a li zed the o th er h a s
,
g a i ned i n becom i ng s p ec i a li zed A nd i f the m ol ecu l a r m oti on

.
th a t wa s c om m on to the two p orti ons whil e they w ere u ndi ffer

enti a t ed b ec om es di vid e d i n t o t w o c om pl em ent a ry ki n d s o f
,
m ol ec u l a r m oti on ; t h en be t ween th es e p orti ons th ere will be

a c on t ra s t o f m ol ecu l a r m oti on s s u ch th a t w h a t ev er i s l us
p
i n the on e will be m i n us i n the o th er ; a nd h enc e th ere w ill be
a s p ec i a l t end en cy t ow a rd s a re st ora ti on of the m ol ecu l a r eq u i
li b riu m b etw een the two : the m ol ec u l a r m oti on c onti nu a lly

p rop a ga t ed a wa y f rom eith er w ill h a v e its li ne of l ea st resi s t
a nce i n th e di r ec ti on Of t h e o th er . I f a s a r gu ed
,
i n the l a s t ch a pt er rep ea t ed res t or a ti on s of m ol ecu l a r equ il i

,
bri u m a l w a y s foll owi ng the l i n e o f l ea s t res i s t a nc e t en d ev er

, ,
t o m a k e it a li ne of dimi n i s h ed res i s t a n ce ; th en i n p r o p or,
ti on a s a ny p a r t s b e com e m ore phy s i ol ogi ca lly i nt eg ra t e d by

th e es t a bli s hm ent O f thi s ch a nnel for th e ea s y t r a n s mi s s i on
o f m ol ec u l a r m o ti o n b et w een th em th ey m a y b ecom e m ore
,
phy s i ol ogic a l ly diff eren ti a t ed . T he c ont ra s t b et ween th ei r

m ol ecu l a r m oti ons l ea ds t o the li ne of di s ch a rge ; the li ne o f
di s ch a rg e once f orm ed p ermits a grea t er c ont ra s t of th ei r
, ,
m ol ecu l a r m oti ons t o a ri se ; th ereu p on the q u a ntiti es of

m ol ecu l a r m oti on t ra ns ferred t o rest ore equ ilib ri u m b ei ng ,
i nc rea s ed the ch a n nel of t ransf er is m a d e m ore p ermea bl e ;

,
a nd i t s f u rth er p erm ea bility so c au s ed rend ers p os sibl e a s till

, ,
m o re m ark ed u nlik enes s of a cti on be t wee n the p a rts T hu s .
the di fferenti a ti on a nd th e i nt egra ti on p rog res s h a nd i n h a nd

a s b ef ore . H ow the s a m e p rin cipl e h olds th rough
P H Y S IOL OGICA L I NT E GRA T I ON I N A NI MA L S .
3 81
ou t th e higher st a ges of nerv ou s d ev el opm ent c a n be s een ,
o n ly s till m or e v a gu e ly N ev erth el es s i t is com p reh ens ibl e

.
,
th a t a s fu n cti ons bec om e fu rth er divi d ed th ere w ill a r i s e the ,
n ee d for s u b c onn e xi on s a l on g w hi c h th ere m a y t a k e pl a c e

-
s econ d a r y equ i l ib r a ti on s s u b or di n a t e t o th e m a i n o nes I t is .
m a n i fes t t oo th a t wh erea s the di ffer en ti a ti on of fu n cti on s

, ,
p rocee d s n ot neces sa rily by divi s i on i nt o two bu t O ft en by

, ,
divi s i on i nt o s ev er a l a nd u su a lly i n s u ch wa y s a s not t o l ea v e

,
a ny t w o f u n c ti on s th a t a r e j u s t c o mpl e m ent a r y t o one a n
o t h er th e r es t or a ti on s O f eq u ilib ri u m ca nn ot be s o s i m pl e a s
,
a b ov e s u pp ose d A n d es p ec i a lly w h en we b ea r i n mi n d th a t
.
m a ny di ffer en ti a t ed fu n c ti o n s a s t h os e O f the s en s es c a n n o t
, ,
be h eld com pl em en t a r y t o a ny oth er fu n c ti on s i n p a r ti cu l a r ;

it bec om es m a ni fes t t h a t the equ ilib r a t i ons th a t h a v e t o be
m a d e i n a n org a ni s m Of m u ch h et erogeneity a re e xt rem ely ,
c om pl ex a nd do n ot t a k e pl a ce b et w een ea ch org a n a nd s om e
,
o th er bu t b e t ween ea c h org a n a n d a l l th e o th er s
,
T h e p een .
l i a ri ty of th e m o l ecu l a r m oti on p r op a g a t e d fr om ea ch orga n ,
h a s t o be n eu t r a liz ed by s om e cou n t er p ecu li a r ity i n th e -
a v er a g e o f th e m ol ecu l a r m o ti on s wi t h w hi ch it i s b rou ght
i nt o rel a ti on A l l th e v a r i ou s ly m odi fi ed m ol ecu l a r m oti on s

.
-
f rom the v a ri ou s p a rt s mu s t h a ve th ei r pl u s es a nd m i nu s es
,
m u t u a lly c a n cell ed : if n ot l oc a lly th en a t s om e cent re t o

,
wh i c h ea c h u n b a l a n c ed m oti on t r a v el s u nt il it m ee t s wi th
s om e Opp os it e u n b a l a n ced m oti on t o d es t r oy it S till i n v olv ed
.
,
a s th es e a c ti o n s m u s t b eco m e it i s p os s ibl e t o s ee h ow the

,
g enera l p r i nc ipl e illu s t ra t ed by th e s i m pl e ca se a b ov e s u p

.
p os ed will con ti nu e t o h old

,
F or a l w a y s th e m ol ecu l a r
.
m o ti o n p r ocee di n g fr o m a ny o ne di f f erenti a t ed p a rt w ill ,
t ra v el m os t r ea dily t ow a r d s t h a t pl a ce w h er e a m ol ecu l a r
m oti on m o s t compl em en t a ry t o it i n ki n d exi s t s— n o m a tt er
wh eth er thi s c om pl em en t a r y m ol ecu l a r m ot i on be th a t p r o
cee di n g fr om a ny on e o th er or g a n or t h e r es u l ta n t O f th e
,
m ol ecu l a r m oti on s p r o ceedi n g fr om m a n y o t h er o r g a n s SO .
th a t th e t end en c y w ill be for ea ch ch a nn el o f c o m m un i ca ti on

o r ne r v e t o u nit e it sel f with s o m e cen t re or g a ngli on wh ere it
, ,
co m es i nt o l ti on w ith oth er nerv es A n d if th ere be a ny

re a .
p a rts of i ts p ecu li a r m ol ecu l a r m oti on u n ca n cell e d by th e

m ol ecu l a r m oti ons i t m ee t s a t thi s cent re ; or i f a s w ill p rob ,
a bly h a pp en t he a v er a ge m ol ecu l a r m ot i on w hi c h it t h ere

,
un i tes t o p ro d u ce di ffers fr o m t h e a v era ge m o l ecu l a r m ot i on

,
e l sew h er e ; t h en a s b e fore t h ere w ill a ri s e a di sc h a r g e a l o ng

, ,
a n o th er ch a nn e l o r n erv e t o a n oth er c ent r e or g a n gli on ,
w h ere t h e re sid u a ry di fferenc e m a y be ca ncell ed by th e

di fferen c es it m eet s ; or wh ence it m a y be s till fu r th er
p rop a g a t ed till it i s s o ca n cell ed T hu s there will be a ten .
d eney t o a genera l nerv ou s i nt eg ra ti on k eepi ng p a c e with

t he di fferenti a ti on .
Of cou rs e thi s mu s t be t a k en a s n othi n g m ore th a n th e

i ndica ti on of i niti a l t end enc i es— not a s a n hyp oth es i s s u ffi
c i en t t o a ccou n t for a l l th e fa ct s I t l ea v es ou t of s ight the
.
o r igi n a n d f u nc t i on s of g a n gli a c on s id er ed a s s om et hi n g m ore

,
t h a n nerv e j u ncti ons

-
Were th ere on ly th ese li nes of ea sy
.
o f m ol ecu l a r di s t u rb a nc e a ch a n g e s et u p i n ,
o n e org a n c ou ld nev er do m ore th a n p r o d u c e i t s eq u iv a l en t
O f c h a ng e i n s om e oth er or o th ers ; a n d t h ere c ou ld be n one
o f th a t l a r g e a m ou n t o f m ot i on i niti a t ed by a s m a ll s en s a ti on
,
whi c h w e h a bit u a lly s ee Th e fa ct s s h ow u nm i s t a k a bly t h a t

.
, ,
th e s light di stu rb a nce com mu n i ca t ed t o a g a ngli on c a u s es a n ,
o v er th r ow o f th a t highly u n s t a bl e nerv ou s m a t t er c on t a i n ed
-
i n it a nd a di s ch a r g e f ro m it o f the g rea tly i ncr ea s e d qu a nt i ty

,
-
o f m ol ec u l a r m oti on s o g en era t ed T hi s h owev er i s b ey on d

.
, ,
ou r i m m edi a t e t opic A l l we h a v e h ere t o n ot e i s th e i n t er

.
dep end enc e a nd u nifi ca t i on of fu ncti ons th a t n a t u ra lly f oll ow

t he di fferenti a ti on of th em .
3 09 . S om
thi ng might be a dd ed c onc erni ng the f urth er
e
c l a ss of i nt eg ra ti ons by which o rg a ni s m s a re c on s tit u t ed

m ech a n ic a lly c oh eren t wh ol es
-
Ca rryi ng fu rther c ert a i n of
.
t he a r gu m en t s c on t a i ned i n the l a s t ch a pt er it might be ,
n ot u n rea s on a bly i n fer red th a t th e bi ndi n g t og eth er o f p a rt s
by b ones m u s cl es a nd lig a m ents i s a s ec ond a ry res u lt of

, , ,
P HY SIOL OGI CA L INTE GRA T ION I N A NIMA L S .
3 83
th os e s a m e a cti ons by whi ch b ones mu scl es a nd lig a m ent s

, ,
a r e s p ec i a li zed B u t a d equ a t e t rea tm ent o f thi s divi s i on of

.
t he s u bj ect i s a t p res en t sca rcely p os s ibl e .
Wh a t littl e of fa ct a nd i nference h a s b een a b ov e s et d own ,
will h owev er s erve t o m a k e c omp reh en s ibl e th e genera l t ru th s

, ,
r es p ec ti ng whi c h i n th ei r m a i n ou tli ne s th ere c a n be n o

, ,
qu es ti on Begi nni ng wi th the feebly di fl erenti a ted s p onge

.
-
,
Of w hich the in t eg ra ti on is a l s o s o feebl e th a t c u tti ng off a

pi ece i nt er feres i n no a pp reci a bl e d eg ree w i t h th e a ctivity
a n d g r owth of t h e res t it i s u nd en i a bl e th a t the a dv a nce i s
,
th rou gh s t a ges i n whi c h the m u ltipli ca ti on of u n lik e p a rt s

h a vi ng u n lik e a c ti on s i s a c com p a ni ed by a n i ncrea s i ng i nt er
,
d ep end ence of the p a rt s a n d th ei r a cti on s ; u ntil w e c om e t o

,
st ru ct u res lik e ou r own i n which a s ligh t ch a nge i niti a t ed i n

,
one p a rt will i ns t a n tly a n d p owerfu lly a ffec t a l l o th er p a r t s
will c onv u l se a n i m m ens e nu m b er Of mu scl es s end a wa v e of ,
c ont ra cti on th rou gh a l l the bl ood v es s el s a wa k en a c r owd o f

-
,
id ea s wi th a n a cc omp a nyi ng gu s h of em oti on s a ffec t the ,
a c ti on o f th e l u ng s o f t he s t o m a c h a n d Of a l l th e s ec reti ng
, ,
or g a ns A nd whi l e it i s a m a n ifes t nec ess ity th a t a l ong w ith

.
thi s su bdivi s i on o f fu n cti ons whi ch th e high er org a nism s sh ow

u s th ere m u st be thi s c l os e co or di na ti on of th em
,
-
t he f ore ,
g oi ng p a ra gr a ph s s u gges t h ow thi s neces s a ry c orrel a ti on is

b rou ght a b ou t F or a grea t p a rt of th e phy s i ol ogic a l u ni on
.
th a t a ccom p a ni e s th e phy s i ol ogic a l sp ec i a li za ti on th ere a p ,
p ea rs t o be a su ffici ent c a u s e i n th e p roces s of di rect equ il i

b ra ti on a nd i ndi rect equ ilib ra ti on m a y be f a i rly p resu m ed a
su flE ici ent c au s e for th a t which rem a i ns .
CH A P T E R X .
S U M M A RY OF P H Y S IO L O G I CA L D E VE L OP M E N T .
310 . I N T E RC O U RS E b et ween h p a r t a n d the p a rticu l a r

ea c
c ondi ti ons t o whi ch it is e xp osed eith er h a bit u a lly i n the

,
i ndividu a l or occa s i ona lly i n the ra ce thu s a pp ea rs t o be the

,
o rigi n o f phy s i o l o gi ca l d ev el op m en t ; a s we f ou n d it t o be th e
origi n o f m orph o l o gi ca l d e v el op m en t T he un lik enes s es of

.
f or m th a t a ri s e a m on g m em b er s of a n a ggreg a t e th a t w ere
or i g i n a lly a lik e we t r a c e d t o u nlik en esses i n th e i ncid ent
,
f orces A nd i n the f oregoi ng ch a pt er s we h a v e t r a c ed t o

.
u nlik enes s es in the i n cid ent f or c es th os e u nlik en es ses of

,
mi nu t e st ru ct u re a nd ch em ic a l c omp ositi on th a t s i m u l ta
n eou s l y a r i s e a m ong th e p a rt s .
I n s u m m i n g u p t he s p eci a l t r u th s ill u s t ra tiv e of this

g enera l t ru th it will be p r op er h ere t o c ont em pl a t e m ore
,
e s p eci a lly t h ei r d ep en d en c e on fi rs t p r i n c ipl es D ea lin g wi th

.
bi ol o gi c a l ph eno m ena a s ph enom ena o f ev ol u ti on we h a v e t o ,
i nt erp ret n ot only the i ncrea s i ng m orph ol o gi ca l h eterog eneity

o f o r ga n i s m s bu t a l s o th ei r i n crea s i ng phy s i o l ogi ca l h et ero
,
g e nei ty i n t erm s Of th e re di s t rib u ti on o f m a tt er a n d m oti on

-
.
,
Wh il e we m a k e ou r r a pid re s u rv ey of the fa ct s l et u s th en
-
,
m ore p a rt i cu l a rly ob s erve h ow th ey a re s u b ordi na t e t o the

u n iv ers a l c ou r s e of thi s r e di s t r ib u ti on
-
.
i s t a bility o f th e h om og eneou s
T he st i tl
§ 31 1 . n or r c y , ,
sp ea ki ng th e i nevit a bl e l a p s e of the m ore h om og eneou s i nt o

,
th e l es s h om ogeneou s which we b ef ore sa w endl essly exem

,
S U MMAR Y OF P H Y SIOL OGICAL D E V E L OP ME NT .
3 85
p l i fi e d by the m orph ol ogic a l diff erenti a ti ons Of the p a rt s of

o rg a ni s m s we h a v e h ere seen a fresh e x emplifi e d i n w a y s a ls o
,
c ou ntl es s by the phy s i ol ogic a l di ff erenti a ti ons of th ei r p a rt s

,
.
An d i n the one c a s e a s i n t he oth er this ch a nge f ro m u ni,
f or mity to mu lti form ity i n org a ni c a gg reg a t es i s cau s ed a s , ,
i t i s i n a l l i n org a nic a ggreg a te s by t he nec es s a ry exp os u re

,
Of thei r c omp onent p a r t s t o a c ti ons u nlik e i n ki nd or qu a n

ti ty or b oth Genera l p roof of thi s i s f u rni s h ed by the ord er
.
i n w hich th e di ff eren ces a pp ea r I f p a rt s a re ren d ered

.
phy s i ol ogic a lly h et er ogeneou s by th e h et erogeneity of th e i n

e id en t f or c es then the ea rli es t c ont ra s t s s h ou ld be b et ween
,
p a rt s th a t a re the m os t s t rongly cont ra s t ed i n thei r rel a ti on s

t o i ncid ent f orc es ; th e next ea r li es t c on t ra s t s s h ou ld occu r
wh ere th ere a re th e next s t ronges t con t ra s t s i n th es e rel a ti ons
a nd s o on I t t u rns ou t th a t they do s o
. .
E v eryw h er e the di fferen ti a ti on Of ou tsid e f ro m i ns id e

c om es fi r s t I n th e s i m pl es t pl a n t s the u nlik enes s of the
.
c ell w a ll t o the cell c ont ent s i s the c ons picu ou s t ra it of

- -
s t ru c t u re .The con t ra s t s seen i n t he s impl es t a n im a l s a re

of th e s a m e ki n d : th e film th a t c ov ers a Rhi z op o d a n d th e
m ore i ndu ra t ed coa t of a n I n f u s ori a n a re m ore u nlik e the,
c ont a i ned s a rc od e th a n th e oth er p a rts of thi s a re f rom one

a n oth er ; a n d the t en d enc y d u ri n g t h e life o f th e a n im a l i s
for the u nlik enes s t o b ec om e grea t er .Wh a t i s t ru e

o f P r o to h ta a nd P r o toz oa i s t ru e o f th e g er m s o f a l l or g a n
p y ,
is m s u p t o the high est : the di fferenti a ti on of ou t er from i nner

i s the fi rs t st ep Wh en the p rot opl a s m o f a n A l ga c ell h a s
. -
b roken u p i nt o the clu s t ers of g ra nu l es which a re ev ent u a lly

t o bec om e s p or es ea c h of th es e qu i c kly a c qu i res a m em
,
bra nou s c oa ti n g ; con s tit u ti ng a n u nlik en es s be t ween s u r f a c e

a n d cen t r e S i m il a r ly with the o vu l e of ev ery high er pl a n t :
.
th e m a s s of c ell s f ormi ng it ea rly e xhibit s a n ou t s id e l a y er of

,
c ell s di s ti ngu i s h ed f rom the cell s withi n With a nim a l germs .

- -
it i s the s a m e B e it i n a cili a t ed g em mu l e be it i n the

.
,
u n fertili z ed ova o f A hi des a n d o f th e C eci dom i a or be it i n

p y ,
t ru e ova the p ri m a ry differenti a ti on c onf orms t o the rel a

,
71
3 86 P H Y SIOL OGICA L D E V E L OP ME NT .
ti ons of ext eri or a nd i nt eri or I f we t u rn t o a d u lt

.
org a nis m s v eg et a l o r a ni m a l we see th a t wh eth er th ey do or

, ,
-
do n ot di s pl a y oth er c ont ra s ts o f p a rt s th ey a l w a y s di s pl a y
,
this c ont ra s t T h ou gh oth erwi s e a l m os t h om ogeneou s s u ch

.
,
F u ngi a s th e p u ff b a ll or a m ong A l gee a l l which h a v e a

-
, , ,
th a llu s of a ny thickn es s p res ent m a rk ed di fferences b et ween

,
th os e of th ei r c ells which a re i n i m m edi a t e con t a ct w ith th e

envi ronm en t a nd th os e whi ch a re not S u ch di fferenc es th ey
.
p res ent i n c om m on with ev ery high er pl a n t ; whi ch h ere i n ,
the sh a p e of b a r k a nd th er e i n the s h a p e of c u ti c l e h a s a n ,
en v el ope i ncl o si n g it e v en u p t o its p et a l s a nd s t a m ens In .
lik e m a nner a m ong a nim a l s th ere is a l wa ys either a t ru e

,
ski n or a n ou t er c oa t a n a l og ou s to one Wh erever a ggreg a t es .
of th e fi r st o r d er h a v e u nit ed i n t o a gg reg a t es of th e s ec on d
a nd thi r d ord er s— wh er ev er th ey h a v e b ec om e th e m or ph o
l ogic a l u nits of s u ch high er a ggreg a t es— th e ou t erm os t of

th em h a v e gr own u nlik e th os e lyi n g withi n E v en the .
S p onge is not with ou t a l a y er th a t m a y by a na l ogy be c a ll ed

d erm a l.
T hi s l a ps e of the rel a tiv ely h om ogen eou s i nt o the rel a

ti vel y h et erogeneou s fi rs t sh owi n g its elf a s on th e hyp oth es i s
, ,
of e v ol u ti on it m u s t do by th e r is e of a n u nlik enes s b et ween

,
ou t s id e a n d i ns id e g oes on next t o sh ow it s el f a s w e i nfer

, ,
th a t it mu st do by the es t a bl i s hm ent of sec on d a ry c ont ra s ts

,
a m ong t h e ou t er p a rt s a n s weri ng t o s ec ond a ry c ont ra s t s
a m on g th e f orc e s f a lli n g on th em S o l ong a s th e wh ol e s u r

.
fa ce o f a pl a nt rem a i ns si m il a rly rel a t ed t o the envi ronment ,
a s i n a P r o toc occus it rem a i n s u nif or m ; bu t wh en th ere c om e

,
t o be a n a tt a ch e d s u rf a c e a nd a f ree su r f a c e th es e b ei n g s u b , ,
jc et to u nlik e a cti on s a re r end ered u nlik e T his is vi s ibl e

,
.
ev en i n a u nic ell u l a r A l ga wh en it b ec om es fi x ed ; it i s
s h own i n th e disti ncti on b et ween th e u n d er a nd u pp er p a rt s
o f or di na ry F u ngi ; a nd we s ee it i n th e u n iv er s a l diff eren ce
between the imb edd ed end s a nd the exp osed end s of the
high er pl a nt s . A nd th en a m ong th e l ess m a rk ed c on t ra st s
of s u rf a c e a n sw eri ng t o th e l ess m a rk ed c ont ra sts i n the i nci
S U MMA R Y OF P H Y S I OL OGICA L D E V E L OP ME NT .
3 87
d ent f orces c ome th os e be t ween the u pp er a nd u nd er sid es of

,
l ea v es ; w hich a s we ha v e s een v a ry i n d egree a s th e con

, ,
t ra s ts o f f orc es v a ry i n d egree a nd di s a pp ea r w h ere th es e c on

,
t ra s t s di s a pp ea r E qu a lly c l ea r p roo f i s fu rn i s h ed
.
by a ni m a l s th a t the origi na l u nif or mity of s u rf a c e l a p s es i nt o

,
m ultif ormity i n p rop orti on a s the a cti ons of th e envi ron m ent
,
u p on th e s u r fa ce b ec om e m u ltif or m I n a Wor m b u rrowi ng .

,
th rou gh d a m p s oil whi c h a cts equ a lly on a l l i ts sid es or i n a ,
T a nia u niform ly b a th ed by the con t ent s o f the i nt es tine it

,
i nh a bi ts the p a rt s of the i nt egu m ent do n ot a pp reci a bly

,
di ff er f rom on e a noth er ; bu t i n c rea t u res not s u r rou nd ed by

t h e s a m e a genc i es a s th ose th a t cra wl a n d th os e th a t h a v e
,
th ei r b odi e s p a rt i a lly i n cl os ed th ere a re u nlik eness es of i n ,
t egu m en t c orres p ondi ng t o un lik enes s es of the con diti on s A .
s n a il s foo t h a s a n u nd er s u rf a c e n ot u n if or m w ith the

’
e xp o s ed s u r f a c e of i ts b ody a nd thi s a g a i n i s n ot u n if or m
,
with the p rot ect ed s u r f a ce A m ong a r ti cu l a t e a ni m a l s th ere

.
i s u su a lly a di s ti n cti on b et ween th e v ent r a l a nd the d or s a l

a s pect s ; a n d i n th os e o f th e A r thr op oda w hi c h s u bj ec t th ei r
a nt eri or a n d p os t er i o r en d s t o di f feren t envi ron i ng a g enci es ,
a s do th e a n t li on a nd th e h er mit c ra b th es e be c om e s u p er
- -
,
fi ci a l l y di fferenti a t ed A n a l ogou s genera l c ont ra sts

.
oc cu r a m ong th e Ver tebra ta Fi s h es t h ou gh th ei r ou t s id es a re

.
,
u nif or m ly b a th ed by w a t er h a v e th ei r b a c ks m or e e xp os ed
,
t o li ght th a n th ei r b elli es a nd th e t wo a re c o mm only di s ti nct

,
i n col ou r Wh en it i s not th e b a ck a nd b elly which a re t hu s

.
di s s i m il a rly c on diti oned bu t the s i des a s i n the P l eur on ec ti da

, ,
’
,
t hen it i s the s id es which b ecom e c ont ra st ed ; a nd there m a y

be s ign ific a n c e i n the f a ct th a t th os e a b norm a l i n divid u a l s of
thi s ord er whi c h r ev er t t o the a nc es t ra l u n di s t or t ed typ e a nd ,
s wi m v ert ic a lly h a v e the two s id es a lik e I n s u ch higher

,
.
v erteb ra t es a s reptil es w e see rep ea t ed thi s di fferenti a ti on

,
of th e u pp er a n d u n d er su r f a c es : esp eci a lly i n th o s e of th em
whi ch lik e s na k es exp os e th ese su rfa c es t o the m ost div ers e

, ,
a c ti on s . E v en i n bi r d s a nd m a mm a l s whi ch u su a lly by ,
ra i s i n g the u n d er s u r f a c e c on sid er a bly a b ov e the g rou nd .

g rea tly dimini s h the c on tra s t be tw een i ts conditi ons a nd the

c onditi ons t o w hi c h th e u pp er s u rfa ce i s s u bj ect t h ere s till ,
r em a i n s s om e u n lik enes s Of cl ot hi n g a ns werin g t o t h e r em a i n
i ng u nlik eness b etwee n th e c onditi ons . T hu s with ,
ou t by a ny m ea n s s a yi n g th a t a l l s u ch differen ti a t i on s a re
di rectly ca u s ed by differences i n the a cti on s of in cid ent for ces ,
whi ch a s b efore s h own

,
t h ey c a nn ot be it i s c l ea r
,
t h a t m a n y of th em a re s o c a u sed I t i s cl ea r th a t p a rt s of
.
th e s u r fa c e e xp os ed t o v ery u n lik e en vi r on i ng a g enc i es b ecom e ,
v ery u n lik e ; a nd thi s i s a l l tha t need s t o be s h own .
Com pl ex a s a re t h e t ra n s f orm a ti ons o f th e i nn er p a rts

o f or g a n i s m s f r o m th e rel a tiv ely h om og eneou s i nt o th e r el a
-
ti vel y h et erogen eou s w e s till see a m on g th em a con for mi ty

,
t o the s a m e gener a l or d er I n b oth pl a n t s a nd a ni m a l s th e

.
f erenti a ti ons a ns wer t o th e s t rong er con

e a r li er i n t ern a l di f
tra s t s Of c on diti on s . P l a n t s a b s or bi ng a l l th ei r
,
nu t ri m en t th r ou gh th ei r ou t er s u r f a c es a re i n t ern a lly m odi

,
fi ed m a i nly by the t r a ns f er o f m a t er i a l s a nd by m ech a nic a l

s t r ess
. S u ch of th em a s do n ot ra i s e th ei r f r ond s a b ov e th e
su r f a c e h a v e th ei r inn er ti s s u es s u bj ect t o n o m a r k ed c on
,
t ra s ts s a v e t h os e ca u sed by cu rr en t s of s a p ; a nd th e li nes
of l en gth ened a nd o th erwi s e ch a ng ed cell s whi c h a re f o r m ed
w h ere th es e cu rren t s ru n a nd a re m os t cons pi cu ou s wh er e

,
th ese cu rrent s m u s t Obvi ou s ly be the s t ronges t a re th e only ,
d ecid ed di fferenti a ti ons of th e i n t eri or B u t w h ere a s i n

.
,
the high er Crypt og a m s a nd i n t e n og a m s the l ea v es a re ,
u ph eld ,
a nd th e s u pp orti n g s t em i s t ra ns v er s ely b en t by
t he w i nd th e i nner ti ssu es s u bj ec t t o di fferen t a m ou nt s o f

, ,
m ech a n ic a l s t ra i n di fferenti a t e a ccordi n gly : the d ep os it of

,
d ens e s u b s t ance c om m ences i n th a t regi on wh ere the s a p

c ont a i ni n g cell s a nd c a n a l s s u ffer the grea t es t i nt ermitt ent
c om p res s i ons . A nim a ls or a t l ea s t su ch of th em
,
a s t a k e f oo d i nt o th ei r i n t eri ors a r e s u bj ect t o f or ces of

,
a n oth er cl a ss t endi n g t o d es t roy th ei r or igi na l h om og eneity .
F ood i s a foreign su b s t a nce which a ct s on th e i nt eri or a s a n

en vi roni ng obj ect which t ou ch es it a ct s on the e xt eri or— is
S U MMA RY OF P H Y S IOL OGICA L D E VE L OP ME NT .
3 89
lit era lly a p orti on o f th e envi ronm en t whic h wh en swa l ,
l owed b ec om es a ca u s e O f i nt erna l diff erenti a ti ons a s the rest

,
o f th e envi r on m en t c o nt i nu e s a c a u s e o f ext ern a l differenti a
ti ons H ow es sent i a lly p a ra ll el a re th e tw o s ets of a c t i ons

.
a nd r ea cti ons w e h a v e see n impli ed by th e p ri m or di a l id entity

,
o f th e en d od er m a n d e c t od er m i n s impl e a n im a l s a nd of th e
,
s ki n a nd m u c ou s m em b r a ne in c o mpl ex 28 8 28 9 ) ,
H ere we h a v e fu r th er t o ob s erv e th a t a s f o od i s th e or igi na l

s ou r ce o f i n t ern a l di f feren ti a ti ons th es e m a y be exp ect e d t o
,
s h ow th ems e lv es fi r s t w h er e t h e i n fl u enc e Of th e f oo d i s
grea t es t ; a nd t o a pp ea r l a t er i n p r op orti on a s th e p a r ts a re
m ore rem ov ed f rom th e i nflu ence of th e f oo d T hey do this . .
I n a ni m a l s of l ow typ e th e c oa ts of th e a li m en t a ry c a vity or
,
c a na l a re m or e di fferenti a t ed th an th e ti s s u e which li es be
t ween the a li m ent a ryca na l a nd the wa ll of the b ody T hi s ti s .
s u e i n the high er C wl en ter a ta fi s a fee bly or g a n i z e d p a r en chym a

-
t ra v ersed by c a na l s li ned wi th s impl e cili a t ed cell s ; a n d i n the

l ower M ol l u s ca th e s t ru ct u res b ou ndi ng the p erivi s cera l ca vity
an d i t s ra mifyi n g s i nu ses a re s imil a rly imp er f ec t
,
F u rth er
.
,
it i s ob s erv a bl e th a t th e di fferenti a ti on o f thi s p eri vi s cera l

s a c a nd i t s s i nu s es i n t o a v a s c u l a r s y s t em p r oc eeds c ent ri
,
fu g a lly from th e r egi on wh ere th e a b s orb e d nu t ri m en t ent ers

th e m a s s o f c i rcu l a ti n g liq uid a n d wh ere this liqu id i s qu a li
,
t a ti vel y m ore u nlik e th e ti ss u es th a n it is a t th e r em ot er

p a rts of the b ody .
P hy s i ol ogic a l d ev el op m ent th en i s i niti a t ed by th a t i n

, ,
st a bility of th e h om ogen eou s whi ch w e h a v e s een t o be every

wh ere a c a u se of ev ol uti on ( F i rs t P ri n ci p l es ,
1 49
T h a t th e p a s s a ge fr om c omp a ra tiv e u n if or mity o f comp os i
ti on a nd mi nu t e st ru ct u re t o c omp a ra tiv e m u ltif or m ity i s s et ,
u p i n org a n ic a ggr eg a t e s a s i n a l l o th er a gg re g a t es by th e
, ,
n eces s a ry u n lik enesses o f th e a cti ons t o w hi c h th e p a r t s a re
s u bj ect is s h own by th e u n iv er s a l r i s e o f th e p r im a r y diff er en

,
t i a ti on i nt o th e p a rt s th a t a re un iv ers a lly m os t c on t r a s t ed i n
th ei r ci rcu mst a nc es a nd by the ri s e of s econd a ry differen
,
3 90 P HY S IOL OGI CAL D E VE L OP ME NT .
ti a ti on s
Obvi ou sl y l t d
re a e in th ei r ord er t o s ec ond a ry c ont ra sts
o f c onditi on s .
§ 312 . H ow
ph y s i ol ogi ca l d ev el op m ent h a s a l l a l ong b een
a id ed by t h e m u ltiplic a ti on of effect s— h ow ea ch di ffer en
t i a ti on h a s ev er t en d ed t o be c om e th e p a re nt Of new di fferen
t i a ti ons we h a v e h a d i n cid ent a lly v a ri ou s ill u s t ra ti on s
, , ,
L et .
u s h ere revi ew th e w or ki ng o f thi s c a us e .
A m ong pl a nt s w e see it i n the p rod u cti on of p r ogres siv ely

mu ltiplyi ng het erogenei t i es of t i ssu e by p rog res s iv e i nc rea s e
o f bu l k .Th e i n t egra ti on of f ronds i nt o a x es a nd of a x es i nt o
gr ou p s of a x es sets u p u nl i k enes s es of a cti on a m ong the
,
i nt eg ra t ed u ni t s foll owed by u nlik enes s es Of mi nu t e st rue

,
t u re E a ch g us t t r a ns v ersely s t ra i ns the v a ri ou s p a rts of the

.
s t em i n v a r i ou s d eg r ees a n d l on git u di n a lly s t ra i n s i n v a ri ou s

,
d egrees th e root s a nd whil e th ere i s i nequ a lity of s t ress a t

ev ery pl a ce i n s t em a nd b ra nc h so a t ev ery pl a ce i n st em
, ,
a n d b r a n ch th e ou t er l a y ers a nd th e s u c c es s iv ely i nner l a y ers

,
a r e s ev er a lly ext en d e d a nd com p res s ed t o u neq u a l a m ou nts ,
a n d h a v e u neq u a l m o difi ca ti ons wr ou ght i n th em L et the .
t ree a dd t o i ts p eriph ery a noth er genera ti on of the un it s

c omp os i ng it a nd imm edi a t ely the m ech a ni ca l s t ra i ns on the
,
s u pp or ti n g p a rt s a re a l l c h a n g e d i n diff erent d e grees i ni ti a t ,
i ng new di fferenc es i n t ern a lly E xt ern a lly t oo new di ffer

.
, ,
en ces a r e i n iti a t ed S h a d e d bv th e l ea f bea ri ng ou t er st ra t u m

.
-
o f s h oo ts th e i nner st ru ct u re s c ea s e t o bea r l ea v es or t o pu t
, ,
ou t s h oot s which b ea r l ea v es ; a n d in s t ea d o f th a t gr ee n c ov er
i ng whi ch th ey origi na lly h a d b ec om e c ov ered w ith b a rk o f

,
i ncrea s i ng thi ckn es s Ma nifes tly th en the l a rg er i nt egr a ti on

.
, ,
o f u ni ts th a t a re origi n a lly si m pl e a nd u nif o r m e n t a il s ,
phy s i ol ogi ca l ch a nges o f v a ri ous ord ers v a ryi ng i n th ei r ,
deg rees a t a l l p a rt s o f th e a ggr eg a t e E a ch b ra nch whi ch .

,
fa v ou ra bly c i rc u m s t a nc ed fl ou ri s h es m ore th a n i ts neigh

,
b ou rs b ecomes a c a u s e o f phy s i ol ogi ca l differenti a ti ons not

, ,
only i n i t s neighb ou rs f r om whi ch it a bst r a cts s a p a nd pres

S U MMA R Y OF P H Y SI OL OGI CAL D E V E L OP ME NT .
3 91
en tly t u rns from l ea f b ea rers i nt o f ru it b ea rers bu t a l s o i n

- -
,
the rem ot er p a rts .
T h a t a m ong a ni m a ls phy s i ol ogic a l d ev el opm en t i s fu r

th ered by the m u ltiplic a ti on of effect s we h a v e l a t ely s een ,
p rov ed by the m a ny ch a ng es i n oth er o rg a n s which the ,
g rowth or m od i fic a ti on of ea ch exc reti ng a nd s ec reti ng orga n

i niti a t es By the a b s t ra ct ed a s well a s by the a dded m a t e
.
ri a l s i t a l t er s t h e q u a lity Of th e bl oo d p a ss i ng th rou gh a l l
,
m emb ers of the b ody ; or by t he liqu id it p ou rs i nt o t h e
a li m ent a ry c a na l it a ct s on the foo d a nd th r ou gh it on t h e

, ,
bl ood a nd th rou gh it on the s y s t em a s a wh ol e : a n a ddi

,
ti ona l di fferen ti a ti on i n one p a rt thu s s etti n g u p a dditi ona l

di fferenti a ti on s i n m a ny oth er p a r t s ; f rom ea ch of which ,
a ga i n ,
s eco n d a r y di ffe r en ti a ti ng f orc e s r ev er be ra t e th r ou gh
th e org a ni s m Or to t a k e a n i nfl u ence of a noth er ord er we

.
, ,
ha v e seen h ow the m odifi ed m ec h a n i ca l a cti on of a ny m emb er

no t only m odifi es th a t m em ber but becom es by i t s rea cti ons, , ,
a ca u s e o f s ec on d a ry m odi fi ca ti on s— h ow for ex a mpl e the , ,
b u rrow i ng h a bits o f th e c om m on m ol e l ea di n g t o a n a lm ost ,
ex cl u s iv e u se o f th e f ore limb s h a v e ent a il ed a dwi n dli ng

,
o f t he hin d li m bs a nd a c on c o m it a nt d w i n dli n g o f th e
,
p elvi s w hi ch bec om i ng t oo s m a ll for the p a s s a g e of the

, ,
y oung h a s i niti a t ed still m ore a nom a l ou s m odific a ti ons

, .
S o t ha t th rou gh ou t phy s i ol ogic a l d ev el opm en t a s i n evo ,
l u ti on a t l a rg e the m u ltiplic a ti on of efl ects h a s b een a f a ct or

,
c ons t a n tly a t w ork a nd worki ng m ore a ctively a s the d ev el op

,
m en t h a s a dv a nced
. T he s ec ond a ry c h a ng es w rou ght by
.
ea c h p rim a ry ch a ng e h a v e nec es sa rily bec om e m ore nu m erou s

,
i n p r op orti on a s org a ni s m s h a v e bec om e m ore c om pl ex A nd .
ev ery i n c r ea se d m u ltipli c a ti on o f e f fec t s fu rth er di fferen ,
ti a ti ng th e org a ni s m a nd by con sequ enc e fu rth er i nt eg ra tin g

, ,
it ha s p rep a red the wa y for s till high er differenti a ti ons a nd

,
i nt egra ti on s s i m il a rly c a u s ed .
313 . Th e genera l t ru th next to be res u m ed i s th a t th es e ,
p roces ses h a ve for thei r li m it a s ta t e of eq u ilibri u m— p ro xi

3 92 P HYSIOL OGI CA L D E VE L OP ME NT .
m a t ely a m ovi ng equ ilib r i u m a nd u ltim a t ely a c ompl et e

equ ilib r i u m Th e ch a nges we h a v e c ont em pl a t ed a re bu t the
.
c oncom it a nt s of a p rogr ess i ng equ ilib ra ti on I n every a ggre .
g a t e which we c a ll l ivi ng a s well a s i n a l l o th er a gg rega t es

, ,
th e i ns t a bility o f t he h om ogeneou s i s bu t a noth er na m e for

th e a b s enc e of b a l a nc e b etween the i ncid en t f orces a n d th e
f orc es whi c h the a gg reg a t e opp os es t o th em ; a nd th e p a ss a ge
i nt o h et er ogeneity is th e p a ss a ge t owa r d s a s t a t e of b a l a nc e .
A nd t o s a y th a t i n ev ery a ggreg a t e org a ni c or oth er th er e

, ,
g oes on a mu ltipli ca ti on of effects i s bu t t o s a y th a t one p a rt

,
whi c h h a s a f resh f orce i m p r ess ed on it m u st g o on ch a ngi ng

,
a nd c om m u n ic a ti ng s ec ond a ry c h a n ges u ntil th e wh ol e of th e

,
imp ress ed force h a s b een u s ed u p i n g enera ti ng equ iv a l ent

r ea ctiv e f or ces .
Th e p ri nc ipl e th a t wh a t ev er new a cti on a n org an ism is

s u bj ect t o mu s t eith er ov erth row the m ovi ng equ ilib ri u m of
,
i ts fu ncti ons a n d c a u se t h e s u dd en equ ilib ra ti on c a ll ed d ea th ,
or e l s e m u st p r og ressiv ely a lt er th e or g a nic r hyt hm s u ntil ,
by the est a blishm ent Of a new rea cti on b a l a nci ng th e new

a cti on a n ew m ovi ng equ ilib ri u m is p ro d u ce d a ppli es a s ,
mu c h to ea c h m emb er of a n org a ni s m a s t o the org an i s m i n

i t s t ot a lity A ny f or c e f a lli ng on a ny p a rt n ot a d a pt e d t o
.
b ea r it m u s t eith er c a u s e l oc a l d es t ru cti on of tiss u e or mu s t

, , ,
with ou t d es t royi ng the ti ssu e c onti nu e t o ch a nge it u n til it

,
ca n ch a n ge it no f u r th er ; th a t i s— u n til th e m odifi ed r ea cti on
o f th e p a rt h a s b ec om e equ a l t o th e m o difi ed a cti on Wh a t .
ev er th e n a t u re of th e f orc e this m u s t h a pp en I f it is a.
m ech a n ic a l f orce th en the i m m edi a t e effect i s s om e di s t orti on

,
o f th e p a rt — a di s t orti on h a vi ng for i t s limit th a t a ttit u d e
i n which th e res i s t a n c e of the s t ru c t u r es t o fu r th er ch a nge of

p os iti on b a l a nces the f orc e t ending t o p rod u ce the fu rth er
'
c h a n g e ; a n d t he u ltim a t e effect s u pp os i ng th e f or c e t o be con

,
ti nu ou s or recu rrent i s su ch a p erm a nen t a lt era ti on of f or m

, ,
or a lt er a ti on o f s t ru ct u re or b oth a s es t a bli s h es a p er m a nen t

, ,
b a l a n c e I f th e f orc e i s phys ic o ch emic a l or ch emi ca l the

.
-
, ,
genera l resu lt is s till the s a m e : the c omp onent m ol ecu l es of

SUMMA RY OF P HYS IOL OGICA L D E VE L OP ME NT .
393
the ti s s u e m u s t h a v e th ei r mol ec u l a r a rra ngem ent s ch a ng ed ,
a nd the ch a ng e i n th ei r m o l ec u l a r a rra n ge m ent s m u st g o on
un til th ei r m o l ec u l a r m oti ons a re so re a dj u s t ed a s t o equ il i

-
b ra t e the m ol ecu l a r m oti on s of the new phy s ic o ch emic a l or -
chem ic a l a gent I n oth er wor d s th e org a nic m a tt er c om

.
,
p os i ng the p a rt if it c onti nu es t o be org a n ic m a tt er a t a l l

, ,
mu s t a s su m e tha t m ol ec u l a r c omp os iti on whi ch ena bl es it to

b ea r or a s we sa y a d a pt s it to th e i ncid ent f orc es
, , .
N or i s it l es s c ert a i n th a t th rou gh ou t th e org a n i s m a s a

wh ol e equ ilib ra ti on is a lik e the p roxi m a t e limi t of the ch a nges
,
wrou ght by ea c h a cti on a s well a s th e u lti m a t e limit of the

,
ch a ng es wrou ght by a ny rec u rren t a cti ons or c onti nu ou s

a cti on . The m o ve m ent s ev ery i ns t a nt g oi ng on a re m ov e ,
m ent s t owa r d s a new st a t e Of eq uilib ri u m Ra i s i ng a limb .
c au s es a s i m u lt a neou s s hifti ng of the c ent re of g ra vity a nd ,
s u ch a lt ered t en s i ons a nd p res su res th rou gh ou t the b ody a s

re a dj u st the di s t u r b e d b a l a nc e
-
. P a s s a ge o f liqu id i n t o or
out Of a ti s s u e impli e s s om e exc es s Of f orc e i n one di rec ti on
,
th ere a t w or k ; a nd c ea s es only wh en th e f or ce s o di m i ni s h es or
the c ou nt er f orces s o i nc rea se th a t th e e xc es s di s a pp ea r s
-
A .
nerv ou s di s ch a rg e is refl ect ed a n d r e refl ected f ro m p a rt t o

-
p a rt u ntil it h a s a l l been u sed u p i n the re a rra ngem ents pr o

,
du ced— e qu ilib ra t ed by th e r ea cti ons c a ll ed ou t A n d wha t .
is thu s ob vi ou s ly t ru e of ev ery n orm a l ch a nge i s eq u a lly t ru e ,
of ev ery a b n or m a l c h a ng e — e v ery di s t u rb a nc e of th e es t a b
l i shed rhythm of th e f u ncti ons I f s u ch di s t u rb a n ce i s a

.
si ngl e one the p ert u rb a ti ons s et u p by it rev erb er a ti ng

, ,
th rou gh ou t the s y s t em l ea v e i t s m ovi ng equ ilib ri u m s lightly

,
a lt ered I f the di s t u rb a n c e i s rep ea t ed or p ers i s t ent its s u c

.
,
c es si ve ef fect s a cc u mu l a t e u ntil th ey h a v e p rod u c ed a new

m ovi ng equ ilib r iu m a dj u s t ed t o the n ew f orc e .
E a c h re b a l a nci ng of a cti ons h a vi n g for i ts n eces s ary c on

-
,
c omit a nt a m odifi ca ti on of ti s su es it is a n obvi ou s c oroll a ry

,
th a t orga ni s ms s u bj ec t ed t o su cc es s ive ch a nges of con diti ons ,
mu s t u nd ergo s u ccess iv e di fi erenti a ti ons a nd re differenti a -
ti ons D i rect equ ilibra ti on i n org anism s with a l l i ts a ec om

.
,
3 9 4: P H YSI OL OGICA L D E VE L OP ME NT .
p a nyi ng s truct u ra l a l tera ti ons i s a s cert a i n a s i s th a t un i

,
v ers a l p rog res s t owa r ds eq u ilib r iu m of w hich it f or m s p a rt .
A nd j u s t a s c ert a i n i s th a t i ndi rec t equ ilib ra ti on i n orga ni sm s

t o whi c h the rem a i ni ng l a rge c l a s s of differenti a t i ons i s du e .
T h e d ev el op m ent of fa v ou ra bl e v a ri a ti on s by th e killi ng of
i ndivid u a l s i n whi ch t h ey do n ot occu r or a re l ea s t m a r k ed ,
i s a s b efore a b a l a n ci ng b et ween c ert a i n l oc a l st ru ct u res a nd

, ,
th e f orc es th ey a re exp osed t o ; a nd is no l ess i nevit a bl e th a n

the oth er .
314 . In al l iv er s a l l a w s we fi nd ou rs elv es a g a in
whi ch un ,
b r ou ght d own t o th e p ers i s t en ce of f orce a s the d eep es t ,
kn ow a bl e c a u s e of th ose m odific a ti on s whi ch c on stitu t e

phys i ol ogic a l d ev el op m ent ; a s it i s the d eep est k now a bl e
c a u s e o f a l l oth er e v ol u ti on H ere a s el sew h ere th e per
.
, ,
e t u a l l a p s e f rom l es s t o g r ea t er h et erog eneity t h e p erp et u a l

p ,
be getti ng of s ec ond a ry m odifi ca ti on s by ea ch p rim a ry m odi

fi ca t i on a nd th e p erp et u a l a pp roa c h t o a t emp ora ry b a l a nc e
,
on th e wa y t ow a r d s a fi n a l b a l a nc e a r e nec ess a ry impli c a

,
ti ons of the u ltim a t e fa ct th a t f orc e c a nnot di s a pp ea r bu t c an

o nly ch a n g e its f orm .
I t i s a n u nqu es ti ona bl e d ed u cti on fro m the p er s ist enc e of

f orc e th a t i n ev ery i ndividu a l org a ni s m ea ch new i ncid ent
,
f orc e mu s t work i ts equ iv a l en t Of ch a nge ; a nd th a t wh ere it

i s a c on s t a n t or recu r ren t f or c e th e limit of the ch a nge it
,
w orks m u s t be a n a d a pt a ti on of st ru c t u re s u ch as opp oses t o

the new ou t er f orc e a n equ a l i nner f or c e T h e on ly thi ng
.
op en t o qu es ti on i s w h eth er s u ch re a dj u stm en t is i nh erit

,
-
a bl e ; a n d fu rth er c on sid era ti on w ill I thi nk sh ow th a t t o

, , ,
sa y it i s n ot i nh er it a bl e i s i n di r ectly t o s a
y th a t f orc e d oes
not p ers i s t .I f a l l p a rt s of a n org a nism h a v e th ei r fu nc
ti ons c o ordi na t ed i nt o a m ovi ng equ ilib ri u m su ch th a t ev ery
-
,
p a rt p erp et u a lly i nfl u ences a l l oth er p a rt s a nd c a nnot be ,
c ha ng ed with ou t i n iti a ti ng ch a n ges i n a l l oth er p art s— i f the
li m it of ch a nge i s th e es t a bli s hm ent of a c om pl et e h a r m ony

a m on g th e m ov em ents m ol ec u l a r a nd oth er o f a l l p arts ; th en
, ,
S UMMA RY OF P H YS IOL OGI CA L D E VE L OP ME NT .
395
a m ong oth er p a rt s th a t a re m odified m ol ec u l a r ly or oth er

,
wi se mu s t be th os e which c a s t off th e ger ms of n ew orga n

,
i s m s T he m ol ecu l es o f th ei r p rodu ced ger m s mu s t t end

.
ev er t o c on f or m th e m o ti on s Of th ei r c o mp onen t s a nd th ere
,
f ore th e a rra n gem ents of th ei r c o mp onent s t o the m ol ec u l a r

,
f orc es of the orga ni s m a s a wh ol e ; a nd if thi s a gg reg a t e

o f m ol ec u l a r f orce s be m odifi ed i n its di s t rib u ti on by a l oc a l
ch a nge of s t ru ct u re t he m ol ecu l es of the g er m s mu s t be

,
gra du a lly ch a nged i n the m oti on s a nd a rra ngem ents of thei r

c omp onents u ntil th ey a re re a dj u st ed t o the a gg reg a t e of
,
-
m olec u l a r f orces
.
CH A P T E R X ‘
.
TH E I N T E GRA T IO N OF T H E O RGA N I C W O RL D .
§ 3 1 4a TH AT
f ro m the b egi nni ng of life there h a s been
.
a n ev er i nc rea s i ng h e t erog e ne ity i n th e E a r th s Fl ora a n d

’
-
F a u na i s a t ru th recogni zed by a l l bi ol ogi s t s wh o a cc ept the

,
d oct r in e of ev ol u ti on I n di s cu ss i ng th e origi n of s p eci es

.
Mr D a rwi n a n d oth ers h a v e been m a i nly occu pi ed i n ex

.
pl a i n i ng the genes i s of n ow t h i s a nd now th a t f or m of

org a n i s m c on sid ere d a s a m emb er o f on e or o th er seri es a n d
, ,
reg a r d e d a s b ec omi n g different i a t e d f r o m i t s a lli es Bu t by .
i m pli c a ti on i f not a v owedly th ere h a s b een simu lt a neou s ly

, ,
a cc ept ed t he b eli e f th a t t h e f orm s c onti nu a lly p r od u c e d by
div ergences a nd re divergenc es h a v e c ons tit u t ed a n a s s em

-
,
bl a g e i ncrea s i ngly m u ltif orm i n i ts i ncl u d ed ki nd s A nd .
thi s whi ch we a re s h own by the p r ocess of org a ni c ev ol u ti on

,
a s foll owe d ou t i n i ts d et a i l s i s a c oroll a r y f r om th e d o ct r in e

,
o f ev ol u ti on a t l a rg e a s wa s p oi nt ed ou t i n § 1 5 9 o f F i rs t
,
P ri n ci p l es .
Mea nwhil e th ere h a s b een littl e if c g iti on of a n a ny r e o n
a cc o m p a nyi ng ch a n g e n o l ess f u nd a m en t a l
,
I n th e genera l .
t ra ns form a ti on whi c h cons tit u t es E v ol u ti on di fferenti a ti on ,
a n d i nt eg ra ti on a dv a nce h a n d i n h a n d ; s o th a t a l ong w ith
the p rod u c ti on of u nlik e p a rt s t h ere p rog res ses the u ni on o f

th es e u nlik e p a r t s i nt o a wh ol e E x a mpl es o f v a ri ou s ki nds
.
b ef ore giv en will rec u r t o th e rea d er a nd a n a dditi on t o th em,
h a s j u s t b een s et f or th i n the ch a pt er on P hy s i ol ogic a l

”
I nt egra ti on One m ore ex a mpl e w orld wi d e i n i ts rea ch
.
,
-
,
h a s s till to be na m ed .
396
THE INTE GRA TI ON OF T H E OR GA NIC WORL D .
397
F or here it rem a i ns to p oi nt ou t th a t a l ong w ith the i n

c rea s i n g m u ltiplic a t i on o f typ e s o f o r g a n i s m s c ov er ing th e
E a rth s s u rfa c e th er e h a s b een ev er g oi ng on a n i ncrea s i ng

’
,
mu t u a l d ep en d ence of th em — a n i nc rea s i ng i nt egra ti on o f

the en ti re a gg reg a t e of liv in g thi ng s .
Ma ny fa ct s which a re Obvi ou s a n d m a ny which a re qu it e

fa m ili a r will be n a m ed a s e vid enc e B u t I m u s t be exc u s ed
.
for remi ndi ng t he rea d er Of thi ng s th a t he know s a n d thi ng s

th a t he m ay ea s ily ob serv e s i nc e u nl es s th e evid ence t rit e
, , ,
a s it m a y be i s g a th ered t ogeth er a n d p rop er ly m a rs h a ll ed

, ,
the genera li z a ti on enu nci a t e d will n ot be th ou ght v a lid .
§ 3 1 4 b Resp ecti ng the physi ol ogi ca l ch a ra ct ers of th e

.
ea rli es t for m s th ere i s a n a ss u m pti on f ro m w hich n o es c a p e
s ee ms p oss ible —the a s su mpti on th a t they u nit ed a ni m a l a nd

v eget a l ch a ra ct ers E v en a m ong exi s ti ng mic r os c opic typ es
.
of t he l owes t cl a s s es th ere is s u ch c om m u nity of pl a nt

,
t ra its a nd an im a l t ra it s th a t d ou bts res p ecti ng the i r p rop er

-
pl a c es i n one or the other ki n gd o m a re c o ntinu a lly ra i s ed

— d ou bt s too wh eth er if reg a r d e d a s v eg et a l th ey a re t o be
, , , ,
g rou p ed a s a lg oid or fu ngo id .
H ere h ow ev er with ou t ent eri ng on m oot qu esti on s we

, , ,
m a y d ra w th e a p r i ori concl u s i on th a t th ese ea rli es t livi ng
thi n gs were d ou bl e na t u red i n s o fa r th a t th ey m u s t h a v e

-
,
h a d th e a bility t o a s s imil a t e f rom th e i n orga n i c w orld a l l th e

m a t eri a ls of whi c h p r ot opl a s m c ons i s t s— m u s t th eref ore ,
a l ong w ith th e p ower o f a pp rop r i a ti ng ca rb o n from i ts g a s e
ou s comp ou n d a ls o h a v e h a d t h e p ower o f a pp rop ri a ti ng n i

,
t rogen eith er from one of its com bi ned oxid es or di rec tl y from
,
th e a i r with w hi ch w a t er i s m ore or l ess c h a rged F or b ef ore .
o rg a n ic s u b s t a n ces exi s t ed th er e c ou ld h a v e b een n on e bu t
i norg a ni c s ou rces f rom whi ch nit rogen c ou ld be obt a i n ed .
T h i s c oncl u s i on c oncern s u s only b ec a u se it i m pli es

h om ogen eity of na t u re i n th es e p rim or di a l for ms of life .
T h ere c ou ld n ot a t fi rs t h a v e exi s t ed a m ong th ese m i nu t es t

o f P r otoz oa ev en s u ch v a ue di s tincti on s a s a re now p res ent ed
g
398 P H Y S IOL OGI CAL D E VE L OP ME NT .
in h a d owy wa y by th ei r m odern rep resent a tiv es A nd the

a s .
impli ca ti on i s th a t d u ri ng the peri od t h rou gh ou t w hi ch these

s m a ll es t l owes t a nd s impl es t livin g thi ng s a l one exist ed
, , ,
th ere c ou ld h a v e been i n the a b senc e Of ki nds no mu t u a l

, ,
d ep end en ce .
S i nc e a m ong v a ri ou s o f t he l owes t typ es now known t o

,
us ,
the s a m e i ndivid u a l e xhibits a li f e which i s n ow pre
d omi nantly v eget a l a nd n ow p red om i na ntly a ni m a l we ,
c ann ot err i n a s su m ing th a t th ere event u a lly t ook pl a c e

di fferen ti a ti ons of thi s origi na l pl a nt a ni m a l typ e i nt o type s -
p erm a nently u nlik e : s ome i n whi c h the t ra its w ere m ore

m a rk edly v eget a l a nd oth ers i n which th ey were m ore
m a rk edly a nim a l A s f a s t a s thi s differenti a ti on a ros e
.
,
th ere c a m e the b egi nnings of c oop era ti on b et ween the pre

d omi na ntly v eg et a l typ es whi ch by the a i d Of light f or med
org a nic m a tt er f rom the in or g a n ic w orld a n d the p redom i ,
n a ntl y a nim a l typ es w hich i n chi ef m ea s u re u tili zed th e

, ,
m a tt er so f orm ed E vid ently with the ri s e of s u ch a dif

.
ferenti a ti on c a m e a n i ncipi ent m u t u a l d epend enc e I f to .
the impli ed a lg oid typ e a nd the a nim a l typ e th ere be a dd ed

the fung oid typ e s om ewh a t i nt erm edi a t e i n c h a ra ct er which
, ,
i n a l a rge p r op orti on Of c a ses liv es on th e d ec a yi ng rem na nts

of th e o th er t wo we a re f urn ish ed with a ru d e c on c epti on of
,
the p rim a ry differen ti a ti on s a nd th e a c c omp a nyi ng v a gu e

mu tu a l d ep end enc es l .
S p ec u l a ti on a s i de i t s uffi c es t o s a y th a t ea rl y i n the hi s t ory

,
of life th er e m u st h a v e a r i s en the di s ti ncti on b et wee n P r o
toz oa a nd P r o top hy ta a nd th a t thi s di s ti ncti on f oresh a d owed

,
th a t wid es t c on t ra st whi c h the high er org a nic w orld p resents

the c ont ra s t b et ween pl a nt s a nd a nim a l s I t i s n eedl es s.
t o do m ore th a n n a m e the m u t ua l d epe nd enc e b et ween th es e

two g rea t divi s i ons T h a t a s b ei ng res p ectiv ely d ec omp osers
.
,
o f c a rb on di o xid e a nd exh a l c rs o f ca rb on di oxid e th ey a ct ,
recip r oc a lly a s a l so i n s o m e m ea s u re by i n t er ch a ng e of n it ro
,
g enou s m a tt ers ; a n d th a t the i m pli ed genera l coop era ti on

s erv es in a n i ndi rect way t o u nit e th ei r liv es a nd i n th a t ,
T HE INT E GRA T I ON OF T HE ORGA NI C WORL D .
3 99
s
s en e i nt eg ra t e the two ki ngd oms ; needs not to be i ns i s t ed
to
u p on F u rth er c omplica ti ons of the mu tu a l d ep endence will
.
be m enti oned by a nd by For th e p resen t it s ufli ces to

.
r ec o gni ze thi s divi s i on o f org a nic f u nc ti ons a s t he fi rs t which
a ros e a nd a s c on ti nu i n g t o be th a t fu n d a m en t a l on e w hi c h
m ore th a n a l l others bind s o rga ni s ms a t l a rge t ogeth er .
3 1 40 . It will be
th ou gh t by m a ny rea ders th a t i n sp ea k
i ng of th e c ont ra s t ed vit a l a ctiviti es o f pl a nt s a nd a n i m a l s a s
”
con s tit u ti ng a divi s i on Of org a ni c f un cti ons I a m s t ra i ni ng
“
,
w or ds b ey on d th ei r m ea ni ng s ; s i n ce the c onc epti on of org a n i c

fu ncti ons p os t u l a t es a n orga n iz ed w h ol e i n whi ch th ey exist
’
a nd pl a nt s a nd a ni m a l s c on s tit u t e n o s u ch org a n i zed w h ol e .
Bu t th ere i s a t h a n d a n u n exp ec t ed d efen ce for thi s c oncep

ti on— a d efence n ot f orthc omi ng a gen era ti on a g o bu t whi c h ,
now a l l bi o l o gi s t s w ill re c o gni ze a s rel ev a nt I refer t o the

.
ph en om ena of s ymbi os i s T h es e p res ent v a ri ou s c a s es i n

.
whi c h the pl a nt f u ncti on a nd th e an i m a l f u n c t i on a re ca rr i e d

- -
on i n the s a m e b ody — c a ses i n whi c h th e c oo p er a ti on is not

,
be t w een s ep a ra t e v eg et a l or g a n i s m s whi c h a cc u m u l a t e n u t ri
tiv e m a tt er s a nd s ep a ra t e a n im a l orga ni s m s whi ch c onsu m e

th em bu t i s a coop era ti on be t ween v eg et a l el em en t s a nd
,
a nim a l el e m en t s f or mi ng p a rts o f th e s a m e or g a n ism .
A s i nt ro d u ct ory t o ex a mpl es of th es e mu s t fi r s t h owev er , ,
be n a m ed a n ex a mpl e of s u c h c oop er a ti on b e t w een th e two

grea t cl a s s es of v eg et a l org a ni sm s— the fung oid a nd the a l
g oi d . I n c r ed ibl e a s th e s t a t em en t on c e s ee m e d it is a s t a t e
,
m ent n ow a cc ept ed th a t wh a t w e k n ow a s lich ens a n d u s ed t o

, ,
c ons id er a s pl a nts f orm i ng a cert a i n l ow cl a s s a re now f ou nd ,
t o be n ot pl a nt s i n th e ordi na rv s ens e a t a l l bu t com p ou nd ,
growth s form e d of mi nu t e a lgae a nd mi nu te fu ngi ca rr yi n g ,
on th ei r liv es t og e th er : t h e a lg a e fu rn i s hi ng t o t h e f un gi
c ert a i n cons tit u ent s th ev need bu t c a nnot di rectly obt a i n .
a n d the fu n gi p r o fiti ng bv c ert a i n m a t er i a l s th ey obt a i n f r om
th e a lgae eith er w hil e l i vm g or whil e i ndivid u a lly d ec a yi ng

,
.
Whence it wou ld s ee m t h a t a fter the m i cr os c opic v eget a l

4 00 P H Y SI OL OGI CAL D E VE L OP ME NT .
typ e h a d b ec om e i n a l a rge d eg ree di fferen ti a t ed i nt o two

m a i n typ es i n a d a pt a ti on t o di ffer ent c o nditi on s o f lif e a nd
, ,
h a d a cqu i red a pp rop ri a t e sp ec i a li ti es of na t u re th er e grew u p ,
thi s c om m u ni s tic a rra ngem ent b et ween c ert a i n of th em en ,
a bli ng ea ch t o ben efit by th e p ow er s w hich th e o th er h a d
a c q u i re d : evid ently a n exch a ng e of s ervic es a phy s i ol ogi ca l ,
divi si on o f l a b ou r a m u tu a l d ep end en ce O f fu ncti on s a na l o

,
g ou s t o th a t which exi s ts b etween f u ncti ons i n a n ordi na ry

pl a nt or a n i m a l .
N ot di ffer i ng i n p r i ncipl e bu t only i n a pplic a ti on i s th a t ,
s ymbi os i s a b ov e ref err ed t o a s e xisti n g b e t ween P r o top hyta
a n d m a n y P r o toz oa a s w ell a s b et ween s u ch P r o top hy ta

,
a nd th e l owes t ki n d s o f M eta zoa A recent s t a t em ent th a t

.
c ert a i n a m oeb ae m a d e g reen by c onta i n ed chl orophyl l con

‘
, ,
t i nu e t o gr ow a n d m u ltiply a ft er th ey h a v e c ons u m ed wh a t
nu t ritiv e m a tt er m a y be a t h a n d i s i n h a rm ony w ith v a ri ou s
,
f a cts a ll eged of other P rotozoa — v a ri ou s oth er ki nds of Rhi zo

p ods v a ri ou s H el i oz oa nu m erou s cili a t ed a nd fl a gel l a ted
, ,
I nfu s ori a A m ong M eta z oa the lik e a ss oci a ti on oc cu r s i n

.
one o f the s p ong es i n the H ydra vi ri di s,

i n v a ri ou s t u rbe l ,
l a ri a ns i n a rotifer a nd ev en i n tw o m oll u s cs
, ,
I n th es e .
ca s e s the p a rt ners hip b et ween th e v eget a l c ell s a nd the

a n im a l c ell s ( e xi s ti n g eith er a s u nit s or a s a n org a ni zed grou p
s u ch a s a p olyp e ) i s a p a rt ners hip w hi ch a s b ef ore p rofit s

, , ,
ea ch o f th e p a r t ner s— a n i nferenc e s u pp ort e d by th e fa ct th a t
Meta z oa c ont a i nin g th es e a lg oid c ell s u s u a lly pl a ce them

s elv es wh ere the light f a ll s u p on th em a nd ca n th eref ore ,
f u rth er th e p rod u cti on of the c a rb o hyd ra t es whi ch ev ent -
n a lly b ec om e u s ef u l t o the a n i m a l c ell s whil e th es e i n s om e

-
,
wa y recip r oc a t e t h e b enefit .
H ere th en we h a v e exch a n ge of s ervic es bet ween a ss o

, ,
c i a ted pl a n t e l em ent s a n d a nim a l el e m ent s— a p er f orm a nce

- -
by th em of different orga nic func ti on s for the b enefit of the

a gg reg a t e which th ey u nit e t o form H ence wh en th es e.
,
v eget a l el em ent s a n d a nim a l el em ents a re s ep a ra t ely em

b odi ed i n pl a nts a nd a nim a ls whi ch p rofit by one a n oth er,
,
T HE I NT E GRA TI ON OF T H E ORGA NIC WORL D .
4 01
we m ay s till p rop erly rega rd th ei r re s p ec tiv e lives a s m u t u a lly

d ep end ent org a nic f uncti ons a s s a id i n the p rec edin g s ecti on
,
.
We a re ena bl ed th e be tter to s ee h ow th e E a rt h s Fl ora a nd ’
F a u na whi ch a re res p ectively a c cu m u l a t ors of m o ti on a nd

,
expe nd ers o f m oti on f orm m u t u a lly d ep en d en t p a rt s o f a

,
-
w h ol e a n d a re i n th a t s en se i nt eg r a t ed
,
A n d we s h a ll be
.
p rep a red t o see how a l l oth er rel a ti ons bet ween org a ni s m s
whi c h m a k e th em s u b servi en t on e t o a n other s i m i l a r ly con ,
s ti tu te el em en t s i n a g ener a l i n t eg ra ti on o f the or g a n ic w or ld .
§ 3 1 4d . f or m of m u t u a l d ep en denc e a ri d cons e
A noth er
qu ently of i nt egr a ti on i s c ons pic u ou s— th a t whi ch a ec om
pa ni ed the p r og res s iv e i ncrea s e of s iz e i n org a n i sm s of the
high er cl a s ses We h a ve bu t t o con t em pl a t e the p os sibiliti es
.
t o s ee th a t li fe m u s t neces s a rily h a v e c om m enced w ith m i nu t e

f orm s a nd th a t the p rog ress t o l a rger ones m u st h a v e b een
,
by s m a ll st ep s .
F or h a d c rea t u res O f a pp reci a bl e s i zes be en the fi rs t t o

a ck of
i
ex s t th ey w ou ld i nevit a bly h a v e di sa pp ea r ed f rom l
f ood H a vi ng no res ou rce bu t to d ev ou r one a nother they
.
,
wou ld q u i c kly h a v e b rou ght life to a n en d T h ere m u s t h a v e

.
been s m a ll er typ es s er vi n g a s p rey for l a rg er ones b efore

th es e c ou ld c onti nu e t o exi s t a nd t o m u ltiply : mic rob es
a ff ordi n g f o od t o i n fu sori a i nf u s ori a a f
,
f ording f ood to s u ch
si zed c rea tu res a s th e E n tom os tra ca th ese a ga i n s u pplyi ng
,
f ood to s m a ll fi s h es s u ch a s l och t rou t a nd th es e l a s t yi eldi ng

,
-
,
t o l a rger fi sh es m as s es s uffi c i ently grea t for th ei r need s : ea ch

high er gr a d e req u i ring l ower g ra d es Of a pp rop ri a t e b u lk I t .
n ee d s bu t t o a s k w h a t w ou ld be co m e of tig ers i f th ere w e re
n o m a m m a l s l a rg er th a n mic e t o s ee th a t th e a n i m a l w or ld i s
,
a li nk ed a s s embl a ge of whi c h the c onnect ed m em b er s s t a nd

,
withi n c ert a i n ra ti os of m a s s ; a nd th a t d u ri ng th e ev olu ti on

o f high er a nd l a r g er typ es th e li nki ng o f g ra d es h a s b ec om e
c l oser .
T h a t a m ong pl a nt s c ons id ere d a s a n a ggrega t e rel a ti on s

of lik e kin d th ou gh fa r l ess disti nct ones h a ve a l l a l ong
, ,
72
4 02 P H YS IOL OGIC A L D E VE L OP ME NT .
b ee n growi ng m ay be rea s ona bly c on cl u ded I n a w or l d .
p eopl ed on ly by mic roscopic typ es there c ou ld not h a v e

e xi s t ed t h e c on diti ons nee d fu l for l a r g e t ree s Gra d u a l di s .
i nt eg ra ti on o f rock su rfa c es p a rtly effect ed by phy s ic a l

-
,
a g en c i es a nd p a rtly by l ow f or ms Of pl a n t s h a d t o p rep a r e ,
the wa y for s u p eri or pl a nt s T he p rod u cti on of su ffi c i en t

.
s o il by mi nera l ogi ca l d ec a y a s w ell a s by th e d ec a y of
org a n i s m s pl a nt a nd a nim a l m ay be reg a rd ed a s h a vi ng

, ,
b een a p relimi na ry to l a rger pl a nt growth ; a nd th ou gh a t -
p res ent the d ep en d ence is far l ess cl os e th a n th a t a m on g

a n im a l s y,et t h e b en efit s yi eld ed t o m et a h
p y t es by t h e de
com p osi ng a c ti on s c a rri e d on by p rot ophyt es a s well a s th os e ,
c a rri ed on by mi crob es p erm ea ti ng the s oil imply a con ,
t i nu ed g enera l i n t erd ep end enc e th rou gh ou t th e a gg reg a t e o f

pl a nt f orms a p a rt f rom m ore s p eci a l i nt erd ep endences A n d
-
, .
then a l ong with thi s in d ebt edness of the g rea t er pl a nt s t o

the sm a ll er d u ri ng th e p r oc es s of ev ol u t i on th ere m u s t be ,
na m e d th a t i nd ebt ed nes s o f pl a nt lif e t o a n i m a l —li fe which -
Mr D a rwi n h a s s h own i n h i s b ook on the a gency of w orms

.
a s p ro d u c er s o f m ou ld .
3 1 4e . S ervic e s o f o ne
th er a nd c ons equ ent un i ons
to a no , ,
of m ore s p ec i a l ki nd s a re i nfini t ely v a ri ed a lik e w ithi n ea ch ,
ki ngd om a nd b et ween th e two ki ngd oms I refer to th os e .
seen i n p a r a s itism c om m en s a lism a nd oth er f or ms o f a s s o

, ,
ci a ti on . Whil e th ey do not c ond u c e t o u n i ons of the ki nd

thu s fa r c ons id ered th es e neverth el ess c onstit u t e i nnu m er
,
a bl e li nks w h er eby th e liv e s o f org a ni s m s pl a n t a nd a n im a l , ,
a r e ti ed t og eth er ; s om etim e s for th e a dv a n t a ge of b oth bu t
i n m os t c a s es for the b enefit of one to th e i n j u ry of th e oth er .
A m ong pl a nt s th e d egrees of d ep en d enc e a re v a ri ou s U n .
a bl e t o ra i s e th em s elv es i nt o th e a i r a nd light s om e cli m b , ,
l i k e the ivy by m odifi ed r ootl et s or spi ra lly c oil th em selves

, , ,
or h a ng by t end rils Oth ers th ere a re which g ra d u a lly

.
st ra ngl e th e t rees th ey emb ra c e or which lik e lich ens in , ,
d a mp cl im a t es fest ooni ng th e sm a ller t rees by and by c au s e

, ,
TH E I NTE GRA T ION OF T HE ORGA NIC WOR L D .
4 03
th ei r d ec a y Of hi gh er typ es of epiphyt es whi ch u s e t ree s

.
on ly t o g a i n el ev a ti on th e orc hid s m a y be i n s t a n c e d
,
A nd .
then we h a v e pl a n t s whi ch lik e th e m i s tl et oe fix th em selv es

, ,
on th e b a r k o f th ei r h os t s u tili z i ng th em p a rtly for p u rp ose s

,
Of el ev a ti on a nd p a r tly by a pp r op r i a ti on o f th ei r j u ic es A ft er .
th es e m a y be n a m ed th ose ext rem e ca ses i n whi ch th e p a ra

s i ti c pl a n t s c ea s i ng t o h a v e a ny chl or ophyll b ea rin g l ea v es
,
-
,
liv e wh olly on th e j u ic es of the i n v a d ed pl a n t s A t h om e .
th e c om m on d odd er a n d i n the t ropic s th e Ra fll es i a cece

, ,
bel ong t o this gr ou p T h ere mu st be a dd ed th e n u m er ou s

.
f orm s of mi nu t e fu n gi whi ch i n lik e m a nner th r iv e a t th e

e xp en se of th e pl a nt s th e y i n f es t I n a l l th ese ca s es th e .
int erd ep end ence i s one sid ed th ou gh a s we s h a ll p res en tly

-
, ,
see w hil e d e t r im ent a l t o one o f th e t w o c onc erned it i s n ot

, ,
a l wa y s d et r i m en t a l t o th e or g a nic w orld a s a w h ol e .
T h a t u tili z a ti on o f one by a n oth er a m ong a n im a l s whi ch

c a u s es i m m edi a t e d ea th i s fa m ili a r en ou gh i n th e rel a ti on s
,
b et w een ca rn iv ores a nd h erbiv ores A l m ost a s f a mili a r a re .
th ose s een i n p a ra s iti s m L e s s fa m ili a r a r e th o s e s een i n

.
c om m en s a l i s m a n d th e l ea s t fa m ili a r a re th o se which s h ow
u s exch a n g e o f s er vic e s A m ong th es e l a s t— th e m u t u a lly
.
benefi ci a l r el a ti on s— th a t b et w ee n the c r oc odil e a nd th e bi r d

whi c h picks p a r a sit es ou t of i t s t eeth i s a s t riki ng on e ; a nd
no l es s s o i s th a t o f the piqu e g on ffe a n A fric a n bi rd w hi c h -
,
’
pi erces the t u m ou r on a b u ffa l o s b a c k th a t i n cl os es a p a ra
sit e T h en of a n oth er kin d we h a v e th e conn exi on b et w een
.
a phid es a nd a nts : th e one p r ofiti ng by b ei ng ca rri ed t o

bett er p a st u res a nd the o th er by i ncrea s ed s a cc h a r i ne ex ore
ti on N ext c om es th e cl a s s of m es s m a t es th e c onnexi on s
.
,
bet ween s om e of which a re rel a tiv ely i nnocen t a s w it nes s ,
the S ea a n em on e which s ettl es it s elf on th e s h ell occu pi ed

- -
by a H ermit c ra b or a s wit nes s the R em ora fix ed on a

-
,
sh a rk s s ki n L es s i nnocent i s the rel a ti on u nd er which one

’
.
o f t h e tw o se i zes a sh a re o f the f oo d obt a i ne d by th e o th er ,
lik e th e a nnelid whi ch i ns i nu a t es its elf b e t ween the H er mit

c ra b a nd the wh elk s h ell it i nh a bits or lik e the s m a ll fishes
-
,
4 04, P H YS IOL OGICA L D E VE L OP ME NT .
i n h a biti ng certa in Medus ee o r th os e w hich nes tl e i n the

,
b ra n chi a l s a c of the L op hi us A fter th es e m ay be na m ed the

.
l es s i nj u ri ou s f orm s of p a ra s it es p rop er— th os e which di s ,
ti ngu i s h ed a s E pi z oa fix th em selv es on th e s ki ns of th ei r
,
h os t s p er m a nently or t em p ora r ily s u ch a s of the one ki nd

, , , ,
th e L erncea on fi s h es a n d o f th e oth er ki nd t he T i c k on
,
m a m m a l s a n d bi rd s T h en ther e c om e the oth er c l a s s o f

.
p a ra s it es m os t of the m high ly i n j u ri ou s di s ti ngu i s hed a s

, ,
E n toz oa livi ng w ithi n th e b odi es of th ei r h os t s n ow i n p a rt s

, ,
o f th ei r a li m ent a ry c a n a l s n ow on o th er O f th ei r m u c ou s
,
s u rf a c es a nd n ow i n v a r i ou s o f th ei r o r g a n s : th es e l a s t t wo
,
g rou ps b ei ng s o nu m erou s i n th ei r ki nd s th a t th ere a re

c omm only m ore sp eci es th a n one p rop er t o ea ch l a rger a ni m a l .
One s t a g e f u rth er i n the c ompli ca ti on m eet s u s i n th e p a ra

s it es u p on p a r a s it e s .
Bu t n ow t he genera l fa ct t o which th es e b ri ef i ndi c a ti ons

,
a re i n t r o d u ct ory is th a t t he u se m a d e of one org ani s m by

,
a n oth er h a s b een ev er w id eni ng a nd b ec om i n g m ore i n v olv ed .
A m ong pl a n t s u tili za ti on o f the l a rger by th e s m a ll er— O f

t rees by epiphyt es a nd p a ra sit es— m u s t h a ve a ri s en s i nc e the
ti m es wh en the l a rg er c a m e i nt o exist ence — ti m es rel a tiv ely
l a t e i n the c ou rs e of org a nic ev ol u ti on Mor eov er m os t of .
the pl a nt s whi c h u tili ze oth er s eith er by climbi ng u p th em or

,
s ettli n g th em s elv es high u

p on th ei r s t em s or s u cki ng th ei r
j u ic es a re phaenog a ms a nd the pl a nts th ey u tili ze a re a l s o
, ,
phaenog a ms ; s o th a t th es e i nnu m era b l e i nt erd ep end en ces

m u s t h a v e b een es t a bli s h ed si nc e th e ph aenog a mic typ e h as
b ec om e s o p red om i n a nt i n res p ec t o f b oth s i ze a nd ki nd .
S imil a rly a m ong a nim a l s T h ou gh th ere a re m any p a ra s it es

.
b el ongi ng lik e the T rem a t od es t o v ery l ow cl a s s es th ere a re

, , ,
m a ny whi ch b el ong t o the A r thr op oda a nd b ei ng d eg ra d ed , ,
f orm s of th a t cl a ss m u st h a v e c om e i nt o exi st ence a ft er

,
A rth r op od s of c ons id era bl e s t ru ct ure h a d b een ev olv ed .
A g a i n a l a rge p a r t o f th e a ni m a l s i n fes t e d by E pi zoa a nd

,
E n tozoa a re v ert eb ra t es— m a ny of the high es t typ es ; a nd a s

thes e a re rel a tively m od ern a l l thi s p a ra s itism m u s t be of
T HE I NT E GRA T I ON OF TH E ORGA NI C WORL D .
4 05
l a t e d a te .S o t oo of mu ch c omm ens a lism a nd m a ny

, ,
mu t u a lly benefi ci a l a s s oci a ti ons The recip roca l s ervic es of

-
.
a n t s a nd a phid es m u s t h a v e origi na t ed sin c e t he H y m en op tera
a n d H em i p tera b ec a m e es t a bli s h ed typ es a n d s i n c e t h e d a ys ,
wh en c ert a i n i n sect s of the a nt type h a d bec om e s oci a l a nd

-
,
s in c e th e d a ys w h en a phi d es h a d b ec o m e d eg ra d e d m emb er s
o f th ei r or d er : b o th d a t es be i n g r el a tiv ely r ec en t A nd s till .
m ore rec ent m u s t h a v e b een th e c omm ens a lis m b etween the

a nt s a n d t he m a ny s p eci es of oth er i n s ect s which i nha bit
th ei r nes ts .
L e a vi ng ou t rel a ti ons o f t he ki n d s j u st n a m e d it s eems ,
th a t d own f rom th os e b et ween c a rn iv ores a nd th ei r p rey t o

th os e b et ween li ce a n d thei r h ost s s u ch rel a ti ons p rofit one ,
of th e t wo s p eci es c on cerned a nd i n j u re the oth er a n d th a t ,
there the m a tt er en d s B ut it d oes not end th ere ; for th a t

.
mu ltiplic a ti on of effect s t o which p eopl e a re u s u a lly bli nd ,
b ri ng s a b ou t ch a nges w hi ch a s hi nt ed a b ov e th ou gh i nj u
, ,
r i on s t o th e i ndivid u a l a re b enefi c i a l t o th e s p eci es a nd ,
which wh en n ot b enefici a l to th e sp eci es a re oft en b enefici a l

, ,
to th e a gg reg a t e of s p eci es .
E v en wh ere a nim a ls of on e cl a s s liv e by d ev ou ri ng a nim a ls

o f a n oth er cl a s s we s ee on l ooki n g b ey on d th e i m m edi a t e
, ,
res u lt s ,
c ert a i n rem ot e resu lt s th a t a re a dv a nt a geou s In .
th e fi rs t pl a ce th e p roc es s is one by which i nf eri or i ndivid u a ls

—the l ea s t a gil e s wift st rong or s a g a ci ou s— a re pi c k e d ou t
, , ,
a n d p rev en t ed f rom l ea vi ng p ost erity a nd l oweri ng the

a v er a g e qu a lity o f th ei r ki n d A t th e s a m e tim e i n divid u a ls
.
m a d e f eebl e by i nj u r y or ol d a ge a re a m ong th os e t o be ,
kill ed and s a v ed f rom s uffer ing p rol onged p a i ns : th e evils of

d ea th by di s ea s e a nd st a rv a ti on b ei ng thu s l im it ed to th e pre
da tory a nim a ls rel a tiv ely sm a ll i n th ei r n u mb ers
,
Mea n .
whil e a ch eck i s pu t on un du e mu ltipli ca ti on Wh ere a t ra ct .
of coun t r y h a s been o v erru n by r a bbits w ea s els th rivi ng on , ,
the a b u n d a nt s u pply o f f oo d p res en tly bec om e nu m erou s

,
en ou gh t o b r i ng th e p op ul a ti on o f ra bbits wit h in m od era t e
limit s ; a nd by d oi ng t hi s b enefit not only a l l th os e ki nd s o f

pl a nts which a re b ei ng ea t en d own a n d a l l th os e oth er a ni

,
m a ls which liv e on su ch pl a nt s bu t a l so the ra bbit s them

,
s elv es ; si nc e i ncrea s i ng bey on d the m ea ns of su b s i s t en ce a

, ,
l a rge p a r t Of th e m wou ld if not killed di e of hu nger Be

, ,
.
t ween a phid es a nd l a dy bi rd s we s ee a c onnexi on of lik e

-
n a t u re : g rea t i n crea s e O f the fi rs t yi eldi ng a b u nd a nt f ood t o

l a rvae of the s ec ond endi ng a ft er a s ea son or s o i n s wa rm s o f
,
l a dy—bi rd s a nd c ons eq u ently o f th ei r l a rvae wh ereby th e

, ,
a phid e s,
i m m ens ely dimi n i sh ed c ea s e s o g rea tly t o inj u re
,
v a ri ou s pl a nts a nd the a nim a l s d epend ent on them E v en .
mi nu t e p a ra sit es by the evils th ey i nfli ct on one sp eci es

, ,
p rofit oth ers : in s t a nc e the enorm ou s d es t ru cti on of fli es

whi ch a mic ros c opic fu ngu s c a u s ed a few yea rs a g o— a
d es t ru cti on which reli ev ed not only m a n bu t a l l the a ni m a l s
which fli es i rrit a t e : o ft en s o m u ch a s t o hi nd er th em f rom
f eedi ng S u ch i ns t a nc es rem i n d u s h ow nu m erou s a re the
.
b ond s by which the liv es of org a ni s m s a re ti ed t ogeth er .
§ 3 1 4 f I h a ve res erved to the l a s t the cl ea res t and m os t

.
s t riki n g ill u s t r a ti on o f thi s p rog res si n g in t eg r a ti on th r ou gh
ou t the o r g a nic w o r ld I re fer t o the m u tu a lly be nefi ci a l

.
-
r el a ti on s es t a bli s h e d b et ween pl a nt s a nd a ni m a l s th r ou gh the
a g ency o f fl ow er s a n d i n s ec t s .
E v ery one n owa d a ys h a s b een m a d e f a mili a r w ith the pro

c es s of pl a nt fertili za ti on a nd knows th a t ( l ea vi ng ou t o f
-
,
c ons id era ti on pl a nt s fertili zed by wi nd b orne p oll en ) the -
a bility t o be a r s eed d ep ends l a rg ely on the a i d giv en by b ee s ,
b u tt erfli es a nd m oths T he exch a nge of s ervic es h a s been

,
.
g row i ng ev er m ore v a ri ous a nd c omplic a t ed d u ri ng l ong p a s t

p eri ods . We h a v e the a cqu i rem en t by fl owers of b righ t
c ol ou rs s ervi ng t o gu id e th es e i ns ects to pl a c es wh ere h oney
i s t o be f ou nd ; a nd we h a v e th ei r p er fu m es a ls o s ervi ng for,
guid a nc e T h en we h a v e the m a ny different a rra ngements

.
,
o ft en c omplic a t ed by which the visiti ng i ns ec ts a re obliged

,
t o c a rry a wa y p oll en a nd d u st with it th e s tigm a s of fl owers

on which th ey s u bs equ ently s ettl e : th u s effecting cros s
THE INTE GRA TI ON OF T H E ORGA NI C WORL D .
4 07
f ertili za ti on P a ri pas s u h a ve g one on i ns ec t dev el opm ents

.
-
m a d e p os sibl e by th es e a rra ng em ent s a nd fu rth eri ng them .
E s p ec i a lly m u s t be n a m ed th e m odific a ti on of c ert a i n H ym en

op tera i nt o h oney st ori n g be es : th e impli c a ti on b ei n g th a t
-
the ent i re economy es t a bli s h ed by th es e s oci a l i nsect s h a s

been s equ ent on the gr owt h of thi s s yst em of recip roc a l
b en efit s A nd th en j u s t inst a nci ng the d ep end enc e b et wee n
.
,
a p a rti cu l a r fl ow er h a ving a l on g t u b u l a r c oroll a a n d a p a r ,
ti cu l a r m oth h a vi ng a n a pp rop ri a t ely l ong p r ob os ci s it ,
s uffic es t o s a y th a t i nnu m er a bl e s p eci a liti es of t hi s g enera l
rel a ti on ev eryw h ere m u ltiply the li n k s by w hich t h e v eg et a l
wor ld a nd th e a ni m a l w or ld a re h ere c onnect ed T h a t th e .
ef fects of the c onnecti on s t ell l a rgely on th e p r os p erity of

b oth i s s u gges t ed by s om e i ns t a nc es Mr D a rwi n giv es a nd
,
.
,
by a s t a t em ent rec en tly m a d e i n the Unit ed S t a t es by D r L ,

. .
0 H ow a r d th a t the g rea t er f ost eri ng of b ee s w ou ld m u ch

.
,
i nc rea s e c ert a i n of the c rop s .
Bu t n ow ob s erv e th e b roa d f a ct t o which th es e few d et a ils

con c erni ng pl a nt f ertili z a t i on a re in t rod u ct ory
-
A l l th es e
.
genera l a nd s p eci a l rel a ti on s b et ween pl a nt s an d a nim a l s

h a v e a r i sen s i nc e the ph aenog a mic typ e c a m e i nt o exist enc e
h a v e i nd eed a r is en s inc e th e hi gh er m embers o f th a t typ e
, , ,
t he A ngi osp er m s h a v e a pp ea red ; for the Gym nosp erms do

,
not pl a y a ny p a rt i n this i nt er c o mm u ni on Bu t s o fa r a s we
.
c a n j u dg e o f p re s en t r es u lts of g eol ogic expl ora ti ons there ,
were n o A ngi os p er m s d u ri ng the E ozoic a nd P a l eozoic

p eri od s S O th a t this cl a s s of c onnexi ons be t ween a n im a l s
.
a nd v ege t a l s m u s t h a v e b een est a bli s h e d si n c e c a rb oniferou s
tim es— a p er i od l on g i nd eed bu t fa r s h ort er th a n th a t which

, ,
or g a n ic ev ol u ti on a t l a r g e h a s occu pi ed .
3 1 4g . I h a v e bu t j u s t t ou ch ed
s om e s a li ent p a rt s of a
on
s u bj ect imm ens e i n ext ent a nd ext rem ely i nv olved which it
, ,
wou ld t a k e a v ol u m e t o set forth a d eq u a t ely E nou gh h a s .
b een s a id h owever t o i ndic a t e the t ru th which it is the

, ,
p u rp os e of the ch a pt er t o b ri ng i nt o Vi ew a nd emph a s i ze
408 P HYSIOL OGICA L D E VE L OP ME NT .
the t ruth th a t b oth of the two grea t l a ws of ev ol u ti on a re

e x emplifi ed i n th e org a ni c w or ld a s a w h ol e a s th ey a re
,
ex emplifi e d i n ev ery or g a ni s m a nd i n a l l oth er thi ngs

, .
T h e rea d er h a s l on g s i nc e b ec ome f a mili a r with th e gene

ra l i z a t i on th a t whil e E v ol u ti on is a ch a ng e f r om th e h om o
g eneou s t o t he h et erog eneou s it is a ls o a ch a n ge f ro m the

,
i nc oherent t o th e c oh erent ; a nd thi s ch a nge f rom th e i nco

h erent t o th e coh erent h a s b een a b ov e exhibit ed a s g oing on
ev en th r ou gh ou t th a t v a s t a ss embl a g e o f org a ni s ms pl a nt
,
’
a nd a ni m a l ,
which c ov er the E a rth s s u r f a c e I n wh a t we
.
a re oblig ed t o c on c eiv e a s th e e a rli est st a g e wh en th e m os t

,
mi nu t e typ es of life a l one exist ed th e a gg reg a t e of livi ng

,
thi ng s wa s a t onc e h om ogeneou s a nd i nc oh erent I n the .
c ou rs e of ep och s imm ea su ra bl e i n d u ra ti on thi s u niform

,
a gg reg a t e of b ei ng s h a s been b ec om i n g m ore m u ltif or m .
A nd now we s ee th a t inst ea d of f or ms o f life ev erywh ere

with ou t the slight est u n i on c au s ed by mu t u a l d ep en denc e ,
th ere h a v e sl ow ly a ris en f orms of li fe a m on g whi c h mu t u a l

d ep end ences h a v e ent a il ed vit a l c onn exi ons c orres p ondi ngly
m a r k ed. A l on g with p rog res s i ng differen ti a ti on th ere h as
ev er been p rog ressing in t eg ra ti on . S O th a t we m a y rec og
ni ze s om ethi ng lik e a g rowin g life of the enti re a ggreg a t e of
org a ni s ms i n a dditi on t o the liv es of i ndivid u a l org a ni s m s
a n e xch an ge of s ervic es a m ong p a rts enh a nci ng th e life of
the wh ol e .
I n this fi na l gen era liz a ti on the l aw of E v ol u ti on is m a ni

fested und er its m ost t ra nsc end ent a l f orm .
P A RT VI .
L A WS OF MUL T I P L I C A T I ON
C H AP T E R I .
*
TH E P A C T OBs .
g a ni s m s h a v e been ev olv e d th ei r resp ectiv e

§ 31 5 . IF or ,
p ower s of m u ltipli ca ti on m u st h a v e b een d et er mi ned by

n a t u r a l c a u s es Gra nt th a t the c ou ntl ess sp eci a liti es of
.
st ru ct ur e a nd fu ncti on i n pl a nt s a nd a nim a ls h a v e a ri s en ,
from the a cti ons a nd rea cti on s b et ween th e m a nd th ei r

envi ron m ents c onti nu e d f rom genera ti on t o g ener a ti on ; a nd
,
it f oll ow s th a t f rom th es e a cti ons a nd rea cti on s h a v e a l s o

a ri s en th ose c ou ntl es s d egrees of fertility whi ch we see
a m on g th em . A s in a l l oth er resp ect s a n a d a pt a ti on Of ea ch
s p eci e s t o its c on diti on s of e xist enc e i s di r ectl i di ctly
y o r n r e
b rou ght a b ou t ; s o mu st th ere be di rectly or i ndi rectly

b rou ght a b ou t a n a d a pt a ti on of i ts rep roduc tiv e a ctivity t o
its c onditi ons of exi s t enc e .
We m a y exp ect to fi nd t oo th a t p erm a nent a nd t emp ora ry

, ,
An outl i ne of th e doct ri ne set forth i n th e fol l o wi ng ch ap te r s , wa s

o r i g nal
i l y p ubl i sh ed i n th e Wes tm i ns ter Revi ew for A p ril , 1 8 5 2 , unde r th e
t i tl e— A Theory of P op ul ati on deduc ed from the Gener a l L a w of A n i m a l
F er ti li ty ; a nd wa s sh ortl y a fterwa rds republ i sh ed wi th a p re fa tory note

s ta t i ng th at it m u st be a ccepte d I hoped a t s om e future
as a s k etch wh i ch
th e to el abora te In ne w re vi s ing a nd com pl eting i t I ha v e o m i tted a
.
,
non es s enti al p a rt of th e a rgum ent whil e I h ave e x pan de d th e rem ai nder

-
,
by a dding to th e num b er of fa c ts put i n e vi dence b y m eeting ob j ect ion s ,
wh i ch w a nt of s pa ce be fore obl i ged m e to p as s ov er a nd by dra w ing v a r iou s ,
s ec onda ry concl usi ons . The ori gi na l p ap er, wi th om i ss i ons , wi l l be found i n

A ppe ndi x A to Vol um e I of thi s work .
41 2 L A WS OF MU L T I P L ICA T ION .
di ff erenc es of f erti l ity h a v e the s a m e genera l i nt erp re ta ti on .
I f the s m a ll v a ri a t i on s Of s t ru ct u re a nd f un cti on t h a t a ri se
withi n the limits of ea ch sp eci es a re du e t o a cti ons lik e t h os e,
whi ch by th ei r l ong a c cu m u l a tin g effec t s h a v e p rod u c ed th e

,
-
,
imm en se c ont ra s ts be t ween the v a ri ou s typ es ; we m a y con

e l u d e th a t s i m il a rly
, th e a cti ons t o which ch a nges i n th e
,
ra t e o f mu ltiplic a ti on o f ea c h speci es a r e du e a l s o p rod u c e , ,
i n grea t p eri ods o f tim e the enorm ou s differen c es b et ween

,
th e ra t es of m u ltiplic a ti on of diff ere nt s p eci es .
Before i n qu i ri ng i n wh a t w a ys the r a piditi es of in crea s e

a r e a dj u s t ed t o t h e req u i rem en ts b o th t emp or a ry a n d p erm a
,
nent it will be n ee d fu l t o l ook a t th e fa ct ors

, L et u s s et .
d own fi rs t th os e which b el on g t o the envi ronm ent a nd th en ,
th os e which b el ong to th e org a nism .
§ 31 6 . E v ery
livi ng a ggreg a t e b ei ng one of which the
i nner a cti ons a re a dj u s t ed t o b a l a nc e ou t er a cti on s it foll ow s ,
th a t the m a i nt ena nce of i ts m ovi ng eq u ili b ri u m d ep end s on

its exp os u re t o th e right a m ou nts of th es e a ct i ons I ts .
m ovin g equ ilib ri u m m ay be overt u rned if one of thes e a cti ons

i s eith er t oo g rea t or t oo s m a ll i n a m ou nt ; a nd it m a y be s o
ov ert u rn ed eith er by exc ess or d ef ect of s om e i n org a nic
a g enc y i n its envi ron m en t or by exc es s or d ef ect of s om e

,
org a n ic a g en cy .
T hu s a pl a n t c onstit u ti on a lly fitt ed t o a c ert a i n w a rm th

,
a nd h u midity i s kill ed by ext rem es o f t em p era t u re a s w ell

, ,
a s by ext rem es o f d rou ght a nd m oi s t u re I t m a y d wi ndl e .
a wa y f ro m w a nt o f s oil or di e f rom the p res enc e o f t o o g rea t

,
or t oo sm a ll a q u a n tity o f s om e mi nera l s u b s t a nc e whi ch th e
s oil s u ppli es t o it I n lik e m a nner ev ery a nim a l c a n m a i n

.
,
t a i n the b a l a nce of i ts functi ons s o l ong only a s the envi ron

m ent a dd s to or d ed ucts f rom i ts h ea t a t ra t es not exceedi ng
d efi nit e limits Wa t er t oo mu s t be a cces s ibl e i n a m ou nt
.
, ,
s uffi ci en t t o c omp ens a t e l os s I f the p a rch ed a i r i s ra pidly

.
a b s t ra cti ng its l iqu id which th ere i s n o p ool or r iv er t o
rest or e its f u ncti on s c ea se ; a n d i f it i s a n a qu a tic c rea t u re

, ,
TH E F A CTORS .
41 3
d rou ght m ay ki ll it e ither by d ryin g u p i ts m ediu m or by

giving it a m edi u m i na d equ a t ely a er a t ed T hu s ea ch orga n .
i sm ,
a dj u st e d t o a c ert a i n a v era g e i n t he a c t i o ns o f i ts
i norga ni c envi ronm ent or ra ther we s h ou ld s a y a dj u s t ed t o

, , ,
cert a i n m od er a t e d evi a ti on s f rom thi s a v era g e is d es t roy ed ,
by ext rem e d evi a ti ons . S o t oo is it w ith th e envi r on

, ,
i ng orga n ic a g enc i es . A m ong pl a nt s only the p ara s itic ki nd s

,
a nd th os e u ni t e d by sy m bi o si s ( a s w ell a s a few i nn ocent

“
l odg ers d ep en d for thei r i ndividu a l p res erv a ti on on the
p res enc e of c ert a i n oth er org a ni s m s ( th ou gh th e p res ence of
c ert a i n oth er org a ni sms is needfu l t o m os t pl a nt s for the p re
s erv a ti on of t h e r a c e by a idi n g f ertili z a ti on ) H ere for the .
,
c ontinu a nc e of i ndividu a l life p a rti cu l a r org a ni s ms m u st

,
be a b sen t or n ot v ery nu m erou s —b ea s ts th a t b row se c a t er ,
pill a rs th a t d ev ou r l ea v es a phid es th a t su ck j u i ces A m ong

, .
a ni m a l s h owe v er t h e m a i nt en a n ce o f t he f u n c ti ona l b a l a n c e
, ,
i s b oth p ositiv ely a nd n eg a tiv ely d ep en d en t on the a m ou nts

of s u rrou ndi n g org a n ic a g ent s T h ere mu s t be a n a cc es s ibl e
.
s u ffi c i ency o f the pl a nts o r a nim a ls s ervi ng for f o od ; a n d of
org a n i s m s th a t a re p red a t ory or p a ra sitic or oth erwi s e det ri
m ent a l the nu mber mu st not p a ss a c ert a i n limit

, .
T hi s d ep end ence Of the m ovi ng eq u ilib ri u m i n ev ery i ndi

vidu a l orga ni s m on a n a dj u stm ent of i ts f orces t o the f orces
of the en vi r on m en t , a n d the o v ert h r ow o f thi s eq u ilib ri u m
by f a il u re of th e a dj u s tm ent i s c omp reh en s iv e of a l l c a ses

, .
A t fi rs t s ight it d oes n ot s eem t o i ncl u d e wh a t we c a ll na t u ra l

d ea th ; bu t only d ea th by vi ol ence or st a r v a ti on or c old or
, , ,
d rou ght Bu t in rea lity n a t u ra l d ea th n o l es s th a n ev ery

.
,
oth er ki n d O f d ea th is c a u s ed by th e f a il u re t o m ee t s o m e
,
ou t er a cti on by a p r op ort i ona t e i nn er a cti on Th e a pp a ren t .
difference is du e to the fa ct th a t i n ol d a g e wh en th e ,
qu a ntity of f orc e ev olv ed i n the org a ni s m gra d u a lly di m i

ni s h es the m o m ent u m o f the fu ncti ons be com es s t ep b st p
, y e
l es s a nd the v a ri a ti ons of the ext ern a l f orc es rel a tiv ely

,
grea t er ; un til th ere fin a lly c om es a n occ a si on wh en s om e

qu i te m odera t e d evi a ti on f rom th a t a v era g e to which the
41 4 L A WS OF M U L TI P L I CA T I ON .
f eebl e m ovi ng equ ilib ri u m is a dj u s t ed p rod u c es

, in it a fa t a l
p ert u rb a ti on .
§ 3 1 7 T h e i ndivid u a l s of every sp eci es bei ng thu s de

.
pe n d ent on cert a i n envi ron in g a c ti ons ; a nd s ev era lly h a vi ng

th ei r m ovi ng equ ilib ri a s ooner or l a t er ov erth rown by one or
o th er o f th ese envi roni ng a c ti on s ; w e h a v e next t o c on s id er
i n wh a t w a y s th e envi r on i ng a cti ons a re s o m et a s t o p r ev ent

e xti n cti on of t h e s p eci es T h ere a r e two es senti a lly di fferent
.
wa y s .T h ere m a y be in ea ch i n divid u a l a s m a ll or g rea t

a bility t o a dj u s t it s elf t o v a ri a ti ons of th e a g enc i e s a rou nd
it a nd t o a s m a ll or grea t nu mb er of s u ch v a ryi ng a genc i es

— th ere m a
y be littl e or m u ch p ow er of p res ervi ng th e
b a l a nce o f the fu ncti on s A nd th ere m a y be mu ch or littl e
.
p ower of p rod u ci ng new i ndividu a ls t o repl a ce th os e wh os e

m ovi ng eq u ilib ri a h a v e been ov erth rown A few f a cts m u st .
be s et d own t o enf orc e th es e a b s t ra ct s t a t em ent s .
T h ere a re b oth a ctiv e a nd p a ss iv e a d a pt a ti ons by which

o rg a n i s m s a re en a bl ed t o s u r viv e a dv ers e i n fl u enc es P l a nts .
sh ow u s bu t few a ctive a d a pt a ti ons : th a t of the P itcher pl a nt -
a nd th o s e o f the r ep ro d u ctiv e p a rt s o f s om e fl ow ers ( whi ch do
n ot h owev er c on d u c e t o s elf p res er v a ti on ) a re exc e pti ona l

, ,
-
i ns t a nc es Bu t pl a nt s h a v e v a ri ou s p a s s iv e a d a pt a ti ons ; a s
.
th orn s s ti ng i n g h a i rs p oi s on ou s a nd a c r id j u ic es rep u gn a nt
, , ,
o d ou rs a nd th e w oolli ness or t ou gh n ess th a t m a k e s th ei r

,
l ea v es u nea t a bl e A nim a l s exhibit fa r m ore nu m erou s

.
a dj u stm en t s b oth p a s s ive a nd a ctiv e I n s om e c a s es they

,
.
su r viv e d es i cca ti on th ey hyberna te th ey a cq u i re t hi ck er

, ,
cl othi ng a nd s o a re fitt ed to bea r un fa v ou ra bl e i norg a nic

,
a cti on s ; a n d th ey a re i n m a ny c a s es fitt e d p a s s iv ely t o m eet
th e a dv er s e a cti on s of oth er org a nisms by b ea ri ng spi nes or ,
a rm ou r or s h ells by s imu l a ti ng neighb ou ri ng Obj ects i n c o l ou r

,
or f or m or b oth by emitti ng dis a greea bl e od ou r s or by h a vi n g

, ,
di s gu sti ng t a s t es I n still m ore nu m erou s w a ys th ey a ctiv ely

.
c on t end with u n fa v ou ra bl e c on diti ons A ga i nst th e s ea s on s .
th ey gu ard by st ori ng u p f ood by s ec reti ng th ems elves i n ,

THE FA C TORS .
41 5
c re vices or by form i n g b u rrows a nd nes t s T h ey s a v e th em

, .
s e lv es f ro m enem i es by d evel op ed p owers of l oc om oti on ,
t a ki ng the s h a p e o f s wi ft ness or a gili ty or a ptit u d e for

c h ang in g th ei r m e di a ; by th ei r s t rength eith er a l one or
a id ed by w ea p ons ; l a s tly by t h ei r i nt elligenc e wi th ou t ,
which i nd eed th ei r oth er su p eri oriti es w ou ld a v a il th em

, ,
littl e . A nd th en th ese v a ri ou s a ctiv e p ow ers s ervi ng for

d ef enc e b ec om e i n oth er c a s es the p owers th a t ena bl e a ni
, , ,
m a l s to a ggress a nd t o p res erv e th ei r lives by the s u c cess

,
o f th ei r a ggr ess i on s .
The se c on d p roc ess by w hi ch exti n cti on i s p rev ent ed— the

f orm a ti on of new i ndi vid u a l s t o repl a c e the i ndividu a ls
d es t roy ed i s c a rri e d on a s d es c ribed i n the ch a pt er on
-
,
Genes i s by tw o m eth od s the s exu a l a nd the a s exu a l

, ,
.
P l a nts m u ltiply by s p on t a neou s fi s s i on by gemm a ti on by , ,
p roli fera ti on a nd by the ev ol u ti on of y ou ng ones f rom de

,
t a ched cell s a nd s c a l es a nd l ea v es ; a nd t h ey a l s o m u ltiply

by the c a sti ng Off of sp ores a nd sp ora ngi a a nd s ee ds I n lik e .
m a nner a m ong a nim a l s th ere a re v a r i ed ki nd s of a g a m o

,
genes is from s p ont a neou s fi s si on u p t o p a rth enogenes is a l l of

, ,
th em c ond u ci ng t o ra pi d i nc rea s e of nu mb er s ; a nd w e h a v e
the m ore fa m ili a r p roc es s of g a m ogenesis a ls o c a rri ed on ,
i n a g rea t v a ri e ty of wa y s T his f orm a ti on of new

.
i ndivid u a l s to repl a ce the ol d i s h owev er i na dequ a t ely con , , ,
cei ved if w e c on t empl a t e onl y th e nu mbe r b orn o r d e t a ch e d
on ea ch oc ca si on T h ere a re f ou r f a ct ors a l l v a ri a bl e on
.
, ,
whi ch the ra t e of m u ltiplic a ti on d ep en d s T he fi r s t is the .
a ge a t whi c h r ep rod u cti on c om m enc e s ; the s ec on d i s the

f requ ency with whi c h b rood s a re p rod u ced ; the thi rd i s th e
nu mb er c ont a i n ed i n ea ch b rood ; a n d th e f ou r th i s the
l ength of ti m e d u ri ng which the b ri ngi ng f orth of b rood s

con tin u es T h ere m u s t be t a k en i n t o a cc ou n t a fu rth er
.
el e m ent— th e a m ou n t o f a i d giv en by th e p a rent t o ea ch
ger m i n the sh a p e of s t ored u p nu trim ent c onti nu ou s f eed -

,
ing wa r mth p r ot ec ti on &c : on which a m ou nt of a i d v a ry

, , , .
,
ing b etween imm en s ely wid e limi t s d ep end s the num b er of ,

41 6 L A WS OF MU L T IP L I CA TI ON .
th e n ew i ndi vid ua l s tha t s u rviv e l ong enou gh t o repl a c e the

ol d a n d p erf orm the s a m e rep ro d u ctiv e p roc ess
,
.
T h u s reg a r di ng ev ery livi n g org a nism a s h a vi ng a m ovi ng

,
eq u ilib r i u m d ep en d en t on envi roni n g f or c es bu t ev er li a bl e ,
t o be ov erth rown by i rregu l a riti es i n th os e f or ces a nd a l wa ys

'
s o o v erth r own s oon er or l a t er ; w e s ee th a t ea ch s pe ci e s Of
or g a nis m c a n be m a in t a i ne d only by the g enera ti on o f n ew
i ndivid u a l s with a c ert a in ra pidity a nd by h elpi ng th em ,
m ore or l es s f u lly to es t a blish thei r m ovi ng equ ili b ri a .
§ 318 . S u ch th e
f a ct ors with which we a re h ere con
a re
c erned I h a v e p res en t ed th em i n a bst ra ct s h a p es for the

.
p u rp ose of s h owi ng h ow th ey a re exp res s ibl e i n genera l

t erm s of f or ce— h ow th ey s t a nd rel a t ed t o th e u ltim a t e l a ws
o f re di s t r ib u ti on of m a tt er a nd m oti on .
F or the p u rp os es of the a rgu m ent n ow to f oll ow w e m a y , ,
h owev er c onv eni ently d ea l wi th th es e f a ct ors u nd er a m ore

,
f a m ili a r g ui s e I g nori ng th ei r oth er a s p ect s we m a y cl a s s

.
,
th e f a ct or s which a ff ec t ea ch ra ce o f org a ni s ms a s f ormi n g

tw o c on flicti ng s ets On the one h a nd by wh a t we c a ll
.
,
n a t u ra l d ea th by enemi es by l a ck of f ood by a tm osph eri c

, , ,
ch anges &c the r a c e i s c ons t a ntly b ei ng d es t roy ed On th e

,
.
, .
oth er h a nd p a rtly by th e end u ra nc e th e st rength the s wif t

, , ,
ne s s a n d t h e s a g a city of its m em b ers a nd p a rt ly by th ei r

, ,
f ertility it i s cons t an tly b ei ng m a i nt a i ned T hes e c on fli cti ng

, .
s et s of f a ct ors m a y be g enera li zed a s— the forc es d es t ructiv e

of ra c e a nd th e f or ce s p res erv a tiv e o f ra c e S o g enera li zi ng .
th em l et u s a sk wh a t a re the nec es s a ry implic a ti ons

, .
CH A P T E R I I .
A P R I OR I I
PR NC PLE I .
319 . TH E mber of a speci es mu st a t a ny time be either

nu
d ec rea s i ng or s t a ti ona ry or i nc rea s in g I f gener a ti on a ft er

.
,
g enera ti on its m em b ers di e f a s t er th a n others a re b orn th e

, ,
s p ec i es m u s t dwi ndl e a nd fin a lly dis a ppe a r I f its ra t e of .
mu ltiplic a ti on is eq u a l t o its ra t e of m ort a lity there c a n be ,
no nu m er ic a l ch a ng e in it An d i f the d ed u cti ons by d ea th

.
a re fewe r th a n t h e a dditi on s by bi rth the s p eci es mu st

,
becom e m ore a bu nd a nt T h es e we m a y s a fely s et d own a s

.
n ecess iti es . T h e f or ce s d es t ru ctiv e Of r a c e m u s t be eith er

gr ea t er th a n the f orces p res er va tiv e of ra c e or equ a l t o th em , ,
o r l es s th a n th e m ; a nd th ere c a nnot bu t r es u lt th es e effects
on nu mbe r .
We a re h ere c oncerned only with ra c es th a t c onti nu e to

exi s t ; a n d m a y th er ef ore l ea v e ou t of c on sid er a ti on th o s e
i n whi ch th e d es t ruc tiv e f orc es rem a i ni ng p er m a nently i n

,
e xce s s o f t h e p res erv a tiv e f or c es c au s e exti n cti on

, P ra c .
ti ca lly t oo we m a y ex cl u d e the s t a ti ona ry c onditi on ; for th e

, ,
ch a n ces a re i nfin ity t o on e a g a i ns t the m a i nt ena nce Of a p er

m a n ent equ a lity b etween th e bi rth s a nd th e d ea ths H en ce .
,
ou r i n qu i r y res olv es its elf i nt o this — I n r a ce s th a t c on
ti nu e t o exist wh a t l a ws of nu m er i ca l v a r i a ti on resu lt f ro m
,
th es e v a ri a bl e c onflicti n g f orc es which a re resp ectiv ely de

,
s t ru ctiv e o f ra c e a nd p r es erva tiv e of r a c e ?
3 2 0 C l ea rly
. if th e f orces d es t ru ctiv e of ra e , c wh en on c e
73 41 7
41 8 L A WS OF MU L TI P L I CA TI ON .
in e xc ess ha d nothi ng t o p rev ent th em f rom rem a i ni ng in

,
exc es s t he ra c e w ou ld di s a pp ea r ; a nd cl ea r ly if th e forc e s
,
p reserva tiv e of ra ce wh en onc e i n exc es s h a d n othi ng t o

, ,
p rev ent them f rom rem a ini ng i n exc ess th e ra c e wou ld go on ,
i n crea s i ng t o i nfi nity I n th e a bs ence of a ny c omp ens a ti n g

.
a cti ons th e on ly p os s ibl e a v oid a n c e o f th es e opp os it e e xt r em es

,
w ou ld be a n u ns t a bl e e qu ilib ri u m b et ween th e c onfli c ti n g

f orc es res u lting i n a p erfectl y c on s t a nt nu mb er of the
‘
, ,
sp eci es : a s t a t e whi ch we k now d oes not exi s t a n d a ga i ns t ,
the e xi s t enc e of wh i ch th e p rob a biliti es are a s a l rea dy s a id , ,
i nfinit e . I t f oll ows th en th a t a s i n e v ery c on ti nu ou s ly

, ,
e xi s ti n g sp eci es n eith er o f th e tw o c on fli c t i n g s et s o f f orc e s

“
rem a i n s p er m a nen tly in ex ces s ; th ere m u s t be s o m e wa y o f
s t oppi n g th a t e xc es s o f the one or th e oth er which i s ev er
o cc u rr i n g .
H ow i s thi s d one ? S h ou ld a ny one a ll ege i n c onf or mity ,
wit h the Old m eth od of in t erp ret a ti on th a t th ere i s i n ea ch ,
c a s e a p r ovid enti a l i nt erp ositi on to rectify the di s t u rb ed

b a l a nc e he c ommit s hi m self to th e su pp os iti on th a t of the
,
milli on s of sp eci es i nh a biti n g the E a rt h ea ch one i s yea r ly ,
re gu l a t e d i n i t s d e gree o f f ert ility by a mi r a cl e ; si nc e i n n o
t wo y ea r s d o th e f or c es whi ch f o st er or the f or c es which ,
ch eck ea ch sp eci es rem a i n the s a m e ; a nd th erefore i n no

, , ,
two y ea rs i s th ere r equ i re d the s a m e f ertility t o b a l a nc e

the m ort a lity Few if a ny will s a y th a t God con ti nu a l l v
.
a lt er s th e r ep r od u ctiv e a ctivity Of ev ery p a ra s itic f u n gu s a n d
ev er y T a p e w or m or T ri chi na s o a s t o p rev ent its e xti ncti on

-
,
or u n d u e m u ltiplic a ti on ; whi ch th ey m u s t s a y if th ey a d op t
th e hyp oth esis of s u p ern a t u ra l a dj u st m ent A n d i n th e .
a b s en c e of this hyp oth es i s th ere rem a i ns only one o th er .
The a lt ern a tiv e p ossibility is th a t th e b a l a n ce of t he p re

,
s erva ti ve a nd d est ru ctiv e f or c es is s elf s u s t a i ni ng— i s o f th e -
ki nd di s ti ngu ish ed a s a st a bl e equ ilib ri u m : a n eq ui lib ri u m

‘
s u ch th a t a ny excess of one of the f orces a t w ork it s elf ,
g enera t es by the d evi a ti on it p rod u c es cert a i n c ou nt er forc es

, ,
-
which ev ent u a lly ou t b a l a nc e it a nd i niti a t e an Opp osit e d evi

-
,
A P R I OR ! P RINCIP L E .
41 9 .
ti on L et u s c ons id er h ow i n the c a s e be fore s u ch

’
a .
, u s, a
st a ble eq uilib ri u m mu s t be c onstit ut ed .
321 . Wh ens ea s on fa v ou ra bl e t o it or a dimi nu ti on of

a ,
c rea tu res d et rim ent a l t o it c a u s es a ny s p eci es to b ec o m e

, .
m ore nu merou s th a n u su a l a n imm edi a t e in c rea se of c ert a i n

,
d es t ru ctiv e i nfl u en ces t a k es pl a c e I f it be a pl a nt the s u p .

,
p os ed g rea t er a bu nd a nc e it self imp l i es fu ll er occu p a ti on of

h
t e pl a c es a v a il a bl e fo r g r ow th— a n occ u p a ti on w hi ch l ea v ,
i ng f ewer s u ch pl a c es a s th e m u ltiplic a ti on g oes on b ec om es ,
a ch eck on f u rth er m u ltiplic a ti on— it s elf c a u s es a g rea t er ,
m ort a lity of s eed s th a t f a il t o root them s elves A nd a ft er .

:
w a rd s i n a dditi on to thi s p a ssive res i s t a nc e t o c onti nu ed

,
i n c rea s e th ere c om es a n a ctiv e res ist a nce : th e c rea tu res which

,
th riv e a t the exp ens e of the sp eci es — th e l a rvae the bi rd s the , ,
h erbiv ores — i nc rea se t oo I f it be a n a nim a l th a t h a s gr own

.
m ore nu merou s then u nl ess by s om e exc epti ona l c oi ncid enc e

, ,
a sim u lt a n eou s a nd p r op or ti on a t e a dditi on t o t he a nim a ls or
pl a n ts s ervi ng for f ood ha s occu rred th ere mu st resu lt a ,
rel a tiv e sc a rcity o f f ood E nemi es t oo be th ey b ea sts of

.
, ,
p rey or be th ey p a ra s it es mu st qu ickly begi n t o mu ltiply

, .
H ence ea c h ki n d Of org a ni s m p r evi ou s ly exi s ti ng i n s om e

, ,
thi ng lik e its nor m a l nu mb er c a nn ot h a v e its nu mb er ra is ed

,
with ou t a ri s e of the d est ru ctive f orc es neg a tiv e a nd p ositiv e , ,
qu ickly c ommenc i ng . B oth neg a tiv e a n d p osi

tive d est ru ctive f orces m u s t a u gm ent u ntil thi s i nc rea s e of
the sp e ci es i s a rrest ed T h e c omp etiti on for pl a c es on which
.
t o g r ow if th e sp ec i es be v eget a l or for f ood if i t be a nim a l

, , ,
mu st becom e m ore i nt ens e a s the over p eopli ng of the h a bit a t -
p rogress es ; u ntil th ere i s r ea ch ed the limit a t whi ch the '
m ort a lity e qu a ls the rep rod u c ti on A n d a s at the s a m e .

,
tim e enem i es will mu ltiply with a ra pidity whi c h s oon

,
bri ng s th e m a b rea s t of the a u g m ent ed s u pply o f p rey th e ,
p os itiv e res t ra i nt th ey ex er t will h elp to b ri ng a b ou t a n

ea rl i er a rre s t o f th e e xp a ns i on th a n p ress u re of p op u l a ti on
a l one w ou ld c a u s e . One m ore i nferenc e m a y be

4 20 L A WS OF MU L TI P L ICA T ION .
d ra wn H a d the s p eci e s to m ee t no rep res si ng i nfl uenc e

.
s a v e th a t neg a tiv e one of rel a tiv ely dimi n i s h ed s p a ce or -
re l a tiv ely dimi n ish e d f oo d s u pply

-
th e ca u s e l ea di ng t o i t s
-
,
i nc rea s e might c a rry it u p t o th e li m it set by thi s a nd th ere ,
l ea v e it : i ts enl a rged nu mb er might be p er m a nent But .
the p os itiv e r ep ress i n g i n fl u en ce th a t h a s b een c a ll ed i nt o

e xi s t en c e wi ll p revent thi s
,
F or th e i n c rea s e of enem i es
.
,
co mm en ci n g a s it m u st ,a ft er th e i n c r ea s e o f the s p eci e s
, ,
a n d a dv a nci n g i n g eom et ric a l p r ogr essi on u ntil it i s it s elf
c h eck ed i n lik e m a nn er w ill end i n a n ex c es s of enemi es

, .
Wh ereu p on m u st res u lt a m or t a lity of the s p eci es grea t er

th a n its mu ltiplic a ti on— a d ecrea s e which will c onti nu e
u n til i ts h a bit a t is u n d er p eopl ed its u ndu ly nu m erou s
-
,
-
en emi es d ecim a t e d by s t a r v a ti on a n d th e d es t royi n g a g en ci es

,
red u c ed t o a mi nim u m Wh ence will foll ow a noth er i n

.
c rea s e .
T hu s a s b ef ore i ndi c a t ed ( F i rs t P ri n
,
85 th ere is .
,
h ere a s wh erev er a nt a g oni s tic f orces a re i n a cti on a n a lt er

, ,
n a t e p red o mi n a n c e of ea ch c a u s i n g a r hyth m i c a l m ov em en t
,
'
-
a rhythmic a l m o v em en t wh i ch con s t i tu tes a m ovi ng equ il i
bri u m i n th os e c a ses wh ere th e f orc es a re n ot di s s ip a t ed wi th

a pp r ec i a bl e r a pidity or a re re s u ppli ed a s f a s t a s th ey a re
,
-
di s s ip a t ed Whil e th erefore on th e one h a n d we s ee th a t

.
, , ,
th e c onti nu e d exi s t enc e of a s p ec i es n ec es sa r ily i m pli es s om e

a c ti on by whi c h th e d es t r u ctiv e a nd p res erv a tiv e f or c es a re
s elf a dj u st ed ; we s ee on th e o th er h a n d th a t s u ch a n a c ti on
-
, ,
i s a n i nevit a bl e c on s equ en c e o f th e u niv ers a l p r oc es s of

equ ilib ra ti on .
§ 3 22 . I s this th e
s ol e eq u ilib ra ti on whi ch m u st exi s t ?
C l ea rly n ot T he t emp ora ry c omp ensa ti n g a dj u s tm en t s o f
.
m u ltiplic a ti on t o m or t a lity i n ea ch s p eci es a re bu t i n t ro ,
du ct ory t o the p erm a n en t c omp ens a ti ng a dj u s tm ents of m u l

t i pl i ca ti on t o m ort a lity a m on g sp eci es i n genera l T h e a b ov e .
rea s on i ng w ou ld h o ld j u s t a s it n ow d oes w ere a l l s p eci es ,
eq u a lly p rolific a n d a l l equ a lly s h ort liv ed I t yi eld s n o -

.
A P RI OR I P RI NCI P L E .
4 2]
a n s wer i nqu i ri es— why do th ei r fertiliti es difl er s o

t o t he
’
en or m ou sly or why d o th ei r m or t a liti es di f

,
fer s o enorm ou sly ?
a nd h ow is t h e g enera l fertility a d a pt ed t o th e g ener a l m or
t a li ty i n ea ch ? T he b a l a nci ng p roc es s we h a v e c ont empl a t ed

c a n g o on on ly wit hi n m odera t e li m i ts— m u s t fa il enti re ly i n
t he a b s enc e Of a du e p rop orti on between the ordin a ry bi rth

ra t e a n d the or di na ry d ea th r a t e
. I f the rep ro d u cti on of
-
.
mic e p roceed ed a s sl owly a s the rep rod u cti on of m en mic e ,
w ou ld be exti nc t b ef ore a new g enera ti on c ou ld a ri s e : ev en

di d th ei r n a t u ra l li v es ext en d t o fi ft een or s i xt een y ea rs it ,
w ou ld still be ext rem ely imp rob a bl e th a t a ny w ou ld for s o

l ong s ur viv e a l l the d a nger s th ey a re exp os ed t o C on .
ver s el y di d o x en p rop a ga te a s fa st a s i nf u s ori a the ra c e

, ,
w ou ld di e o f st a rv a ti on i n a w ee k H enc e th e mi nor a dj u s t
.
,
m en t of v a ryin g mu ltiplic a ti on t o v a ryi ng m ort a li ty i n ea ch

sp eci es impli es s om e m a j or a dj u stm ent of a v er a ge mu ltip l ic a
,
ti on to a v era ge m ort a lity Wh a t mu st this a dj u s tm ent be ?

.
We h a v e a l rea dy s een th a t the f orc es p res erv a tiv e of ra c e

a re tw o —a bility i n ea ch m emb er o f t he r a c e t o p res erv e
it s elf a nd a bility t o p rod u ce other m emb ers— p ower t o m a i n

,
t a i n i ndi vidu a l life a nd p ower t o g enera t e the sp eci es

,
.
T h es e m u s t v a ry i n v ersely When f rom l ownes s of orga ni

.
,
za ti on t h e a bility t o c ont en d w ith ext ern a l d a n g er s i s s m a ll

, ,
th ere mu s t be g rea t fertility t o c omp ensa t e for th e c on s e

qu ent m ort a lity ; o th erw i s e the ra c e m u st di e ou t Wh en .
,
on th e c ont ra ry high end ow m ents giv e m u ch c a p a city o f

,
s elf p res erv a ti on a c orresp ondi ngly l ow d eg ree of fertility is

-
,
-
req u i sit e Giv en th e d a n gers t o be m et a s a c on s t a n t qu a n

.
tity ; th en a s th e a bility t o m ee t th em mu st be a c on s t a n t
qu a ntity t oo ; a nd a s this is m a d e u p of the two f a ct ors ,
p ower t o m a i nt a i n i ndividu a l life a nd p ower t o mu ltiply ,
th es e c a nnot do oth er th an v a ry in v er s ely : one m u st de

c rea s e a s the o th er increa s es .
I t need s bu t t o c onceiv e the resu lts of n onc onf ormi ty t o

thi s l aw to s ee th a t every sp eci es mu st eith er c onf orm t o it
,
or c ea s e t o exist S u pp os e fi rst a sp eci es wh os e i n divid u a l s

.
, , ,
h a vi ng bu t s m a ll s el f p reserv a tiv e p owers a re ra pidl y de

. '
'
-
,
s t r oy ed t o be a t th e s a me t i m e wi t h ou t rep r od u c tiv e p o w er s
,
p rop orti ona t ely gr ea t T he d efect o f fert ili ty i f ext re m e

.
, ,
wi ll res u lt i n th e d ea th of one genera ti on b ef ore a n oth er h a s

grown u p I f l es s e xt re m e it will enta il a sca rcity s u ch
.
,
th a t i n th e n ext genera ti on sex u a l con gres s wi ll be t oo i n fre

qu ent t o m a i nt a i n ev en the sm a ll nu mb er which rem a i ns ; a nd
t he ra c e will d w i n dl e with i nc rea si ng r a pidi ty I f s till l es s .
ext rem e th e c ons equ en t d egr ee of s p a rs enes s

,
whi l e n ot s o ,
grea t a s t o p rev en t a n a deq u a t e nu mber of p roc rea tive un i ons ,
wi ll be s o g rea t a s t o ren d er s p ec i a l food a bu nd a nt a nd

s pe ci a l enemi es few— w ill thu s dimi nish the d es t ru ctiv e
f orces s o mu ch th a t the s elf p reserv a tive f orc es w ill b ec om e
-
r el ati vel y g rea t : s o g rea t rel a t iv ely th a t wh en c ombi ned

, ,
wi th the s m a ll a bility to p ropa g a t e the sp eci es th ey will ,
s u ffi ce to b a l a nce the s m a ll d es t ru ctiv e f orces S u ppos e .

,
n ext a s pe ci es w h ose i n dividu a l s h a v e high p ow er s Of s elf

,
p res erv a ti on whil e th ey h a v e p ow ers of mu ltipli ca ti on mu ch

,
beyond wh a t i s n eedfu l The exces s of fertility if ext rem e

.
, ,
will c au s e s u dd en exti ncti on o f the speci es by st a rv a ti on .
I f l es s ext rem e it m u s t p rodu c e a p erm a nent i nc rea se i n

,
the nu mb er of th e s pec i es ; a nd thi s f o ll owe d by i nt enser ,
c om pet i ti on for f ood a n d a u gm ent ed nu mbe r o f enem i e s will ,
i nv olv e s u ch a n i n c rea s e of th e d a ngers t o in divid u a l life ,
th a t the g rea t self p res ervi ng p owers of the i ndividu a ls will

-
n ot be m ore th a n s uffi c i en t t o c op e with th e m T ha t i s t o .
s a y if th e fertility is rel a tiv ely t oo gr ea t th en the a bility t o

, ,
m a i n t a i n i ndivid u a l life i nevit a bly becom es s m a ll er r el a ti vel y ,
t o t he requ i rem ents ; a n d the i nv er s e p rop orti on i s thu s

e s t a bli sh ed .
S O th a t wh en f rom c omp a ri n g th e differen t st a t es o f the

,
s a m e spec i es w e g o on to comp a re th e st a t es of different

,
speci es we s ee th a t th ere i s a n a na l og ou s a dj u stm ent— a n a

,
l ogon s i n the s ens e th a t gr ea t m ort a lity i s a ss oci a t ed w ith

grea t m u ltiplic a ti on a nd s m a ll m ort a lity wi th sm a ll m ulti
,
plica ti on A nd we see th a t the u nlik eness of the c a s es con

.
A P R I OR ] P RI NCI P L E .
23
i ts merely in thi s th a t wh a t is a t emp ora ry

s s ,
re a l ti on i n th e
o ne is a p er m a nent rel a ti on i n t h e oth er .
3 23 . m oment it d oe s not c oncern u s to i nqu i re

F o r th e
wh a t i s the origi n of thi s p er m a nent rel a ti on T h a t w hich .
we h a ve now to no t e is simply th a t i n s o m e wa y or oth er

,
there mu st be est a blish ed a n i nverse p r op orti on bet ween the

p ower to s u s t a in i ndivid u a l life a nd t he p ower t o p rod u ce
new i ndi vid u a ls Wh eth er or not thi s p erm a nent rel a ti on
.
‘
‘
i s self a dj u s tin g i n l on g p eri ods of tim e a s th e t emp ora ry

-
,
rel a ti on i s s elf a dj u s ti n g i n s h ort p e r i o d s of tim e

-
is a ,
s ep a ra t e qu es ti on The p u rp os e of thi s ch a pt er i s f ul fill ed by

.
sh owi ng th a t su ch a pe rm a nent rel a ti on m u s t exi s t .
B u t h a vin g rec ogn i zed the a p ri ori p r incipl e th a t i n ra c es

whi ch c onti nu ou s ly s u rv iv e the f orces d es t ru ctive of ra c e
,
m u s t be equ ilib ra t ed by th e f orc es p res erv a tiv e of ra ce ; a nd

tha t s u pp osi n g th es e a re c onst a nt th ere m u s t be a n i nvers e
, ,
p r op orti on b etween s elf p re s erv a ti on a nd ra ce p res erv a ti on ;

- -
we m a y g o on t o i nqu i re h ow thi s rel a ti on nec ess a ry i n ,
theory a ri s es i n f a ct L e a vi ng ou t the u nt ena bl e hyp oth es is

, .
o f a s u pe rn a t u r a l p re a dj u s tm ent-
we h a v e t o a sk i n wh a t
,
w a y a n a dj u s tm ent c om e s a b ou t a s a resu lt of E v ol u ti on .
I s it du e t o the s u r viv a l of v a ri eti es i n w hich the p rop orti on

of fertility t o m ort a lity h a pp e ns t o be the b es t ? Or is th e
fertility a d a pt ed t o the m ort a lity i n a m ore di rect way ? T o
thes e qu esti ons l et u s now a dd ress ou rs elves .
OBVE RS E A P R I OR I I
PR NC PLE I .
3 24 . WH E N d ea li ng with its ph enom ena i nd u ctively ,

we
saw th a t h owev er it m a y be c a rri ed on Genes i s is a p roc es s ,
“
of neg a tiv e or p o sitiv e disi nt eg ra ti on ; a nd is th u s e ss enti a lly
Opp os ed to th a t p roc ess of i nt egra ti on whi ch i s the p rim a ry

”
p roces s i n i ndividu a l ev ol u ti on E a ch n ew i ndi vi
.
d ua l wh eth er s ep a ra t ed a s a ger m or i n s om e m ore d ev el op ed

,
-
f orm i s a d ed u cti on f rom the m a s s of a pre existi ng i ndi vi

,
-
d ua l or of two pre existi ng i ndivid u a l s Wh a t ev er nu t ritiv e

-
.
m a tt er is st ored u p a l ong with the germ if it be d ep os it e d i n ,
th e s h a p e of a n egg is so m u ch nu t ritiv e m a tt er l ost t o th e

,
p a r ent N o d rop of bl ood ca n be a bs orb ed by the foetu s n or

.
,
a ny d ra u ght of mi l k s u ck e d by the y ou n g wh en b orn w ith ou t ,
t a ki ng f rom the m oth er tissu e f ormi ng a nd f orce ev olvi ng - -
m a t eri a l s t o a n equ iva l ent a m oun t A nd a l l s u b sequ ent su p .
pli es gIven to p rogeny i f th ey a re nu rt u red i nv olv e t o a

, , ,
p a rent or p a rent s s o mu ch w a st e i n ex erti on whi ch d oes not

"
b ri ng its ret u rn i n a ssi m il a t ed f ood .
C onv ers ely the c onti nu ed a ggr eg a ti on of m a t eri a l s i n t o

,
one or g a nism ren d ers imp os s ibl e the f o rm a ti on of o th er or

,
g a n i s m s ou t o f th os e m a t eri a ls A s m u ch a s s imil a t ed f oo d
.
a s is u n it ed i nt o a singl e wh ol e is so mu ch a ssimil a t ed f oo d
,
w ithh eld f rom a pl u ra lity of wh ol es which might e ls e h a v e

b een p rodu c ed Given the a bs orb ed nu t rim ent a s a c ons t a nt
.
qu a ntity a nd the l onger the b u ilding of it u p i nt o a con

,
cret e sh a p e g oes on the l on ger m u st be p ostp oned a ny b u ild

,
OB V E RS E A P R I OR I P RINCI P L E .
4 25
i ng of it u p i nt o disc ret e sh a p es A nd s imil a rly the l a rger .

, ,
th e p rop orti on of m a tt er c ons u m ed i n th e fu nc ti ona l a cti on s

o f p a rents th e s m a ll er m u s t be t he p rop orti on of m a tt er
,
whi ch ca n rem a i n t o es t a blish a nd s upp ort the f uncti on a l

a cti ons of offsp ri n g .
T h ou gh t he nec es s ity of th es e u n iv ers a l rel a ti ons is t ol er

a bly o bvi ou s a s thu s st a t e d g enera lly i t will be u s efu l t o ,
d well for a b ri ef sp a c e on thei r l ea di ng a s p ects .
3 25 . T ha t
di s i nt egra ti on w hich c ons tit u t es g enes is m a y ,
be s u ch a s t o di s p er s e enti rely the a ggr ega t e whi c h i nt eg ra

ti on ha s p revi ou s ly p rod uced— the p a rent m a y di ss olv e wh olly
i nt o p rogeny T hi s di ss ol u ti on of ea ch a gg reg a t e in t o tw o
.
or m a n y a gg reg a t es m a y occ u r a t v ery s h ort i nt er v a ls i n

, ,
w hich c a se th e b u lk a tt a i ned ca n be bu t ext re mely s m a ll ; or

it m a y oc cu r a t l onger int erv a l s i n which c a s e a l a rger b u lk ,
m a y be a tt a i ned .
I ns t ea d of qu i ckly l osi ng its own i ndividu a lity in the

i ndivid u a liti es o f i t s offs p ri ng ea ch m ember of th e ra c e m a y , ,
a ft er gr ow i n g for a ti m e h a v e p or ti ons o f i t s s u b s t a n c e be gi n
,
t o d ev el op i nt o t h e p a rent a l s h a p e a n d p res ently d et a c h

th em selv es ; a nd the p a ren t m a i nt a i ni ng its own id entity , ,
m a y c on ti n u e i n d efi n it ely s o t o p rod u c e y ou n g ones Bu t .
c l ea rly ,
t he ea rli er it comm enc es d oi n g thi s a nd the m ore ,
ra pidly it d oe s it th e s oon er mu s t the i nc rea s e of its own

,
b u lk be s t opp ed .
Or a g a i n gr owth a n d d ev el opm ent c onti nu in g for a l ong

,
p eri o d with ou t a ny d edu cti on of m a t eri a l s a n i n dividu a l of ,
c ons id era bl e si ze a nd org ani za ti on m a y res u lt ; a nd th en th e

a b s t ra c ti on of s u bst a n c e for th e f or m a ti on Of n ew i n divid u a ls
,
or r a th er the eggs of th em m a
y be s o grea t th a t a s s oon a s
,
th e eggs a re l a id th e p a rent di es of exh a u s ti on— di es th a t is , ,
f rom a n exc es siv e l oss of th e nu t ritive m a tt er s need ed for its

*
o w n a ctiviti e s .
I was h e re th i nki ng onl y of the ca s es whi ch a re g enera l a m ong i ns ect s,

but i t s eem s th a t v ert ebrat e a ni m al s , too , furni s h c as es . Mr Cunni ngham
.
wri te s “
Th ere i s a curi ous i nsta nc e of t hi s i n th e Conger : th e fem al e
4 26 L A WS OF MUL TI P L ICA TI ON .
On c e m ore the d ed u c ti on o f m a t eri a l s for the p rop a g a ti on

,
o f th e s p eci es m a y be p os tp on ed l on g enou gh t o a ll ow of grea t
b u lk a nd c omple x s t ru ct u re bei ng a tt a i ned The p rocrea tiv e .
s u bt ra cti on th en setti ng i n whi l e it check s a nd p res ently ,
st op s g rowth m a y be s o m od era t e a s t o l ea v e vit a l c a pit a l

,
s uffi ci ent t o c a rry on t he a ctiviti e s of th e p a rent ; m a y g o

“
“
on a s l ong a s p a rent a l vig ou r s u ffic e s t o f u rn ish with ou t f a t a l ,
res u lt t he m a teri a ls need ed t o p rod u c e y ou ng on es ; a n d m a y

,
c ea s e when s u ch a su rpl u s c a nnot be s u ppli ed l ea vi ng the ,
p a renta l life t o c onti nu e .
3 26 . pp os it e side of this a nt a gonism h a s a l s o

T he o
s ev era l a s p ects P rog ress of org a nic ev ol u ti on m a y be s h own

.
i n i nc rea s ed b u lk i n i n c r ea s ed st ru ct u re i n i ncrea se d a m ou nt
, ,
or v a r i e ty of a cti on o r i n c ombi n a ti ons o f th es e ; a nd u nd er

,
a ny Of its f orms this c a r ryi n g high er of ea ch i ndivid u a lity ,
i m pli es a c orrel a tiv e ret a r d a ti on in the est a bli s hm ent of new

i ndivi du a liti es .
Oth er thi ngs equ a l ev ery n orm a l a dditi on t o the b u lk of

,
a n org a ni s m is a n a u gm ent a ti on o f its li fe

*
Bes id es b ei ng a n .
a dv a nc e i n i nt eg ra ti on it impli es a g rea t er t ot a l of a ctiviti e s

,
g one th rou gh i n the a s s imil a ti on of m a t eri a ls ; a nd it i mpli es ,
therea ft er a grea t er t ot a l of th e vit a l ch a nges t a ki ng pl a ce

,
f rom m om en t to m oment i n a l l p a rts of the enl a rged m a s s .
Moreov er whil e i n c rea s ed s i ze is thu s i n s o fa r the exp res

, , ,
s i on Of i nc rea s ed life it i s a l s o wh ere the or g a nism i s a ctiv e

, , ,
th e exp res s i on of i nc rea s ed a bility t o m a i nt a in life— in c rea s ed

s t rength A gg rega ti on of s u bst a nc e is a l m ost the only m od e
.
i n which s elf p res erving p ower is sh own a m ong the l owest

-
typ es ; and even a m ong the high es t s u s t a i ni ng the b ody i n ,
i ts i nt eg rity is th a t in which s elf p res erv a ti on fund a m ent a lly -
c ons i s t s— is the end which the wid est i nt elligenc e i s i ndi

grows to 6 or 7 feet l ong and a weight of 6 0 l bs . a nd u pwa rds a nd th e n
c ea s e s to fee d for 6 m onth s wh i l e t he eggs de v el op, a nd wh en th e e ggs a re
”
s h ed di es .
I s ay norm al for the purpos e x

of e cl udi ng not onl y m orbid growth s
bu t x
e ces s of fa t .
OB V E RS E A P R I OR ] P RINCI P L E .
4 27
rec tl y m a d e t o s u bs erv e Whil e on th e one h a nd the i n

.
, ,
c re a s e of ti s su e c ons tit u ti ng growth i s c ons erv a tiv e b oth i n

es s en ce a n d i n res u lt ; on the o th er h a n d d ec r ea s e O f ti s s u e , ,
e ith er fr o m i n j u ry di s ea se or Old a g e i s i n b oth es s en ce a n d

, , ,
re s u lt t h e re v ers e A nd if so ev ery a dditi on t o i ndi vid u a l

.
,
life thu s i m pli ed nec es s a rily d el a y s or dimini s h es the c a s t i ng

,
of f of m a tt er t o f orm new i ndivid u a ls .
Oth er thi ng s eq u a l t oo a g rea t er d e gree o f org a niz a ti on

, ,
i nv olves a sm a ll er d eg ree of th a t di sorg a ni za ti on s h own by

the s ep a ra ti on of r ep ro d u ctiv e gem m m a n d germ s D et a ch .
m ent of a livi ng p or ti on or p orti ons f rom wh a t wa s p revi ou s ly

a livi ng w h ol e i s a cea sing o f c o or di n a ti o n ; a n d i s th ere f ore
,
-
es s e n ti a lly a t v a r i a nc e with th a t e s t a bli s hm en t of g rea t er c o
o rdi n a ti on which i s a chi ev e d by st ru ct u ra l d ev el op m en t In .
th e ext rem e ca s es w h ere a livi ng m a ss i s c onti nu a lly dividi ng

a nd s u bdividi ng it i s m a n ifes t th a t th er e c a nnot a r i s e m u c h
,
phy s i ol ogi ca l divi s i on of l a b ou r ; s inc e p rog res s t ow a rd s

mu t u a l d ep en d ence of p a rt s i s p revent ed by the p a rt s be
c o mi n g i n d ep en d en t C ont ra r i wi se it i s eq u a lly cl ea r th a t
.
,
i n p rop ort i on a s th e phy s i ol ogi ca l divi s i on of l a b ou r is c a rri ed

fa r t he s ep a ra tiv e p roces s m u s t be l oc a li zed i n s om e c om
,
p a r a t i ve l y s m a ll p o rti on o f t h e o r g a n i s m where it m a y g o ,
on with ou t a ffect i n g th e g enera l st ru ct u re— m u s t be c om e
rel a tiv ely s u b ordin a t e T he a dv a nc e th a t is sh own by

.
g rea t er h et erogeneity mu s t be a hi nd ra nce t o mu ltiplic a ti on

,
i n a noth er wa y . F or orga ni za ti on ent a il s c os t T h a t t ra ns .
fer a nd t r a ns form a ti on of m a t eri a l s impli ed by differenti a

ti on ca n be effect ed only by expe ndi t u re of force ; a nd thi s
,
s u pp oses c ons u m pti on of digest ed a nd a b s orb ed f ood w hich ,
might oth erwi se h a v e g one t o m a k e new org a n i s m s or the ,
g erm s of them . H ence th a t i n divid u a l evol u ti on which

,
c on s ist s i n p rogr es s ive di fferenti a ti on a s w ell a s th a t which ,
c on si s t s i n p r ogres s iv e i n t egra ti on n ec es s a rily dimi ni s h es

,
th a t Sp ec i es Of di ss olu ti on genera l or l oc a l which p r op a ga

, ,
ti on of the ra ce exhibit s .
I n a ctive org a nisms we h a v e yet a fu rth er opp ositi on

428 L A WS OF M UL T IP L I C A T I ON .
be t ween s elf m a i nt en a nc e a nd m a i n t en a nc e o f the ra c e A l l

-
.
m oti on s ens ible a n d i ns en s ibl e g enera t ed by a n a ni m a l for

, ,
th e p res erv a ti on of i t s li fe i s m oti on l ibe ra t ed f r o m decom

,
p ose d nu t ri m ent— n u t ri m en t w hi ch i f n ot thu s d ec om p os ed , ,
w ou ld h a v e been a v a il a bl e for r ep r od u c ti on ; or ra th er
might h a v e b een r epl a ced by nu t ri m ent fitt ed for rep rod u c tiv e
p u rp os es a b s orb ed f rom other kin d s of f ood H ence i n p ro
, .
,
p ort i on a s th e a c tiviti es i n crea s e— i n p rop orti on a s by its ,
m ore v a ri ed c om pl ex ra pid a n d vigorou s a cti ons a n a nim a l

, , , ,
g a i ns p ow er t o s u pp ort it s elf an d t o c op e with s u rr ou n ding

d a ngers it mu s t l os e p ow er t o p rop a g a t e
, .
§ 3 27 . H ow
thi s a nt a g oni s m be b es t exp res sed i n a
m ay
b ri ef w a y ? I f s elf p r eserv a ti on di s pl a yed it s el f i n the

-
high es t org a ni s m s a s it d oes i n th e l owes t i n littl e els e bu t

, ,
c ont i nu ou s g rowt h ; a n d i f r a ce p res erv a ti on c ons i s t ed a l wa y s

-
,
a s it d oes Oft en o f nothin g b ey on d d et a ch m en t of p orti on s

,
f rom th e p a rent a l m a s s ; then the a nt a g oni s m w ou ld be ,
th r ou gh ou t th e Obvi ou s ly neces s a ry one of in t eg ra ti on a nd

,
-
di s i nt egr a ti on Ma int ena nce of th e in divid u a l a nd p rop a g a

.
ti on of the s p eci es bei ng res p ec tiv ely a ggr eg a tiv e a n d s ep a

,
r a t i ve it w ou ld be a s s el f evid en t th a t th ey v a ry i nv ers ely

,
-
,
a s it i s s el f evid en t th a t a dditi on a n d s u bt ra c ti on u nd o one

-
a n oth er B ut th ou gh t he s i m pl es t typ es s h ow u s the o pp os i

.
ti on of s el f m a in t ena nce a nd ra ce m a i nt ena nce a l m os t wh olly

- -
u nd er thi s f orm ; a n d th ou gh high er typ es u p t o th e m os t ,
c om pl ex exhibit it t o a g rea t ext ent u nd er thi s f or m ; yet a s

, ,
we h a v e j u s t s een this i s n ot its only f orm,

T he t ot a l .
m a t eri a l m onop ol i zed by th e i ndi vidu a l a nd wi thh eld f rom

t h e ra ce m u st be st a t ed a s th e qu a ntity u nit ed t o f or m its
,
f a b ric p l us th e q u a ntity exp end ed i n diff ere n ti a ti n g i ts

,
f a b ric p l us th e qu a n tity exp end ed i n its s elf c ons ervi ng

,
-
a c ti ons . S imil a rly th e t ot a l m a t eri a l d ev ot ed t o th e r a ce a t

,
'
th e expen s e O f th e i n divid u a l i n cl u d es th a t whi ch i s di rec t ly

,
s u bt ra ct e d f r om th e p a rent i n th e s h a p e o f egg or f oet u s p l u s ,
th at which i s di rectl y s ubt ra ct ed i n the sh a p e of milk p l us

'
,
OB VE RS E A P R I ORI P RINCI P L E .
4 29
th a t which i s i ndi rectly s u bt ra ct ed i n the s h a p e of m a tt er

c onsu m ed i n ex erti ons for f os t eri ng the y oung H enc e .
this i nv ers e v a ri a ti on i s n ot exp res s ibl e i n s impl e t er ms of

a ggr eg a ti on a n d sep a ra ti on A s we a dv a nc e t o m ore highly
.
e v o lv ed org a n i s m s th e t ot a l c os t Of a n i n divid u a l b ec om es
,
v ery mu ch g rea t er th a n i s impli ed by the a m ou nt Of ti s s u e

c om p os i ng it S o too the t ot a l c os t of p rodu ci ng ea c h new
.
, ,
i n divid u a l b ecom es v ery mu ch gr ea t er th a n th a t of i ts m ere

su b s t a n c e. A n d it i s bet ween th es e t wo t ot a l c osts th a t the
a n t a g on i s m exi s t s .
We m ay i nd eed red u ce the a nt a goni s m t o a f orm com

, ,
p reh ens iv e of a l l c a s es if we c ons id er it a s exi s ti ng bet w een

,
th e s u ms of the f orces l a t ent a nd a ctiv e u s ed for t he two

, ,
p u rp os es T he m ol ec u l es whi ch m a k e u p a pl a nt or a ni m a l
.
,
h a v e been f orm ed by the a b s orpti on of f orces di rectly or

i n di rectly d er ived f rom the S u n ; an d h enc e th e qu a ntity of
m a tt er r a i s ed t o th e f or m c a ll ed org a n i c which a pl ant or ,
a n i m a l p r es en t s i s equ iv a l en t t o a c ert a i n a m ou nt of f orc e

, .
A no th er a m ou nt of for c e i s exp res s ed by th e t ot a lity of its

di fferenti a ti ons A fu r th er a m oun t of f or ce i s th a t di s s ip a t ed
.
i n i t s a cti ons A n d in th es e th ree a m ou n ts a dd ed t og eth er

.
,
w e h a v e the wh ol e exp en s e of the i ndividu a l life S o t oo .

, ,
t he wh ol e exp ense of es t a bli s hi ng ea ch n ew i ndi vid ua l i n

e lu d es— fi r st th e f or c es l a t ent i n th e s u b s t a n c e c o mp os i ng
it when b orn or h a t ch ed ; s ec on d the f orc es l a t en t i n the

p rep a red nu t rim ent a ft erwa r d s s u ppli ed ; a n d thi r d the f or ces
e xp en d ed i n fe edi n g a n d p r ot ecti n g it T h es e two s ets of
.
f orces b ei ng t a ken f rom a c omm on f u nd it i s m a nifes t th a t ,
ei t h er s et c a n in crea s e on ly by d ec rea s e o f th e oth er I f of .

,
th e f or ce whi c h th e p a r en t obt a i ns fr o m th e envi ronm ent ,
m u ch is c onsu m ed i n i ts own li fe littl e rem a i n s t o be c on ,
s u m ed i n p ro d u ci n g o th er liv es ; a n d c on v ers ely if th ere i s

, ,
a gr ea t con s u m pti on i n p r od u c i n g o th er liv e s it c a n on ly be ,
wh ere c om p a ra tiv ely littl e i s res er v ed for p a ren t a l life .
H ence th en I ndivid u a ti on a nd Genes is a re nec essa rily

, ,
a nt a g oni s tic Grou pi ng u nd er the w or d I ndividu a ti on a l l

.
4 30 L A WS OF MUL TI P L I CA TION .
p roc es ses by which i ndivi du a l life i s c ompl et ed and m a i n

'
ta i ned ; a nd enl a rgi ng the m ea ni ng Of the w or d Genes is s o

'
as t o i nclu d e a l l p roc ess es a idi ng th e f orm a ti on a nd p erf ectin g

o f new i ndivid u a l s ; we s ee th a t t h e t wo a re fun d a m ent a lly
o pp o s ed .A s su min g oth er thi ngs t o re m a i n th e s a m e

a s su mi ng th a t envi ron i ng c onditi on s a s t o clim a t e f ood , ,
e nemi es &c , c onti nu e c ons t a nt ; th en i nevit a bly ev ery

.
, , ,
hi gh er d egree o f i n dividu a l ev ol u ti on is f oll owed by a l ower

deg ree of ra c e m u ltipli c a ti on a n d vi ce vers a
-
P rogress i n
, .
bu lk c om pl exity or a ctivity i nv olv es ret rog ress i n fertility ;

, , ,
a n d p ro gres s i n f ertili ty i n v olv es ret r ogres s i n b u lk c om ,
p l exi ty or a ctivity
, .
T hi s s t a t em ent need s a s light qu a lifi ca ti on F or r ea s on s .
t o be h erea ft er a ss igned t he rel a ti on d es crib ed i s nev er c o m

,
p l e t el y m a i n t a i n e d ; a n d i n the s m a ll d ep a rt u re f rom it we ,
sh a ll fin d a rem a rk a bl e s elf a c tin g t end ency t o fu rth er th e -
s u p rem a cy of the m os t d ev el op ed typ es H ere h owev er t hi s

-
.
, ,
hi nt mu s t s uffi ce : expl a na ti on wou ld c a rry u s t oo fa r out of

ou r li n e o f a rg u m en t For th e p res en t it w ill n ot l ea d u s
.
a s t r a y if we re g a r d thi s i nv ers e v a ri a ti on o f I n divid u a ti on
a nd Genes is a s ex a ct .
§ 3 28 .th T hu s ,
en t h e c on diti on which ea c h r a c e mu st
,
fu lfil if it i s t o s u rvive is a Condi ti on whi ch i n the na t u re of

, ,
t hi ngs it ever t ends to fu lfil I n the l a s t ch a pt er we s aw

,
.
th a t a sp eci es c a nn ot be m a i nt a i ned u nl es s the p ower t o

p res erv e i ndivid u a l li fe a nd the p ower t o p r op aga t e oth er
'
i ndividu a ls v a ry i nv er s ely A nd h ere we h a v e s een th a t .

,
i rr es p ectiv e of a n end t o be s u bs erved th ese p owers c a nn ot ,
d o oth er th a n v a ry i nv ersely On the one h a n d giv en a .

,
c ert ai n t ot a lity of d es t royi n g f or ces w ith whi ch th e s p ec i es

h a s t o c ont end ; a nd i n p r op orti on a s its m emb er s h a v e
s evera lly bu t sm a ll a bili ty t o res i s t th es e f orce s it i s requ i s it e ,
th a t th ey s h ou ld ha ve g rea t a bility t o f or m new i ndivid u a ls

'
a n d vi c e ver s a On th e oth er h a nd giv en the qu a nt ity Of

'
.
,
forc e a bs orb ed a s f ood or o th erwis e which the sp eci es can

, ,
OB VE RSE A P R I ORI P RINCI P L E .
43 1
u se to c ou nt erb a l a nce th es e d es t royi ng f orces ; a nd i n p rop or

ti on a s m u c h of it i s exp end ed i n p res erv i ng the i ndividu a l ,
littl e of it ca n be res erv ed for p rodu ci ng new i n divid u a l s ,
a nd vi ce ver s a . T h ere i s thu s c om pl et e a cc or d a nc e b et w ee n

t he requ i rem en t s c on s id ered un d er ea ch a sp ect T h e t wo
.
nec es s iti e s c orresp ond .
We might res t on thes e d ed u cti ons a nd th ei r s ev era l c orol

l a ri es
. With ou t goi ng further we might with s a fety a s sert
the genera l t ru ths th a t oth er thi ng s eiqu a l a dv a nci n g evol u
, ,
ti on m u st be a ccomp a ni ed by d ecli ni ng fertility ; a nd th a t i n ,
the high es t typ es fertility m u s t still f u rth er d ec rea s e if evol u

,
ti on still f u rth er i nc rea ses We might be s u re th a t if other

.
,
thi ngs equ a l the rel a ti on s b et ween a n org a ni s m a nd i ts en

'
vi ronm ent bec om e s o c h a nged a s p erm a nently t o dimi ni s h
the di fficu lti es of s elf p res erv a ti on th ere will be a p er m a nen t

-
,
.
i nc rea s e i n the ra t e of m ul tiplic a ti on ; a nd c onv ers ely th a t

, ,
a d ec r ea s e o f fer tility w ill res u lt wh ere a lt ere d ci rc u mst a nc es
m a k e s elf p res erv a ti on m ore l a b or i ou s

-
.
Bu t we need n ot c on t en t ou rs elv e s with th es e (i p ri ori

i nferenc es I f th ey a re t ru e there m u s t be a n a gr ee m ent
.
,
b etween them a nd the Ob serv ed f a cts L et u s see h ow fa r.
s u ch a n a greem ent i s t ra c eabl e .

C H A P TE R I V .
I I
D F F C UL T E S O F I I N D U C T IVE VE RI F I CA T IO N .
WE RE
l
§ 3 29 . p eci es s u bj ect t o the s a m e ki nds a nd

al l s
a m ou nt s Of d es t ru ctiv e f or c es it w ou ld be ea s y by com p a ri n g
, ,
di fferen t sp ec i es t o t es t th e i nv ers e v a ri a ti on of I ndivid u a ti on

,
a n d Gen es i s Or if e ith er the p ower of s elf p res erv a ti on or

.
-
t he p ower of m ul tipli ca ti on w ere c ons t a n t th er e w ou ld be ,
li ttl e diffi cu lty i n s eei n g h ow th e oth er ch a nge d a s the

d es t royi ng forces ch a nged Bu t comp a ri sons a re n ea rly
.
a l w a ys p a rti a lly viti a t e d by s om e w a n t of p a r ity E a ch .
f a ct or b es id es b ei ng v a ri a bl e a s a wh ol e i s c omp ou nd ed of
, ,
f a ct ors th a t a re s evera lly v a r i a bl e N ot s i m ply i s the su m

.
o f th e f or c es d es t ru c tiv e of ra c e di f ferent i n ev ery ca s e ; an d

n ot s i m ply a re b o th s et s o f f orces p res erv a tiv e o f ra c e u nlik e
i n th ei r t ot a liti es i n ev ery c a s e ; bu t ea c h i s m a d e u p o f a c ti ons

t h a t bea r s u ch ch a ngi n g p r op orti ons t o one a noth er a s t o
p reven t a ny p ositiv e es tim a ti on of its a m ou nt .
B ef ore d ea li n g w ith th e fa cts a s w ell a s we c a n it wi ll be ,
b es t t o gl a nce a t the chi ef di ffic u lti es ; s o th a t we m ay see

th e ki n d o f v erific a ti on w hich i s a l one p oss ibl e .
3 30 E
ith er a b s ol u t ely or r el a tiv ely to a ny s p eci es every
.
, ,
envi r onm en t di ff er s m ore or l ess f r om ev ery o th er .
T h ere a re th e u nlik eness es o f m edi a — a i r w a t er ea rth , , ,
org a ni c m a tt er ; s ev era lly i nv olvi n g s p eci a l resist a nces t o
m ov em ent a nd sp eci a l l oss es of h ea t T h ere a re the con .

D I FF I CU L T IE S OF I ND U CT I VE V E RI FICA TION .
433
t ra sts li m a t e : h ere grea t ex pen dit u re for the m a in t ena nce

of c
Of t emp er a t u re is needed and th ere v ery littl e ; in one

,
z one a n org a ni s m i s s u ppli e d wi th a b u n d a nt light a l l t h e
yea r rou nd a nd i n a nother only for a few m onth s ; this

,
regi on yi eld s a n a lm os t u nf a ili n g s u pply of w a t er w hil e th a t ,
e n t a il s t h e ex erti on of t ra v el l i ng m a ny mil es ev ery n ight for a
d ra u ght .
P erm a nent di fferen ces i n th e na t u res a nd di s t ribu ti on s of

a lim ent gre a tly i nt er f ere w i t h t h e c omp a r i sons T h e S wa l .
l ow g oes th rou gh m ore ex erti on th a n the S p a rr ow i n s ec u ri n g

a giv e n w eight of f ood ; bu t th en th ei r f ood s a re di ss imil a r
i n nu t ritiv e qu a liti es T h ere i s a w a nt of p a ra ll elism be t ween

.
t he ci rcu m st a nces of th os e h erbiv ores which liv e wh ere th e

pl a i ns a re a nnu a lly cov ered for a tim e with rich h erb a g e bu t ,
a fterw a r d s b ec om e p a rc h ed u p a nd o f th os e i nh a biti ng m ore

,
te m pera t e regi ons I ns ec t s wh os e l a rv ae fee d on a n a b u nd a n t

.
pl a nt a s do s ev era l of the genu s Va n es sa on the N ettl e h a v e

, ,
p ra ctic a lly a n en vi ronm ent v ery u nlik e t h a t of i ns ects s u ch

a s D ei l ep hi l a E up h or bi aa w h os e l a rv ae fee d on a c om p a ra tiv ely
,
ra re pl a n t— th e S ea S p u rg e -
.
A ga i n c om p a ri s ons b et w een c rea t u res oth erwi s e a ki n i n

,
th ei r c ons tit u ti ons a nd ci rcu m s t a nces a re hi nd ered by i ne ,
qu a liti es i n th ei r rel a ti ons to enemi es T wo a n im a ls of.

~
whi c h on e is p re d a t or y a n d h a s n o f oes bu t p a ra sit es w hil e

th e oth er is m u ch pu rsu ed c a nno t p rop erly be cont ra s t e d
,
with a vi ew to d et ermi ni ng the i nfl u ence of si ze or com

p l ex i ty .
With ou t mu ltiplyi n g i nst a nc es it will cl ea r enou gh, be

th en th a t the a ggreg a t e of d es t ru ctive a cti ons p os itiv e a nd ,
n eg a tiv e which ea ch s pe ci es h a s t o c on t en d with is s o

, ,
u ndefin a bl e i n the a m ou n t s a n d ki n ds of its c om p onent s

,
th a t nothi ng bey ond a v a gue id ea of its rel a tive t ot a l ca n be

f orm ed .
§ 331 . Besid es th es e imm ens e v a ri a ti on s i n the ou t er

a cti ons to be c ounte r b a l a nc ed th ere a re imm ens e v a ri a ti ons
-
,
74
43 4 L A WS OF MU L T IP L ICA TI ON .
i n the i nner a cti ons requ i red t o c oun t er b a l a nc e th em E ven

-
.
were s p ec i es s i m il a rly conditi oned s elf p res erv a ti on w ou ld

,
-
requ i re o f t h em e xt re m ely u n lik e exp e n dit u res o f forc e .
T h e c os t Of l oc om oti on i ncr ea ses i n a g rea t er r a ti o th an

th e s i ze I n Vi rt u e of th e l a w th a t th e w eight s of a ni m a l s i n
.
c reas e a s the cu b es o f t h ei r di m en si on s whil e th ei r p owers o f

,
be a rin g s t ra i ns i ncrea se only a s th e s qu a res of th ei r di m en

s i on s p res erv a ti on of i ts v a ri ou s a ttit u des requ i res a
l a rg e a ni m a l t o c onsu m e m ore s u b s t a n ce in p r op orti on t o i t s
weight th a n it req ui res a s m a ll a n i m a l t o c ons u m e ; a n d th ere
,
res u lt s,
oth er thi n gs equ a l ,
a di ffi cu lty of s elf m a i n t en a n c e -
which au gm ent s i n a m ore ra pid r a ti o th a n the bu lk N or .
m u s t we overl ook the f ur th er c omplic a ti on th a t a m on g ,
a q u a tic c rea t u res th e v a r i a ti on of res i s t a n c e of th e m ediu m
t end s to p rodu c e a n opp os it e eff ect .
A ga i n th e h ea t c onsu mpti on i s a c h a n gi ng el em ent i n th e

,
-
t ot a l e xpen s e of s el f p res er v a ti on C rea t u res whi ch h a v e tem

-
.
e ra tures sc a rc ely a b ov e th a t of th e a i r or w a t er m a y oth er

p , ,
thi ngs equ a l a ccu mu l a t e m ore s u rplu s nu t rim ent th a n c rea

,
t u res which h a v e t o k eep th ei r b odi es wa r m spit e of the con

ti nu a l l os s by ra di a ti on a n d c on d u cti on T his difference of
.
c ost i s m odifi ed by th e p res enc e or a b senc e of na t u ra l cl oth

i ng ; a nd it i s a l s o m odifi ed by u nlik enes s es of s ize H ere .
th e b u lky a ni m a l s h a v e th e a dv a nt a ge : sm a ll m a s s es c oolin g
m ore ra pidly th a n l a rge ones .
D i s s imil a riti es of a tt a ck a n d d ef enc e a re a l s o c a u s es of

v a ri a ti on i n the ou tl a y for s elf m a i nt ena nc e A c rea t u re th a t
-
.
h a s t o h u nt ; a s c omp a red with a n oth er th a t g et s a suffi ci ency

of p rey by lyi ng i n w a it or a c rea t u re th a t es c a p es by s p ee d
,
a s c o mp a red with a n oth er th a t e s c a p es by c onc ea lm ent ,
o b vi ou s ly l ea ds a life th a t is phy s i ol o gic a lly m or e c os tly .
A ni m a l s whi ch p rot ect th ems elv es p a s siv ely a s th e H edge h og ,

-
by its s pi nes or a s the S kunk a nd the Mu sk ra t by th ei r i n -
t ol era b l e od ou rs a re rel a tiv ely ec onomic a l ; and ha v e the

,
m ore vit a l c a pit a l for oth er p u rp os es .
A mplific a ti on is nee dl ess T h es e i nst a nce s will sh ow th a t

.
D I F FI CU L TIE S OF IND U CT IVE VE RI FICA T ION .
43 5
anythi ng bey ond v ery g enera l c oncepti ons of th e i n divid u a l

exp endit u re s i n di fferen t c a s es c a nnot be
,
r ea c h ed .
332 . S till m ore

t gl ed a re we a m ong qu a lifyi ng con
en a n
sid era ti ons wh en we c ont ra st s p eci es i n th ei r p owers of m u lti

pli ca ti on T he t ot a l c ost of Genes is a dmits of ev en l es s
.
d efinite estim a ti on th a n d oe s the t ot a l c os t of I ndi vidu a

ti on I do not refer m erely t o the t ru th th a t the d eg ree o f
.
fertili ty d ep ends on f ou r f a ct or s— the a ge of c omm enci ng

rep r o d u cti on the nu mbe r i n ea ch b r oo d , t he f req u en cy of the
,
b rood s a nd the tim e d u ri ng which b roods c onti nu e to be

,
rep ea te d . T h ere a re m a ny f urth er Obst a cl es i n the wa y of

c omp a ri son s .
Were a l l mu ltiplic a ti on c a rri ed on s exu a lly the p robl em ,
w ou ld be l es s i nv olv ed ; bu t th ere a re m a ny ki nd s of a s exu a l

‘
m u ltiplic a ti on a lt erna ti ng with the s exu a l T his a sexua l

.
mu ltipli c a ti on i s i n s ome c a se s p erp et u a l i nst ea d of occ a ‘
si ou a l ; a nd o ft en ha s m ore f or ms th a n one i n the s a me

sp eci es The resu lt i s th a t we h a v e t o c omp a re w h a t is here
.
a p e r i odic p ro cess wi th wh a t is e l sew h e re a cyclic a l p roc ess
p a r tly c onti nu ou s a nd p a rtly p er i odic : the c a lc u l a ti on of

fertilit y i n thi s l a s t c a s e b ei ng next t o imp ossibl e .
We h a ve to a v oid bei ng mi s l ed by the a ssu m pti on th a t the

c ost of Genes i s i s m ea su red by the nu mb er of y ou ng p rod u ced ,
i ns t ea d O f b ei ng m ea su red a s it i s by th e w eight of nu t ri
, ,
m ent a b s t ra ct ed t o f orm the y oung p l us the w eight con

,
s u m ed in c a r ing for th em T hi s t ot a l weight m a y be v ery

.
div ers ely a ppo rti oned I n c ont r a st t o th e Cod with its
.
milli ons Of sm a ll ova sp a wned with ou t p rot ec ti on we m a y ,
pu t the H i pp oca m p us or th e P ip e fi sh w ith i ts few rel a tiv ely

-
, ,
l a r ge ova ca rri e d a b ou t by the m a l e i n a c a u d a l p ou ch or ,
s ea t ed i n h em i s ph eric a l pits in i ts s ki n ; or we m a y pu t the

s till m ore r em a r k a bl e g enu s A ri a s a n d e s p ec i a lly A r
, ias
B oa ken — a fi sh s om e six or s even i nch es l on g which p rod u ce s ,
ten or a d ozen eggs 5 — 1 0 mm in di a m et er th a t a re ca r ri ed

.
,
by th e m a l e i n hi s m ou th till th ey a re h a tched H ere th ou gh .

43 6 L A WS OF MU L TI P L I CA TION .
th e d egrees of f ertility if m ea s u red by the numbe rs of

,
f ertilized germs d ep os i ted a re ext rem ely un l i k e, th ey a re

,
l es s u nlik e if m ea s u red by th e nu m b er s of y ou ng which a re

h a tched a nd su rviv e l ong enou gh to t a k e ca re of th ems elv es ;
n or will t h e t a x on th e p a r en t C od s eem s o imm en s ely di f -
ferent fr o m th a t on th e p a ren t A ri a s if th e m a s s es of th e -
,
ova i n s t ea d o f th ei r n u m b er s a re c om p a red
, ,
A ga in, .
w hil e s om etim es t h e p a rent a l l oss i s littl e el s e bu t the m a t ter

d ed u ct ed to f orm egg s &c a t oth er t i m es it t a k es the
,
.
,
s h a p e Of a sm a ll di rect d ed u cti on j oi ned w ith a l a rg e i n di rect
o u tl a y T h e Ma s on w a s p f u rni s h es a typic a l i nst a nc e

.
-
In .
j ou rn eyings hi th er a nd thith er t o f etch bit by bit the

m a t er i a ls for bu ildi n g a c ell ; i n p u tti ng t ogeth er th es e
m a t eri a l s a s well a s i n s ec reti ng gl utin ou s m a tt er t o a ct a s
‘
c em en t ; a nd th en a ft erwa r d s i n th e l a b ou r of s ee ki ng for ;
, ,
a n d c a rryi ng th e s m a ll c a t erpill a r s wi t h whi ch it fill s u p the

,
c ell t o s erve i ts l a rv a with fo od wh en it em erges f rom the

egg ; th e Ma son w a s p exp en d s m or e s u b s t a n ce th a n i s c on
-
t a i ned i n the egg it s elf A n d thi s s u ppl em ent a ry exp en dit u re

.
i s m a ni fes tly s o grea t th a t but few eggs ca n be h ou s ed a nd

p rovi s i oned .
E s tim a t es of th e c os t of Genes i s a re f u rth er c omplic a t ed

by v a ri a ti on s i n th e ra ti o b orne by the two s ex es A m ong .
Fishes th e m a s s of milt a pp roa ch es i n s i ze the m a ss of sp a wn

bu t a m on g high er Ver tebra ta th e s u b s t a nc e l os t by th e one
s ex i n th e sh a p e of s p er m cell s i s s m a ll c om p a red w ith th a t
-
l os t by the o th er s ex i n th e s h a p e o f a lb u m en st ored u p i n -
the eggs or bl ood s u ppli ed t o the f oet u s or milk giv en t o th e

, ,
y ou ng T h en th ere c om e th e differen c es of i ndi rect t a x

.
o n m a l es a nd fem a l es Whil e f req uently the f ost eri ng of

.
, ,
th e y ou n g d ev olv es enti rely on the f em a l e occ a si ona lly th e ,
m a l e u nd ert a k es it wh olly or i n p a rt A ft er b u ildi ng a .
nes t , t he m a l e S tickl eb a c k g u a r d s th e eggs till th ey a re
h a tch ed ; a s d oes a ls o th e grea t S i l urus g l a ni s for s om e f orty

d a y s d u r in g whi c h h e t a k es n o f ood A nd th en a m ong m os t
,
.
,
bi rds we h a v e the m a l e occ u pi ed i n fee d ing the fem a l e d u ri ng

,
D IF FI C U L TI E S OF IND U CTIVE VE RI FICA TI ON .
437
i ncu b a ti on ,
a nd y oung a ft erw a rds E vidently a l l th es e
t he .
d i fi erences a ffect th e p rop orti on bet ween th e t ot a l c os t of re

p rod u cti on a nd the t ot a l c ost o f i ndividu a ti on .
Wh eth er the sp eci es i s m on oga m ou s or p olyg a m ou s a n d ,
wh et h er t h ere a re m a r k e d diff erences of siz e or o f s t ru c tu re

b et ween m a l es a nd f em a l es a re a l s o qu esti on s not t o be ov er
,
l ook ed I f th ere a re m a ny f em a l es t o one m a l e the t ot a l

.
'
,
qu a ntity of a s s imil a t ed m a tt er d ev ot ed by ea ch genera t i on t o

t he p r od u cti on of a new g enera ti on i s grea t er th a n if th ere
,
i s a m a l e t o ea ch f em a l e S i m il a r ly wh ere t he requ i rem en t s

.
,
a re su c h th a t s m a ll m a l es w ill s u ffic e t he l a r g er q u a n tity o f

,
f oo d l eft for the f em a l es m a k es p os s ibl e a g rea t er s u rplu s

a v a il a bl e for rep ro d u c ti on A n oth er c a u s e h a s a lik e effect
. .
Wh ere th e h a bit s of the r a ce ren d er it nee dl es s th a t b oth

s ex es s h ou ld h a v e d evel op ed p ow ers o f l oc om oti on— wh ere ,
a s i n th e Gl ow wo r m a n d s u nd ry L epi d op tera
-
th e f em a l e is ,
W i n gl es s w hil e the m a l e h a s w i ng s— th e c os t of I n dividu a ti on

not bei ng s o g rea t for th e s p eci es a s a wh o l e th ere a ris es a ,
g rea t er res erv e for Genesi s : the m a t ter whi ch w ou ld other

wi se h a v e g one to the p rod u cti on of wi ngs a nd the u sing Of
th em m a y g o t o the p rod u c ti on of ova
, .
Oth er com pli ca ti on s a s th os e whi ch we see i n Bee s a n d

,
A nt s might be d w elt on ; bu t the f oreg oi ng will a mply s erv e

,
the i nt end ed p u rp os e .
333 . To t i by c omp a ris on of c a s es wh eth er I ndi

s
a c er a n
vid u a ti on a nd Genes is v a ry i nversely i s thu s a n u nd ert a ki ng

,
s o b es et w i th di f
ficu lti es th a t we might d es p a i r of a ny s a t i s
,
fa ct ory r es u lt s wer e n ot th e rel a ti on t oo m a r k ed a on e t o be

,
hidd en ev en by a l l th ese c om pl exiti es S p e ci es a re s o ex

.
t rem el y c ont ra s t ed i n th ei r d egr ees of ev olu t i on a nd s o ,
ext rem ely c o nt r a s t e d i n th ei r r a t es o f m u ltipli ca ti on th a t th e

,
l a w o f r el a ti on b et ween th es e t ra it s b ec om es u nm i s t a k a bl e
wh en the evid ence i s l ook ed a t i n its ens em bl e T hi s we sh a ll
.
s oon fi nd on ra n gi n g i n or d er a nu mb er of typi ca l c a s es .
I n d oi ng thi s it will be c onv eni ent to negl ect t emp ora rily , ,
438 L A WS OF MU L T IP L I CA TION .
al l unl ik enes s es a m ong the ci r cu m s t ances in whi ch org an

i s ms a re pl a c ed A t the ou t s et we wi ll t u rn ou r a tt ent i on
.
,
wh olly t o the a nt a g on i s m di s pl a yed b et ween the i nt egr a tiv e

p roces s whi ch resu lt s i n i ndi vidu a l ev ol u ti on and the di s i nt e
r a t i ve p ro c e s s w hich res u lt s i n m u ltiplic a ti on o f i n divid u a l s ;
g
a n d thi s w e will c onsid er fi rst a s we s ee it u nd er th e s ev er a l
f orms of a ga m ogenes i s a nd th en a s we see it u n der the

,
s ev era l f or ms of g a m ogenes i s . We w ill next l oOk a t the

a n t a g on i s m b et ween p r op a g a ti on a n d th a t ev ol u ti on whi c h is
s h own by i nc rea s ed c ompl exity .A nd t h en we wi ll c on s id er

th e rem a i ni ng ph a s e of th e a nt a g oni s m a s it exi s t s be t w een
,
th e d egree of fer ti l ity a n d th e d egree of ev ol u ti on exp res s ed

by a ctivity .
A ft erwa rd s p a ss i ng t o t h e v a ryi ng r el a ti on s bet wee n

,
or g a n i s m s a n d th ei r en vi ron m en t s we will n ot e h ow rel a tiv e

,
i ncrea s e i n the s u pply of f ood or r el a tiv e d ecrea s e i n the

,
qu a n tity of f or ce exp end ed by th e i n divid u a l ent a il s rel a tiv e

,
i ncrea s e i n the qu a ntity of f orc e d ev ot ed t o mu ltipli c a ti on ,
a n d vi ce vers c
i .
Cert a in inor qu a lifica ti ons t ogeth er with s u nd ry imp or

m ,
t ant c oroll ari es, m ay then be ent ered u p on .

CH A P T E R V .
A N T A G N S M BE T W E E N GR
O I O WT H A N D A SE X UA L I
GE N E S S .
334 . WH E N ill u s t ra ti ng ,
i n P a r t I V, th e
ph ol og ic a l
m or
c omp os i ti on of pl a n t s a n d a ni m a l s th ere were s et d o wn i n

,
g r ou p s nu m er ou s fa cts whi ch we h a v e h ere t o l ook a t from

,
a n oth er p o i n t of vi ew T h en w e s a w h ow by u n i on o f s m a ll
.
,
s impl e a gg reg a t es th ere a r e p r od u c e d l a r g e c om p ou n d a gg re

,
g a t es N ow we h a v e t o ob s erve th e rea ctiv e effec t O f this

.
p roces s on the rel a tive nu m b er s Of the a ggreg a t es Ou r pre .
s ent s u bj ect i s the a nt a g on i s m of I ndivid u a ti on a nd Genes is

a s s een u n d er i t s s i m pl es t for m i n th e self —evid en t t ru th th a t
,
t h e s a m e qu antity of m a tt er m a y be divid ed i nt o m a ny s m a ll
w h ol es or few l a r ge w h ol es ; bu t th a t n u m b er n eg a tiv es l a r g e
ne s s a nd l a r genes s neg a tiv es nu mb er .
I n se tti n g d own s om e ex a mpl es we m a y c on v en i en tly

a d opt th e s a m e a rra n g em en t a s b e f ore We will l oo k a t th e
.
fa ct s a s th ey a re p res en t ed by v eget a l a gg rega t es of th e fi rs t

or d er of th e s ec on d or d er a n d of the thi rd or d er ; a n d th en
‘
, ,
a s t h ey a re p res en t ed by a n im a l a gg r eg a t e s o f th e s a m e th ree
o r d er s .
335 . T he
din a ry u ni cell u l a r pl a nt s a re a t on c e mic ro
or
s copi c a nd enorm ou s ly p r olifi c The Oft en ci t ed S p haer el l a

.
n i va li s whi ch sh ow s its i m m ens e p ow ers o f m u ltiplic a ti on by

,
r edd en i n g w i d e t r a c t s o f s n ow i n a s i ngl e night d oes thi s by ,
d ev el opi ng i n its c a vity a b roo d Of y ou ng c ell s w hi ch being , ,
43 9
44 0 L A WS OF MU L TI P L ICA TION .
p res ently set f ree by the bu r sti ng of the p a rent c ell s ev era lly -
,
g row a nd qu ickly rep ea t t he p roc ess T he lik e occu rs a m ong .
s u n d ry o f th ose ki n d red f orm s o f mi nu t e A l g ae w hi c h by ,
th ei r enor m ou s nu m b ers s om etim es s u dd enly ch a ng e p ool s to

,
a n op a qu e g reen S O t oo th e D es m i di a cece o ft en m u ltiply s o

.
, ,
g rea tly a s t o c ol ou r the wa t er ; a nd a m on g th e D i a tom a ceae

t he ra t e of gen es i s by s elf divi s i on i s s om ethi ng rea lly ex
-
,
t ra or di nary S o s oon a s a f ru s t u l e is divid ed i nto two ea ch

.
,
Of the l a tt er a t on ce p roceeds with the a ct of s elf divi s i on s o -
’
th a t t o u s e P rofess or S mith s a pp ro xim a tiv e c a l cu l a ti on Of
,
t he p os s ibl e ra pidity of m u ltipli c a ti on s u pp os i ng the p roc es s ,
t o o ccu py i n a ny s i ngl e i ns t a nc e t wenty f ou r h ou rs

, ,
-
,
s h ou ld h a v e a s t h e p rogen y of a si ngl e fr us t u l e th e a m a z i ng
, ,
’
n u mb er o f on e th ou sa n d milli on s i n a s i ngl e m on th In .
th es e ca s es the mu ltipli ca ti on i s s o c a rr i ed on th a t the p a rent

i s l os t i n the O ffs p r i ng— th e ol d i ndivid u a lity di s a pp ea rs eith er
i n th e s w a rm s of zoos p ores it di s s olv es i n t o or i n the two ,
or fou r new i n divid u a liti es s im u lt a n eou s ly p ro d u ce d by fi s
si on. Veget a l a gg rega t es of the fi rs t or d er h a v e h ow , ,
e v er a f orm o f a g a m o gen es is i n whi c h th e p a r en t i n divid u a l

,
ity i s n ot l os t : the y ou ng c ell s a ri s e fr om the ol d cell s by ex

t ern a l g emm a ti on T hi s p roces s too rep ea t ed a s it i s a t
.
, ,
sh ort i nt erv a l s resu lt s i n i m m ens e fertility T he Y ea s t

, .
fu ngu s whi ch i n a few h ou rs th u s p r op a g a t es it self throu gh

,
ou t a l a rg e va t of w o rt o f fer s a f a m ili a r ex a mp l e
, .
I n c ert a i n c om p ou nd f orm s th a t m u s t be c l a s s ed a s pl a n ts
of th e secon d ord er o f a ggreg a ti on th ou gh v ery mi nu t e o n es
, ,
s elf divi s i on simil a rly i nc rea ses th e nu m b er s a t high ra t es

-
.
T he S a r ci na ven tri cu l i a p a ra s iti c pl a n t which i n fes t s th e

,
st om a ch a nd s wa rm s a f res h a s f a s t a s p revi ou s s wa r m s a re
v omit ed s h ow s u s a S p ont a neou s fi s s i on of cl u s t ers of c ells
, .
A n a lli ed m o d e of i ncrea s e occu rs i n Ge ni u m p ec tor a l e : ea c h

c e ll o f th e cl u s t er res olvi n g it s elf i nt o a s econ d a ry cl u s t er ,
a n d the s ec on d a ry cl u s t er s th en s ep a ra ti n g S u pp os i ng .
,
w hi c h i s v ery p rob a bl e th a t a y ou ng Goni u m a ft er t wenty

,
f ou r h ou rs is c a p a bl e of d evel opm en t by fis s i on it f oll ow s ,

GROWTH A ND AS E X UA L GE NE SIS .
44 1
th a t und er f a v oura bl e c ondi ti ons a si ngl e c ol ony m a y on th e

s ec ond da y d eve l op 1 6 on the thi rd 25 6 on the f ou rth
, ,
a nd a t t h e en d of a wee k o th er org a n i s m s lik e

”
it s el f
. I n th e Vol voci n ece thi s c onti nu a l di s s ol u ti on of a
p ri m a ry c omp ou n d i ndividu a l in t o s ec ond a ry c omp oun d i ndi
vi du a l s is c a rri e d on en d og en ou sly a n d on a m o difi e d sy s t em
, ,
s om e onl v of the c om p onent c ell s g ivi ng origi n t o y oun g

c ol oni es a n d the p a rent bu rs ting t o libe ra t e them
,
T he .
nu mbe rs a ri s i n g by thi s m eth o d a ls o a r e s om e tim es s o gr ea t,
a s t o ti nt l a rg e b odi es Of wa t er . More fu lly es t a b

l i s hed a nd or ga ni zed a gg reg a t es of the s ec on d or d er s u ch a s ,
the high er T h a ll ophyt es a nd the l ower A rch eg oni a t es do not ,
s a c r ifi ce th e i r i n divid u a liti es by fi ss i on ; bu t n ev erth el es s by ,
the ki n d red p r oc es s of ge mm a ti on a re c on ti nu a lly hi nd ered

,
i n t he i nc r ea s e Of th ei r i n divi d u a liti es T he gem m ae c a ll ed

.
t et ra s p ores a re c a s t off in g rea t nu m bers by th e m a ri ne A l gae .
A m on g th ose s i m pl e J u ng erm a nni a c ece w hi c h c ons i s t of s i ngl e

f rond s th e y ou n g ones th a t bu d ou t g row for a tim e i n c on
,
nexi on w ith th ei r p a ren ts

. s en d rootl e t s f rom th ei r u n d er
,
sid e s i nt o the s oil a nd p resently s ep a ra t e th em s elv es

,
ha bit wh ich a u gm en t s th e nu mb er of i n divid u a ls in p rop or

ti on a s it ch ecks th ei r g rowth s .
P l a nt s o f th e t hi r d or d er o f c omp os iti on a ri s ing by a rrest ,
o f thi s s ep a ra ti on e xhibit a fu rth er c orres p on di n g d ec rea s e

,
i n th e a b u n d a n c e of t he a ggreg a t es f or m ed A rch eg oni a t es of .
i n feri or type s i n w hi ch the a x es p rodu ce d by i nt egr a ti on of

,
f r on ds a re but s m a ll a nd feebl e a re ch a ra ct eri zed by the

,
h a bit of th r owi ng off bu lbil s— bu d s h a p ed a x es which f a lli ng

-
,
a n d t a ki n g root a dd t o the nu m b er o f disti n ct i n divid u a l s

, .
T hi s a g a m ic mu ltipli c a ti on v ery genera l a m ong th e Mos s es

,
.
a n d th ei r ki n d red a n d n ot u n com m on u n d er a m o difi ed f or m

,
i n su ch high er typ es a s the F ern s m a ny of whi c h p r od u c e ,
y ou n g ones fr om the s u r f a c es o f th ei r f ron d s b ec om es verv ,
u nu s u a l a m on g P h aenog a m s The d et a chm ent of b u lbil s

.
,
th ou gh n ot un known a m ong th em i s exc epti ona l A nd w hil e

, .
it i s t ru e th a t s om e fl oweri ng pl ants a s the S t ra wb erry , ,

44 2 L A WS OF MUL TI P L I CA TI ON .
mul tiply by a p roc ess a l l i ed t o g emm a ti on, yet t hi s is not

ch a ra ct eristic of the cl a ss A l ea di ng t ra it of th es e high es t
.
gr ou ps t o which the l a rges t m emb ers Of the veget a l ki ng

,
d om b el on g i s th a t a ga m ogenesi s h a s s o fa r c ea s ed th a t it
, .
d oes not u s u a lly origi n a t e in d epend ent pl a nts T h ou gh the .
a x es whi c h bu ddi ng one ou t of a noth er c omp os e a t ree

, , ,
a r e th e eq u iv a l en t s of a s ex u a lly p r od u ce d i n divid u a ls
; yet
-
th e a s exu a l p ro d u cti on o f th e m st op s s h ort of s ep ara ti on .
T h ese v a s t i n t eg ra ti on s a ri s e w h ere s p ont a neou s disi nt egr a

ti on a nd the mu ltiplic a ti on effect ed by it h a v e c ome t o a n
, ,
e nd .
T hu s , n ot
f orgetti ng th a t c er t a i n P h aen og a m s a s Begonia ,
h
p yl l om a nia ca rev ert t o qu it e p rimitiv e m od es of in crea s e
, ,
w e m a y h old it a s b ey on d qu es ti on th a t w hil e a m on g t he
m o s t minu t e pl a n t s a s exu a l m u ltiplic a ti on i s u n iv ers a l a nd ,
p rod u c es enorm ou s nu mb ers i n sh or t peri od s it b ec om es s t ep ,
by s t ep m ore res t r i ct ed i n ra nge a nd f requ ency a s we a d

v a nce to l a rg e a nd c om p oun d pl ants ; a nd di sa pp ea r s s o
g ener a lly f rom the high es t a n d l a rgest th a t it s occu rr enc e is ,
reg a rd ed a s a n om a l ou s .
336 . x mpl es furni s h ed by a nim a ls m a k e cl ear

P a ra ll el e a
the p u rely qu a ntit a tiv e n a t u re Of thi s rel a ti on u n d er its ori gi

na l f or m A m ong t h e P r o tozoa a s a m ong the P r otop hy ta
.
, ,
th ere occu r s th a t p rocess by which the i ndividua lity of the

p a rent i s wh olly l os t i n p r odu ci ng offs p ri n g— the b rea ki ng
u p o f th e p a r en t a l m a ss i n t o a n u mb er o f g erms S om e Of .
t he I nfus or ia a s for i nst a nc e th os e of the genu s K ol p oda

,
a n d s ev er a l a lli ed g ener a b ec om e ency s t ed a nd s u bs equ ently

,
b rea k u p i nt o y ou ng ones T he m or e f a mili a r m od e

.
o f i n cr ea s e a m on g th e s e a ni m a l a gg reg a t e s o f th e fi r st or d er
-
,
by fi s s i on th ou gh it s a crifi ces the p a rent i n dividu a lity by

,
m er gi ng it i n th e i ndivid u a liti es of the two p rod u ced s a cri ,
fi ces it l es s c ompl et ely th a n d oes the di s s ol u ti on i n t o a grea t

nu mb er o f ger ms Occ u rri ng h owev er a s t hi s fis s i on d oes
.
, , ,
v ery f requ ently a nd b eing c ompl et ed i n s om e c a s es th a t

, ,
GROWT H A ND A S E X U AL GE NE S IS .
44 3
h a v e been Obs erv ed i n the c ou rs e of h a lf a n h ou r it r esu lt s

,
- -
,
i n imm en s ely r a pid mu ltipli ca ti on

-
I f a l l i ts offs p ri ng s u r .
viv e a nd c onti nu e dividing th em s elves a si ngl e P a ra m ceci u m ,
i s sa id t o be c a p a bl e of t hu s origi n a ti ng 2 6 8 m i lli on s i n th e
c ou rs e of a m onth *
N or i s thi s the grea t es t kn own ra te
.
Of i n c rea se A n oth er a n i m a l cu l e vi s ibl e only u nd er a high

.
,
m a gni fyi ng p ower i s c a l cu l a t ed t o genera t e 1 7 0 billi ons

“
,
d th s e en or m ou s p ower s o f p ro p a g a ti on
i n f ou r d a ys 1 A n e
.
'
a re a c c om p a ni e d by a mi n u t enes s s o e xt rem e th a t o f s o m e ,
s p ec i es one d ro p o f w a t er w ou ld c ont a in a s m a ny i n divid u a ls
a s th ere a r e h u m a n b ei n g s on the E a rth ! E v en if w e a ll ow

a l a r g e m a r gi n for ex a gg er a ti on i n th es e es tim a t es it is ,
b ey on d qu es ti on th a t a m on g th ese s m a ll es t Of a ni m a l s the
ra t e o f a s e x u a l m u ltipli c a ti on i s i m m ensely t h e g r ea t es t ;
’
a nd thi s s u ffi ces fo r t h e p u r p o s es o f a r gu m en t .
Of a ni m a l a ggr eg a t es b el ongi n g t o th e sec on d or d er th a t ,
m u ltiply a sexu a lly with ra pidity the f a m ili a r P olyp es ,
fu rn i s h c on s picu ou s ex a m pl es By gem m a ti on i n m os t c a s es .
,
i n ot her c a s es by fi s s i on a n d i n s om e c a s es by b oth the , ,
a ga m o gen es i s i s ca rri ed on a m on g th e s e t r ib es A s s h own i n .
Fig 1 48 th e bu ddi ng of y ou ng ones f rom the p a rent H ydra

.
,
is c a rr i ed on s o a ct iv ely th a t b ef or e th e Old es t of th em is
,
c a st off h a lf—a d ozen or m ore oth ers h a v e r ea ch ed v a r i ou s

-
s t a ges o f g row t h ; a n d ev en whil e s till a tt a ch ed th e fi r s t ,
f orm ed of th e grou p h a v e c om m en c ed bu ddi ng out f rom th ei r

s id es a s ec on d genera ti on of y ou n g ones I n the H ydra tu ba .
To m eet a poss i bl e cri t i ci s m i t sh oul d h e r em a rk e d th a t th i s ca l cul a

ti on as s um es th a t th e po w er of a se x u al r e p rod u cti on i s not e xh a u ste d b y
th e end of t h e m onth . I t h as be en found th a t “
v
th e succ e s s i e fi ss i ons of
P a r a m aeci um ca nno t cont i nu e i ndefi nitel y . After s om e h u ndre ds of ge ner a

t i ons th e p rod u ct s of fi s s i on a re sm al l , h a v e no m outh , a nd di e unl e s s b efor e
thi s v
th ey ha e b een al l owed to con j u
ga te wi th i ndi idua l s v of a nothe r b roo d .
”
It m a y, how ev e r, be fa i rl y t a k en for gra nte d th a t s om e h undre ds of g enera

t i ons woul d ta k e l onge r th a n a m onth .
{E v en thi s num be r i s fa r x
e c eed ed . D r E dw a rd Kl e in, i n
. a l ect ure h e
ga v e at th e Ro yal Ins ti tution on J une 2, 1 8 9 8 , a s s erte d th a t 24 6 b a ct e ri a in

a c ub i c cent i m etr e of nu t ri ti ve l i qu i d woul d m ul t i pl y to
'
i n the
co u rs e o f t wenty -four hou rs a ra t e w h i ch , at th e e nd of t h e thi rd d ay, woul d
gi v e , as th e Offs p ri ng o f o ne i ndi v i dua l ,

444 L A WS OF MUL TIP L I CA T I ON .
thi s gem mip a rou s mu l tipli ca ti on i s f rom tim e t o tim e i nt er

ru pt ed by a t r a n s v ers e s plitti n g u p o f th e b ody i nt o s egm en t s
-
,
whi c h su cces s iv ely s ep a r a t e a n d s wim a wa y : t he resu lt of

t he tw o p rocess es b ei n g th a t i n the c ou r s e of a s ea s on th ere
, ,
a r e p r o d u c e d f ro m a s i n gl e g er m g rea t nu mbe rs o f y ou n g
Med us ce whi ch a re th e a d u lt or sexu a l f or m s of th e s p eci es

, .
Res p ecti n g c oel ent er a t e a ni m a l s of thi s d egree of com p os i

ti on it m a y be a dd ed th a t wh en we a s cen d t o th e l a rger
,
ki nd s we fi nd a sexu a l genes is fa r l es s a ctiv e T h ou gh c om .
p a ri s ons a re i nt erfered with by di fferen ces of st ru ct u re a nd

m od e of life yet the cont ra sts a re t oo s t riki ng t o h a v e th ei r
,
m ea ni ng s mu ch ob s cu re d I f for in s t a nce we t a k e a s olit a ry

.
, ,
A c ti n ozo on a nd a s olit a ry H ydr ozoon w e s ee th a t th e rel a ,
t i vel y g rea t b u lk of th e fi rst g oes a l on g w ith a rel a tiv ely

-
,
sl ow a ga m og enes i s T he c omm on S ea a nem ones a re bu t

.
-
occ a s i on a lly o b serv e d t o u n d er g o self divi s i on : m u ltipli c a ti on -
by b u ddi n g b ei ng i n s om e c a s es l a r gely f oll ow ed bu t th ei r ,
nu m ber s a r e n ot ra pidly i n cr ea se d by ei t h er p roces s A .
high er c l a s s of sec on d a ry a ggr eg a t es ex em plifi es the s a m e

g enera l t r uth with a di fferenc e I n the s m a ll er m em b ers the .
a ga m og enes is is i ncom pl e t e a n d i n t h e l a rger it di sa pp ea rs

,
.
T h e gem m a ti on of th e m i nu t e P ol yz oa th ou gh it d oes n ot ,
en d i n th e s ep a r a ti on o f t he y ou n g i ndivid u a l s h a bit u a lly ,
g oes t o th e ext en t of p r od uci ng f a m ili es of p a rt i a lly i nd e -
p end en t i n divid u a l s ; bu t thei r nea r a lly the P h or om s whi ch , ,
i m m ens ely ex ceed s t h e m i n s iz e i s s olit a ry a n d n ot gemmi ,
p a r ou s S o t oo i s it wi th th e A s ci di oi da A n d th en a m ong
.
, , .
th e t ru e M a l l a s ca whi ch a re rel a tiv ely l a rge no s u ch thi n g

, ,
is known a s fi ss i on or gem m a ti on .
T a k e n ext th e A nnu l os a i ncl u di n g u n d er thi s titl e th e

,
A nn el i da a nd A r thr op oda Wh en t rea ti ng o f m or ph ol ogi ca l

.
c omp o siti on r ea s ons w er e giv en for th e b eli e f th a t th e a nnu

,
l os e a nim a l i s a n a gg reg a t e of the thi rd ord er th e segm en ts ,
o f whi ch p r od u c ed one f rom a noth er by g emm a ti on ori

, ,
g i n a l l y b e ca m e s e p a ra t e ; bu t by p r o gres s iv e int egr a ti on or ,
a rres t e d disi n t egr a ti on th ere resu lt ed a typ e i n whi ch m a ny

,
GROWT H A ND A SE X UA L GE NE S IS .
44 5
g ents were pe rm a nentl y nit ed 205—7

' ’ '
Su ch se m u a nd n o e t
to P a rt of
videnc e th ere a s s igned is evid ence t o
the e ,
be h ere rep ea t ed i n illu s t r a ti on of th e di rec t a nt a g oni sm o f

Growt h a nd A s exu a l Gen es i s We s a w h ow a m ong the
-
.
,
l ower A nneli ds the s t ri ng o f s egments p rodu c ed by gemm a

,
ti on p resently divid es t ra ns v ers ely i nt o t wo st ri ng s ; a n d

how i n s om e ca ses thi s res o lu ti on of th e el ong a ti n g s t ri ng
, ,
of segm en t s i nt o g r ou p s th a t a re t o for m s ep a ra t e i n divid u a l s ,
g oes on s o a ctiv ely th a t a s m a ny a s s i x g rou p s a re fou n d i n

di fferent s t ages o f p rogr es s to u lti m a t e i n d ep end ence —a f a ct
*
implyi ng a hi gh ra t e of fi ss i pa rou s mu ltiplic a ti on T h en we .
s a w th a t i n the s u p er i or a n nu l ose typ es di s ti n gu i s h ed i n th e

, ,
m a s s by i nclu din g the l a r ger s p ec i es fi s s i on d oes n ot occu r ,

.
The high er A nnel id s do n ot p r op a ga t e i n thi s w a y ; th ere i s

no kn own c a se of new i n divid u a l s b ei n g s o f or m ed a m ong
th e Myri a p oda ; n or do th e Cru s t a c ea ns a ffor d u s a s i n gl e

i ns t a nc e of thi s p ri m or di a l m od e of i nc rea s e It is .
,
i ndeed t ru e th a t whil e a rti cu l a t e a n im a ls never mu ltiply

,
a s exu a lly a fter t his s i m pl es t m et h od a n d w hil e th ey a r e ,
ch a r a ct er i zed i n the m a ss by the c es s a ti on of a g a m ogenes i s of

ev ery ki n d th ere n ev erth el ess oc c u r i n a few o f th e i r s m a ll
,
s pe ci es t h ose high er for ms of a g a m o g en es i s k n own a s p a rth e

,
n ogen es i s a n d p seu d o p a rt h en og en es i s -
a n d th a t by th es e
s o m e o f th em m u l t iply v ery ra pidly H erea ft er w e s h a ll .
fi n d i n the int erp r et a ti on of thes e a n o m a li es fu rth er s u pp ort

, ,
for the g enera l d oc t ri ne .
T o the a b ov e e vid en ce ha s t o be a dd ed th a t whi ch th e

Ver tebr a ta p resen t T hi s m a y be v ery b r i efly su m m ed u p
. .
On the one h a n d this cl a s s wh eth er l ook e d a t i n the a gg r e

,
g a t e or i n its p a rticu l a r sp ec i es i m m ens ely exc eed s a l l oth er

,
c l a s ses i n th e s i zes of i t s i n divid u a l s ; a nd on th e o th er h a n d ,
a g a m og en es is u n d er a ny f orm i s a b s o l u t ely u nk n ow n i n it .
I f it be s a id th a t b u ddi ng occ u rs a m on g th e T u ni ca ta whi ch ,
Myr i a nia a f
’
It h as s i nce be en s h own th a t i n i
as c a ta a s m a ny a s 29 at
ta ch ed group s e x i st . See Ca m br i dg e N a tu ral His tor y, Vol II, . Worm s ,

R oti
fers an d P olyzoa , p 2 8 0 . .
44 6 L A WS OF MU L TI P L I CA TION .
un d er th e c omm on titl e of Chor da ta a re i nclu d ed i n the s a m e

,
phyl u m with the Ver tebra ta th en it m a y fi rs tly be repli ed

,
th a t th os e typ es whi c h h a v e no v ert eb rae c a nnot p rop erly be

c a ll ed Ver tebra ta a n d sec on dly th a t if
,
a s b ei ng C h or da ta
, ,
th ey mu s t be rec ogniz ed th en the exc epti on whi ch th ey pre

,
s en t fu rth er illu s t ra t es the t ru th th a t a g a m ogenetic mu lti

plic a ti on occu rs only i n c rea t u res sm a ll i n size or l ow in ,
st ru ct ure or b oth
,
.
§
33 7 . S u ch a re a few
l ea di ng f a cts s er vi ng t o sh ow h ow
d ed u cti on i s i ndu ctiv ely verifi ed i n s o far a s the ant a g oni s m
,
b et ween Growth a n d A s exu a l Genesis is c oncerned In .
wh a t ev er wa y we expl a i n thi s opp o siti on of the i nt egra tiv e

a n d di s i nt egr a tiv e p rocesse s th e f a cts a nd th ei r implic a ti on s
,
rem a i n t h e s a m e I n d ee d we need not c omm it ou r s elv es t o

.
a ny hyp oth es i s res p ecti ng t h e phy s i ca l c a u sa ti on I t s uffic es

.
t o rec ogni ze th e resu lt s u nd er th ei r m ost genera l a sp ects .
We c a nnot h elp a dmit t ing th ere a re a t work thes e two ant a

g on i s t t end en ci es t o a gg reg a ti on a nd s ep a ra ti on ; a nd we
c a nnot h elp a dmittin g th a t the p rop or ti on be tween the a ggre
g a ti ve a n d s ep a ra tiv e t en d enc i es mu s t i n ea ch c a s e d et ermi ne
,
t h e rel a ti on b et ween i ncrea s e i n b u lk of the i ndi vid u a l a nd

i ncrea s e of the ra ce i n n u m b er .
The a ntith es i s is a s m a ni fes t d p os teri ori a s it is nec es sa ry

at p ri ori . Whil e the mi nu t est orga ni s m s mu ltiply a s exu a lly
in th ei r billi ons ; whil e th e I nfus om a thu s mu ltiply i n th ei r
'
mi l li ons ; whil e the s m a ll c omp ou n d typ es next a b ov e th em

t hu s mu ltiply i n th ei r th ou sa nd s ; whil e l a rger and m ore
c omp ou nd typ es th u s mu ltiply i n th ei r h u nd reds a nd th ei r
t ens ; the l a rgest typ es do not thu s mu ltiply a t a l l C on .
ver s el y th ose which d o n ot m u ltiply a sexu a lly a t a l l

, a re a ,
billi on or a milli on tim es the s i ze of th os e whi ch thu s m u l

t i pl y w ith gr ea t es t ra pidity ; a nd a re a th ou s a n d tim es , or a
h u n d red tim es or t en tim es the si ze of th os e whi ch thu s
,
mu ltiply with l ess a nd l ess ra pidity With ou t s a yi n g th a t.
this i nvers e p rop orti on is regu l ar which a s we sh a ll h ere

, ,
GROWT H A ND A SE XU A L GE NE SIS .
44 7
a ft er see it c a nnot be we m a y u nh esit a tingly a ss ert i ts

, ,
a v era g e t ru th
. T h a t th e sm a ll es t org a ni sm s h a bit u a lly re
p rod u ce a se xu a lly with im m ens e ra pidity ; th a t the l a rges t
org a ni s m s n ev er r ep r od u c e a t a l l i n thi s m a n ner ; a nd th a t
betwee n thes e ext rem es th ere is a genera l d ec rea s e of a s exu a l

rep ro d u c ti on a l on g with a n i n c rea s e of b u lk ; a re p r op o si
ti on s whi ch a dmi t of n o di sp ut e .
C H A P TE R VI .
A N T A GON S M B E T WE E N GR
I O WT H A N D S E X UA L GE N E S S I .
§ 338 . IN so fa r as it is
p r oc ess of s ep a r a ti on s exu a l
a ,
g enes i s i s lik e a s exu a l genes i s ; a nd is th eref ore equ a lly with ,
a se x u a l g en es i s o pp os ed t o th a t a ggreg a ti on w hi c h r esu l t s i n
,
gr owth Wh eth er d edu cti on i s m a d e f rom one p a rent or

.
f rom two wh eth er it is m a d e f rom a ny p a rt of the b ody

,
i ndifferently or from a s p eci a lized p a rt or wh eth er it i s m a d e ,
di rectly or i ndi rectly it rem a i ns i n a ny c a s e a d ed u cti on ;

,
a n d i n p r o p or ti on a s it is g rea t or f requ en t or b oth it m u s t

, , ,
r es t ra i n th e i n crea s e o f th e individ u a l .
H er e we h a v e t o g r ou p t ogeth er th e l ea di ng ill u s t ra ti on s
of thi s t ru th We will t a k e th e m i n the sa m e ord er a s
.
b e f or e
.
§ 33 9 T he l owes t v eget a l f orm s or ra th er we m ay say

.
, , ,
th ose f orm s whi ch we ca nnot cl a ss a s eith er di s ti nctly v eget a l

or di s ti nctly a n im a l s h ow u s a p r oce ss of s ex u a l m u ltipli c a
,
ti on th a t di ffer s mu ch l es s f rom the a s exu a l p roces s th a n i n

the high er f orms T he c omm on c h a r a ct er whi ch di s ti ngu i sh es
.
s ex u a l f ro m a s ex u a l g enes is i s th a t th e m a s s o f p r ot op l a s m
,
wh enc e a new g en era ti on i s t o a ri s e h a s been p rod u ced by the ,
u n i on of two p orti on s o f m a tt er which w ere b efore m ore w id e
l y s ep a ra t ed I u se thi s g enera l exp ress i on b eca u s e a m ong

.
,
the s i m pl es t A l gae this is n ot i nv a ri a bly m a tt er s u ppli ed

,
by different i ndivid u a l s : c ert a i n D i a tom a cece exhibit wi thi n

a si ngl e c ell the f or m a ti on of a sp ora ngi u m by a d ra win
, g
44 8
GROWT H A ND SE X U A L GE NE SIS .
44 9 .
t ogeth er of the opp os it e h a lves of the end och rom e i nt o a

b a ll M os tly h owev er s p or a ngi a a re p rod u cts of c onj u g a
.
, ,
t i on. Th e p r ot opl a s m i c c ont ent s of two c ell s u nit e t o form

the ger m m a s s or zyg ot e ; a nd th es e c on j u g a ti ng c ell s m a y be
-
eith er en ti r ely i nd ep end ent a s i n m a ny D es m i di a cece a n d i n

,
t he g a m et es o f m a ny C onfervo i deaz; or th ey m a y be t wo of
t he a dj a cent c ell s f orm i ng a th rea d a s i n s om e C onjuga tece,
a n d t h e g a m et es of C onfer voi dece ; or th ey m a y be c ell s
bel ongi ng t o a dj a c ent threa ds a s i n other C onju ga teae Bu t

, .
wh eth er it i s origi n a t ed by a si ngl e p a rent c e ll or by tw o -

,
p a ren t cell s the zyg ot e a ft er rem a i ni n g qu i escen t u ntil th ere

-
, ,
rec u r th e fit c onditi ons for g r o wth e ith er b r ea ks u p i nt o a

,
m u ltitu de of s p ores ea ch of which p rod u ces a n i ndividu a l

,
th a t u su a lly m u ltipli es a sex u a lly or germi na t es di rec tly to

,
p rod u c e one new i ndividu a l ; a nd the f a c t h ere t o be not e d

i s th a t a s the en ti re c ont en t s of th e p a ren t c ell s u n it e t o
,
-
f or m the zyg ot e thei r i ndivid u a liti es a re l os t i n t he g er ms of

,
a new g ener a ti on I n th es e mi nu t e s impl e typ es s ex u a l

.
,
p rop a g a ti on j u s t a s c ompl et ely s a crifi ces the life of the

p a rent or p a rent s a s d oes th a t form of a s exu a l p rop a ga ti on
,
'
i n w hi c h the p rot opl a s m res olv es i ts el f di rectl y i nt o z oo

sp ores A nd i n th e one ca s e a s i n the o th er thi s s a c rifi ce
.
,
i s th e c oncomit a n t of a p rodigi ou s f ertility S lightly

.
i n a dv a nce o f thi s bu t s till s h ow i n g u s a n a lm ost equ a l l os s

,
o f p a ren t a l lif e i n t h e liv es o f offs p r i n g i s th e p r oces s seen i n

,
s u ch u nicell u l a r A l gae a s B o trydi u m a n d i n mi nu t e F u ngi of

,
th e s a m e d eg ree of c om p os i ti on T h es e exhibit a rel a tiv ely

.
enor m ou s d ev el op m en t o f t he sp or e p r o d u c i ng p a r t -
a nd a n ,
a l m o s t en ti re a b s or pti on of t h e p a r ent a l s u b s t a nce i nt o it .
A s evid ence o f the res u lti n g p owers o f m u ltipli c a ti on we ,
h a v e bu t t o rem emb er th a t the s p rea d of m ou ld ov er s t a l e

f ood th e ra pid d es t ru cti on of c rop s by mild ew a nd oth er
, ,
ki nd red occu rrenc es a re m a d e p o ss ibl e by the i nc a lcu l a bly

'
nu m erou s s p o res th u s g enera t ed a n d u n iv er s a lly di s p e rse d .
P l a n t s a d egree high er i n c om p os iti on s u pply a p a ra ll el

s er i es o f ill u s t r a t i on s We h a v e am ong the l a rger Fu ngi i n
'
.
,
75
4 50 ~
L A WS OF MU L T IP L ICA TION .
whi c h th e p rod u ctiv e a pp a ra t u s i s rel a t iv ely so enorm ou s a s

re
t o c ons tit u te t h e os tens ibl e pl a nt a s imil a r s u b or di na ti on o f ,
t h e i ndivid u a l t o the ra ce a nd a simil a r ly imm ens e fertili ty

,
-
.
T hu s a s q u ote d by D r Ca rp ent er F ri es s a ys
, . i n a s i n gl e i n
,
divid u a l of Reti cu l a ri a m a xi m a I h a ve c ou nt ed ( ca l cu l a ted ? )

,
”
sp oru l es I t nee d s bu t t o n ot e the cl ou d s o f
.
p a rticl es s o mi nu t e a s t o l ook like s m ok e whi ch rip e p u ff

, ,
b a lls giv e off wh en they a re b u rs t a nd then t o rem em b er ,
t h a t ea ch p a rt icl e i s a p ot en ti a l f u ngu s t o be imp res s e d wi t h ,
t he a l m os t i n c onceiv a bl e p owers of p rop a g a t i on w hi c h th es e

pl a nt s p os s es s T he L i c h en s t oo fu rni s h ex a m pl es
.
, , .
T h ou gh th ey a re nothi ng lik e s o p rolifi c a s th e F u ngi ( the

difference yi eldin g a s we s h a ll h erea ft er s ee furth er su pp ort
, ,
t o t he genera l a r gu m en t ) yet th ere is a grea t p r o du ct i on o f

,
g erm s a nd a p r op orti ona t e s a c rifi ce of th e p a rent a l i ndi

,
vi du a l i ty Con s id era bl e a rea s o f t he th a ll u s d ev el op i nt o the

.
f ru it b odi es ch a ra ct eri s tic o f th e v a ri ou s f ungi which c om

-
,
bi ned with a lgae f or m th e different lich ens ( v a ri ou s m e mb ers

,
o f th e A s c om yce tes a nd th e Ba s i di om ycetes ) F rom th es e a re

p rod u c ed grea t nu m b ers of a sc osp ores or b a s idi osp ores a s the ,
c a s e m a y be Very m a n y li ch en s a ls o rep rod u c e th e m s elv es

.
by m ea n s of S or edi a i e littl e m a ss es of a lg a l c ell s cl os ely

, . .
,
w ra pp ed i n a we ft of f u n g a l hyph ae S om e con .
t r a s t s p res ent e d by t h e high er A l gae m a y a l s o be na m ed a s

ex empli fyi n g t h e i n v ers e p r op orti on b et ween the s i z e o f th e
i ndivid u a l a n d the ext ent of the genera tiv e s t ru ct u res Whil e .
i n the s m a ll er ki n d s rel a tiv ely l a rge p orti on s of the f ron d s a re

t ra ns f orm ed i nt o r ep r od u ctiv e el em ent s i n the l a rger ki n ds ,
t h es e p orti ons a re r el a tiv ely sm a ll : i ns t a nce the Ma cr ocys ti s
py fr i em a,
gig a n tic sea w ee d w hi ch s om etim es a t t a i n s a
-
l en gth of feet of whi ch D r Ca rp ent er rem a rks

,
.
T hi s d ev el opm en t o f th e nu t ritiv e s u r fa ce t a k es pl a c e a t
“
the exp en s e of th e f ru ctifyi ng a pp a ra t u s whi c h i s h ere qu it e ,

”
s u b ordi n a t e .
Wh en we t u rn t o v eget a l a ggrega t es of the thi rd ord er of

c omp os iti on f a c ts h a vi ng th e s a m e m ea ni ng a re c on spi cu ou s
,
.
GROWT H A ND S E X UA L GE NE SIS .
451
On th e av era ge th es e higher pl a nt s a re fa r l a rger th a n

pl a nt s of a l ower d eg ree of c omp os iti on ; a nd on the a v era ge
thei r ra t es of s exu a l rep rod u cti on a re fa r l ess S imil a r ly if .
,
a m on g A rch egon i a t es a nd P h aen og a ms we c o mp a re the ,
sm a ll er typ es with th e l a rg er w e fi nd them p rop or ti ona t ely

,
m ore p rolifi c T his i s not m a ni fes t if w e simply c a l cu l a t e

.
the nu mber of s ee ds rip ened by a n i ndivid u a l i n a si ngl e

s ea s on ; bu t it b ec om es m a n i fes t if we t a ke i nt o a cc ount the
f u rth er f a ct or which h ere com pli ca t es the res u lt— the a ge a t
whi c h s exu a l genes i s c ommences T h e s m a ller P h aenog a ms .
a re m ostly eith er a nnu a ls or p erenn i a ls

,
th a t di e d own
a nnu a lly ; a nd s ee di ng a s th ey do a n n u a lly b ef ore th ei r
d ea th s or th e d ea ths of th ei r rep rodu ctive p a rts it res u lts

, ,
th a t i n the c ou rs e of a yea r ea ch gives origi n to a mu ltitu d e

o f p ot en ti a l pl a n t s o f wh i ch every one m a y th e next y ea r if
'
, ,
p res er v ed giv e or igi n t o a n equ a l mu ltit u d e S u pp os i ng bu t

,
.
.
a h u n d red off s p ri ng to be p rodu ced the fi rs t yea r t en ,
th ou s a nd m a y be p rod u ced i n the s ec on d yea r a m illi on i n ,
the thi r d a hun d red milli on s i n th e f ou r th

, Meanwhil e, .
wh a t h a s b een the p os sibl e mu ltipli ca ti on of a l a rg e P h aeno
g a m ? Whil e its s m a ll c ongener h a s b een s eedi n g a nd dyi ng ,
a nd l ea vi ng m u ltit u di n ou s p rog eny t o s ee d a n d di e it h a s ,
simply b een gr ow i ng ; a nd m a y s o c onti nu e t o g row for ten

o r a d oz en y ea r s w ith ou t b ea r i n g fru it B ef ore a Co c oa nu t .
-
t ree h a s r ip en ed i ts fi rs t cl u s t er of nu t s the d es cend a nt s of ,
a w h ea t pl a n t s u pp o s i n g th em
,
a l l t o s u r viv e a nd m u ltiply ,
will h a v e bec om e nu m erou s enou gh t o oc cu py the wh ol e

su r f a ce o f th e E a rth S o th a t th ou gh wh en it b egi n s t o
.
,
‘
b ea r a t ree m a y a nnu a lly sh ed a s m a ny s eed s a s a h erb yet

, ,
i n c ons eq uence o f thi s d el a y i n be a ri n g i ts f ertility i s i ncom ,
p a r abl y l es s ; a nd i t s rel a tiv ely s m a ll fer tility b ec om e s s till

-
fu rth er r ed u c ed w h er e a s i n L odoi cea ca l l i pyge th e s eed s

, ,
t a k e two yea rs f rom the d a t e of fertiliz a ti on t o the d a t e of

‘
‘
germi na ti on .
3 4 0 S om e
. o b s erv er s st a t e th a t in c er a n t i P r o tozoa th ere
452 L A WS OF M U L TIP L I CA T I ON .
o ccu rs p rocess of c onj u g a ti on a ki n t o t h a t whi ch th e

a
P r o top hy ta exhi bit— a c oa l es cence of the s u b s t a nc e o f two

i ndivid u a l s t o form a ger m m a s s T hi s h a s b een a ll eg ed -
.
m ore e s p ec i a lly o f A c ti nop hrys I f this s t a tem ent s h ou ld .
be p r ov ed t ru e t h en of th e mi nu t e f orm s th a t a pp ea r t o be
*
,
m ore a nim a l th a n v eg et a l i n th ei r ch a ra ct ers s om e h a v e a ,
m od e of s exu a l m u ltipli c a ti on by whi ch the p a rents a re

s a c rifi c ed b odily i n th e p r o d u c ti on o f a new g enera t i on .
A m ong s m a ll a ni m a l a gg reg a t es of th e s ec ond or d er th e ,
fi rs t t o be c on s id ered a re of c ou rs e the C ael en tera ta A H ydra

’
.
o c c a s i on a lly d ev ot es a l a r ge p a rt o f i ts s u b s t a nce t o s ex u a l
genes is I n th e w a ll s of i ts b ody g rou p s of ova or sp erm a
.
,
t oz oa or b oth t a k e th ei r r i s e ; a nd d ev el op i n t o m a ss es
, ,
’
grea tly di s t orti ng the c rea t u re s f orm a nd l ea vi ng it m u ch ,
dimi ni s h ed w h en th ey es c a p e H ere h owev er g a m ogenes is is .

, ,
o bvi ou s ly s u ppl em ent a ry t o a g a m o g en es i s— th e i m m en s ely
r a pid m u ltipli c a ti on by b u ddi n g c on ti nu es a s l ong a s foo d is
a b u n d a n t a nd w a r m th s u f fi ci ent a nd is repl a ced by ga m o ,
genes i s only a t the cl os e o f the s ea s on A b ett er ex a mpl e .
To thi s p a s sa ge P rof . Ma cBri de a ppends th e rem a rk Th i s i s q uite

p rov en now, a nd th e s ta te m ent as it s ta nds is q ui t e co rre ct ; bu t fa r b e tt e r
a nd m ore m i nutel y work ed ou t cases a r e to be found a m ongs t th e Infusor i a .
In P a r a m ec i u m for e x a m pl e ,
t h e re a re norm a l l y p re s ent a l a rge m ac ro
nu c l e u s a n d a s m a l l i cro nu cl eus l yi ng a l ongs i de
m of i t Wh en two i ndi
.
vi du a l s a dh ere j
p re p a ra tory to c on uga tion, th e m a cronu cl eus bre a ks u p i n to
fra gm ent s w hi ch a re a b s orbed : the m i cronucl eu s — whi c h h as s om e ti m e p re
v
v i ou s l y di i ded i n to t wo— begi ns t o bre ak u
p furth er a nd e entu al l y form s v
e i gh t bod i es ; a l l of x
th es e e ce pt one di s a ppea r ; thi s l as t p i ece then d i i des v
i nto t wo ; o f th es e t wo one rep rese nt s a m a l e geni ta l ce ll , for i t p a s ses ov e r
i nto t he bodv of th e o ther P a r a m e ci um a nd fus es wi th one of th e t wo cor re
s pondi ng nu cl ei the re ; th us e a ch of th e t wo i ndi idu a l s v whi ch a dh ere
fert i l i ze s th e oth er . Th e t wo i ndi vi du al s t hen s ep a ra t e and the nucl e us
( res ul t of fus i on of m a l e a nd fem a l e nu cl ei ) in e ach di vi des i nto four . Of

th es e , t wo m v to one
o e e nd of the ani m a l a nd two to the oth er . Th e
a ni m a l then di v ides i nto v
t wo t ra ns er s el y— ea ch of t he p roduct s t hus h a v
i ng t wo nu cl e i whi ch form th e m i cro a nd .
m a cro-nucl eus of i t Thu s i t .
a p p e a rs th a t the f uncti on of s e x ua l uni on is s im p l y t o gi v e i ncre a sed vi gour

to al l th e vi t al proce s ses i nc l u di ng fi ssi on . S i nce as m enti oned abo v e ( p .
4 4 3 ) i f i t i s pre v ent ed the , p rod ucts of fi s s ion a re v

e ent ua l l y una bl e t o feed
the m s el v es .
GROWT H A ND S E X UA L GE NE S IS .
453
of th e l ti on b et ween s m a ll s ize a nd a c tiv e ga m ogenes i s

re a
.
a m on g l ow typ es o f t h e M e ta zoa i s s u ppli ed by t h e R o ti e ra

f .
Mi cros copic a s thes e a re th ey h a v e a g rea t ra t e of s exu a l

,
i nc rea se A cc or din g t o E h renb er g H yda ti na s en ta

“
.
, is
c a p a bl e o f a f ou r f old p rop a g a t i on e very t w en ty fou r or
- -
thi rty h ou rs b ri ngi ng forth in thi s tim e f ou r ova whi ch gr ow

, ,
f rom th e emb ry o t o m a t u rity a nd ex c l u d e th ei r fertil e ova

,
i n the s a m e p er i od T h e s a m e i n divid u a l p r odu ci n g i n t en

.
,
d a ys f orty egg s d ev el op e d with the ra pidi ty a b ov e cit ed thi s

, ,
ra t e
,
ra is ed t o t he t en th p ow er giv es on e milli on of i n di
,
vi du a l s f ro m one p a r en t on th e el ev en th d a y fou r milli on s

, ,
”
a n d on t h e t wel fth da y s ixt een milli on s a n d s o o n ,
.
E hrenberg h owev er ch a ra ct eri zed by H u xl ey a s

, ,
the
”
gr ea t es t l oc ker a nd the w ors t ob s erv er i s not a s a fe a u tho ,
rity a n d it i s be t t er t o s t a t e th e e s tim a t e o f L u d w ig P l a t e
, ,
wh o s a y s th a t H yda ti na l a y s fifty egg s i n two t o th ree week s

-
a nu m b er whi c h mu ltiplyi n g i n the m a nn er d es c r ib ed
, ,
will yi eld i n the tim e na m ed a mu ch s m a ll er t ot a l th ou gh s till

a n enor m ou s t o t a l .
The A nnu l osa i ncl u di ng a m on g th em th e i nferi or typ es

, ,
h a v e h a bits a nd c onditi ons of li fe s o v a ri ou s th a t only the

b roa d es t c ont ra sts ca n be i ns t a nc ed i n s u pp ort of the pr o
p ositi on b ef ore u s T he di fferences o f org a n i z a ti on a nd
.
a ctivity gr ea tly c om plic a t e th e i n v erse v a r i a ti on o f fe rtility
a nd b u lk B ea r i ng i n mi nd h owev er th a t th e ra t e o f m u lti
.
, ,
plic a ti on d ep end s mu c h l es s on the nu mb er of ea ch b rood

th an on the qu i c knes s with which m a t u r i ty i s r ea ched a nd a
n ew g enera ti on com m en ced it will be obvi ou s th a t th ou gh
,
A nnel id s rel a tiv ely enorm ou s i n si ze p rod u c e grea t nu m b er s

, ,
o f ova , y et a s th ey d o this a t c omp ara tiv ely l ong i nt er
v a l s th ei r ra t es of i nc rea s e f a ll imm ens ely b el ow th a t j u s t

,
i ns t a nced in the Rotifers A nd wh en a t th e oth er ext rem e

.
we com e t o the l a r ge a rticul a t e a n im a l s s u c h a s th e C ra b ,
a nd th e L obst er th e f u rth er di min u ti on o f f ertility is s een i n

,
t he s till l onger d el a y which occu rs be f ore ea ch new g enera ti on

begin s t o rep rodu c e .
454 L A WS OF MUL T I P L I CA TION .
P erh a ps th e bes t ex a mpl es a re s u ppli ed by vert eb ra t e

a ni m a l s ,
a n d esp ec i a lly th os e th a t a re m o s t f a mili a r t o u s .
C omp a ris on s b etw een Fi s h es a re u ns a tis fa ct ory b eca u se o f ,
ou r ign ora n c e o f t h ei r hi s t or i es I n s om e c a ses Fish es equ a l

.
i n b u lk p rod u ce wid el y di fferent nu mb er s of egg s ; a s the

C od whi c h s p a wns milli on s a t once a n d the S a l m on by ,
w hich nothi ng lik e s o gr ea t a n u m b er is sp a wned Bu t th en .
t h e eggs a re v ery u nlik e i n s i ze ; a nd if th e ov a ri a of th e two

fi s h es be c omp a red the di ff erenc e b etween th ei r m a sses is
,
c om p a ra tiv ely m od era t e T h ere a re i nd eed c ont ra s t s whi ch

.
, ,
s eem a t v a ri a nc e w ith the a ll eged rel a ti on ; a s th a t b et ween

th e C od a n d the S tickl eb a c k w hi ch th ou gh s o m u ch s m a ll er
, ,
’
p rodu ces f ewer ova . T h e S tickl eb a ck s ova h owev er a re-
, ,
rel a tiv ely l a r g e ; a n d th ei r t ot a l b u lk b ea r s a s g rea t a r a ti o t o
th e b u lk of th e S ti c kl eb a ck a s d oes th e bu lk of the C o d s ova

’
t o th a t of the C od . Moreov er i f a s i s not imp rob a bl e the

, ,
r ep r od u ctiv e a g e i s a rr iv e d a t ea rli er by th e S tickl eb a ck th a n
by the C od the fertility of the s p eci es m a y be g rea t er n ot

,
wi th s t a ndi ng the s m a ll er nu m ber p rodu ced by ea ch i ndi

vidu a l . E vid enc e whi ch a dmit s of b ei ng t ol era bly
w ell di s en t a ngl ed i s f u rni s h ed by B i rd s T h ey differ bu t
.
littl e i n th ei r gr a d es of org a ni za ti on ; a nd the h a bit s of life

thr ou gh ou t ext ens i ve grou p s of th em a re s o si m il a r th a t '
c omp a ri s ons m a y be f a i rly m a d e I t i s t ru e th a t a s h ere

.
,
a ft er t o be s h own the di fferenc es of exp endit u r e w hi c h

,
di fferen ces of b u lk en t a il h a v e d ou btl es s m u ch t o do wi th

,
th e diff erences of f ertility .B u t w e m a y set d own u nd er th e

p resent h ea d s om e o f th ose c a s es i n whi ch the a ctivity b ei ng ,
r el a tiv e ly s light d oes n ot grea tly i nt er fere w ith th e r el a ti on

,
w e a r e c on s id eri ng ; a nd m a y n o t e th a t a m ong su ch bi r ds
h a vi ng si m il a rly s light a ctiviti es the sm a ll p rodu c e m ore eggs
,
th a n the l a rg e a n d egg s th a t b ea r i n th ei r t ot a l m a ss a
,
grea t er ra ti o t o the m a s s of the p a rent C on s id er for ex

.
,
a m pl e ,
th e g a l li na c eou s bi r d s ; which a re lik e on e a noth er
a n d u nlik e bi rd s o f m os t oth er g rou p s i n flyi n g c omp a ra tiv ely
littl e T a king fi rst the wild m emb ers of thi s ord er which
.
,
GROWT H A ND SE X U A L GE NE SIS .
55
ra re ly b reed m ore th a n onc e i n a s ea s on we fi nd th a t th e ,
P hea s a n t h a s from 1 0 t o 1 4 eggs th e Bl a c k c oc k fro m 6 t o ,

-
1 0 t he Grou s e 8 to 1 4 the P a r t ridge 1 2 t o 2 0 th e ! u a il

, , ,
s till m or e s om etim es rea c hi ng tw o b r oo d s o f 7 t o 1 2 i n ea c h

,
.
H ere th e on ly e x cepti on t o the r el a ti on b et ween d ec r ea s i n g

b u lk a n d i ncrea s i ng nu mb er of eggs occu rs i n the c a ses of ,
t h e P h ea sa n t a nd the B l a ck coc k ; a nd it i s t o be r em em
-
b ered i n expl a na ti on th a t th e P h ea s a nt i s c ons tit u ti ona lly

, ,
a d a pt ed t o a w a rm er r egi on i s b ett er fe d— o ft en a rt i fi ci a l l y
,
a nd l e a d s a l es s a ctiv e lif e I f we p a s s t o d om es ti ca t ed
.
genera of the s a m e or d er we m eet wi th p a ra ll el di fferences

,
.
F rom the nu mb ers of eggs l a id littl e ca n be i n ferred ; for ,
u n d er the f a v ou ra bl e conditi on s a rtifi c i a lly m a i n t a i n ed t h e ,
l a yi ng i s c a rri ed on i nd efi nit ely Bu t th ou gh i n t he s i zes of

.
thei r b rood s th e T u rk ey a n d th e F ow l do not g rea tly differ ,
the F ow l b egi ns b ree di n g a t a m u c h ea rli er a ge t h a n the

T u rk ey a nd p r od u ces b rood s m ore f r equ en tly : a c ons id er
,
a bly high er ra t e of m u ltipli ca ti on b ei n g th e res u lt N ow .
th es e cont ra s t s a m ong d om es ti c crea t u res whi c h a re s imil a rly

c onditi oned a nd c l os ely a lli ed by cons titu ti on m a y be
,
-
,
h eld t o sh ow m ore cl ea r ly th a n m os t other cont ra s ts the

, ,
i nv ers e v a ri a ti on bet ween bu lk a nd s exu a l g enes i s ; si n ce

h ere the c os t of a ctivity i s dimi ni s h e d t o a c omp a ra tively
sm a ll a m oun t T h ere is littl e exp en dit u re i n flight s om e
.
-
tim es a lm os t none ; a nd the exp en dit u re i n wa lki ng a b ou t

i s n ot grea t : th ere is m ore of s t a n di n g th a n of a ct u a l
m ovem ent . I t i s t ru e th a t y ou ng T u r k eys c omm en ce
th ei r exi s t en ce a s l a rger m a s ses th a n chi ck en s ; bu t it i s
t ol era bly m a ni fes t th a t th e t ot a l weight of th e eggs l a id -
by a T u rk ey d u ri ng ea ch s ea s on b ea r s a l es s r a ti o t o th e ,
’
T u r k ey s weight th a n th e t ot a l w eight of th e egg s whi ch a
,
H en l a ys d u ri ng ea c h s ea s on b ea rs t o th e H en s weight ;
,
’
a n d thi s i s th e f a i r es t wa y o f m a ki n g t h e com p a ri s on T he .
comp a ri s on s o m a d e s h ows a
g rea t er di fferen c e th a n a pp ea r s
lik ely t o be due to the di fferent cos ts of l ocom oti on ; con
s ideri ng th e i nert nes s of the c r ea t u res Rem emb eri ng th a t .
456 L A WS OF MU L TI P L I CA T ION .
th e s i il a ting s u rfa ce i ncrea s es on ly a s the s qu a res of th e

a s m
dimen s i ons whil e th e m a ss o f th e fa b ri c t o be b ui l t u p by the

,
a b s or b e d nu t r i m en t i n c r ea ses a s t h e c u b es o f th e dim ens i o n s ,
it w ill be s ee n th a t the exp ense of g rowth b ec om es rel a tiv ely

g rea ter with ea c h i n crem ent of s i ze ; a n d th a t h ence of two ,
s imi l a r c r ea tu res c om m enc i n g li fe w i t h di f feren t s i zes t h e ,
l a rger one in rea ching i ts s u p eri or a d u l t b u lk will do t h i s a t ,
a m or e th a n p r op ort i on a t e e xp en se ; a n d s o will ei t h er be
d el a yed i n c om m enci ng i ts rep r od u cti on or w ill h a v e a ,
di m i n i sh ed reserv e for rep rodu cti on or b oth Oth er ord ers

,
.
o f B i r d s a c tiv e i n th ei r h a bit s s h o w m or e m a rk ed ly th e con

, ,
nexi on b et wee n a u g m en ti n g m a s s a n d d ecl i n i ng fer tility .
B u t i n th em th e i ncrea s i n g c os t of l oc o m o ti on b ec om es a n
i m p ort a nt a nd p rob a bly the m os t imp ort a nt f a ct or T he
, ,
.
e vid en ce th ey f urni s h w ill th eref ore c om e b ett er u nd er

a n o th er h ea d . C ont r a s ts a m on g Ma m m a l s lik e ,
th ose whi ch B i r ds p res ent h a v e th ei r m ea ni ngs ob scu red by

,
i nequ a liti es of th e exp endit u res for m oti on T he s m a ll er .
f ertility whi ch h a bit u a lly a ccomp a ni es g rea t er b u lk mu s t ,
i n a l l c a ses be p a rtly a scri be d t o thi s S till it m a y be

.
,
w ell if we b ri efly n ot e for a s m u ch a s th ey a re w orth

, ,
th e b roa d er c on t ra s t s .Whil e a l a rge Ma m m a l b ea rs bu t

a si ngl e y ou n g one a t a tim e i s s e v er a l y ea rs be fore it c om
,
m ences d o i ng thi s a n d th en rep ea t s th e rep rod u cti on a t l ong

,
i nt erv a l s ; we fi n d a s w e d escen d t o th e s m a ll er m emb ers cf

,
the cl a ss a v ery e a rly c omm encem ent of b ree di n g a n i ncrea s

, ,
i ng nu m ber a t a bi r th r ea chi ng i n s m a ll Rod ent s t o 1 0 or

,
e v en m ore a nd a m u c h m or e f req u en t r ec u rren c e of b ro od s

,
th e combi ned resu lt b ei ng a rel a tiv ely p r odigi ou s fer tility .
I f a sp ec ific c om p a ri s on be d es i red b et ween Ma m m a l s th a t

a r e s i m il a r i n c on stit u ti on i n f ood i n c onditi on s o f life a n d
, , ,
a l l oth er thin g s bu t si ze th e D eer t rib e s u ppli e s it

,
-
Wh il e .
th e l a rge Red d ee r h a s bu t one a t a bi rth the sm a ll Roe d eer

-
,
-
*
h a s f requ ently two a t a bi r th .
§ 341 . T he t g on i s m be tw een gr owt h

an a a n d se u a x l g ene
A p a s sage t ra ns l a ted for m e from t he Ge rm a n m ay be here gi v en i n
GROWT H A N D SE X U A L GE NE SIS .
457
sis visibl e i n th es e genera l c ont ra st s m a y a l s o be t ra c ed i n the

, ,
hi s t ory of ea c h pl a nt a nd a nim a l S o f a mili a r i s th e f a ct .
th a t sexu a l genes i s d oes n ot occu r ea rly i n li fe a nd i n a l l ,
org a n i s m s w hi c h e xp en d m u ch b e gi n s on ly wh en the li m it of
s ize i s n ea rly rea ch ed th a t we do n ot su ffi ci ent ly not e its

,
signific a nc e I t is a genera l phy s i ol ogi ca l t ru th h owev er

.
, ,
tha t whil e the bu ildi ng u p of the i ndividu a l i s g oin g on

-
ra pidly the rep rod u c tiv e org a ns rem a i n imp er f ectly d ev e

,
l op ed a nd i na ctiv e ; a nd th a t the c om m encement of rep r o

d u cti on a t onc e i n di ca t es a d ecli nin g r a t e of g rowth a nd ,
be c om es a c a u s e of a rres ti ng gr owth A s wa s s h own i n 7 8 .

,
t he ex c epti on s t o t hi s ru l e a re fou nd wh ere the limit o f

g r owth i s i nd efinit e ; eith er b ec a u s e th e org a n i s m exp end s
littl e or n othi ng i n a cti on or exp en d s i n a cti on s o m od era t e
,
a n a m ou nt th a t th e s u pply of nu t rim ent i s ne v er eq u ilib ra t e d
by i ts exp enditu re .
We w ill p a ss over the i nferi or pl a nts an d li m iti ng ,
ou rs elv e s t o P haen og a m s will not d well on the l es s con

,
s p i cu ou s evid en c e w ith t he s m a ll er typ es p res en t A few .
c a s es su ch a s ga r dens s u pply will s erve A l l k n ow th a t .
a P ea r t ree i n crea s es i n si ze for yea r s b ef or e it b egi ns t o

-
b ea r ; a nd th a t p r odu cin g bu t few p ea rs a t fi r st it i s l on g

, ,
b efore it f ru it s a b u nd a ntly A y ou ng Mu lb erry t ree b r an ch .

-
,
i ng ou t l u x u r i a ntly s ea s on a ft er s ea s on bu t c ov ered w ith ,
n othi n g bu t l ea v es a t l en gth bl os s o m s s p a ri n gly a n d s et s

,
s o m e sm a ll a n d imp er f ec t be r r i es w hi c h it d r op s w hil e th ey ,
a re green ; a nd it m a k e s th es e fu til e a tt empts tim e a ft er
verificati on . Dr . D i onys Hel l in i n an ess ay on the ori gi n of Mul ti p ari ty

a nd Twi n b i rth s ,
-
re fe rs to th e th es i s a bo ve s et forth , a nd s ay s th a t “
th e
fac t th a t i t i s ge nera l l y w om en of s m all growth who h ea r t w i ns i s i n co m

”
p l e te a g r eem e nt wi t h i t He ad d s t h a t
. P u ec h i s ri ht i n h i s op i ni on th at
g
t w in p re gna nc i e s a re a di rect re s ul t of rel a ti v el y l a rge ov a ri e s i n com
p a ri s on w ith th e W hol e body) He h a s ob s er ed th a t for th e

. v s a m e s i z e of
v
body the o a ri um of a pl u ri p a rou s a ni m al i s a l wa ys of gre a t er v ol um e th a n
th a t of a uni p a rou s ani m a l a s ow ha s v

o a ri e s as l a rge a s a co w s
’
;
b u t whi l e th e l a tte r bea rs onl y one cal f [at a t im e ] ,
th e sow bri ngs forth
6 — 1 5 a t ea ch l i tt er E v en in a ni m al s of th e
.
sam e s pe c i e s b ut bel onging to
diffe rent ra ce s th ese rel a t i ons m ay be v eri fied ,
”
eg
. .
,
Ba rb a ry sheep and ordi
na ry sh ee p .
458 L A WS or MU L TI P L I CA T I ON .
tim e be f ore it s u ccee d s i n r ip enin g a ny s eeds B ut thes e .
mu lti a xi a l pl a nt s or a ggrega t es of i ndivid u a l s s ome of

-
,
whi c h c onti nu e t o g row w hil e o th ers be c om e a rr es t e d a n d

t ra ns f orm ed i nt o s eed b ea rer s s h ow u s th e rel a ti on l ess de
-
,
fi nit ely th a n c er t a i n pl a nt s th a t a re s u b s t a nt i a lly if n ot ,
lit era lly u ni a xi a l

,
-
Of th ese th e C ocoa nu t m a y be i n
.
-
s t a n c ed F or s om e y ea r s it g oes on s h oot i n g u p with ou t

.
m a ki n g any s ign of b ecomi ng f ertil e A b ou t the s i xth y ea r .
it fl owers bu t the fl owers wi th er with ou t resu lt I n the .
s ev enth y ea r it fl owers a nd p ro du c es a few nu t s ; bu t th ese

p rov e a b ortiv e a nd d rop I n the eighth yea r it rip ens a .
m odera t e nu m b er of nu t s ; a nd a ft erwa r d s i nc rea ses the

nu m b er u ntil i n th e t enth y ea r it co m es i nt o f u ll b ea r i n g
, ,
.
Mea nwhil e f rom th e tim e of its fi rs t fl oweri n g its gr ow th

,
b egi n s t o dimi ni s h a nd g oes on di m i ni s hi ng till th e t enth

,
yea r wh en it cea ses H ere we s ee th e a nt a g oni s m b et ween

,
.
g rowth a nd s exu a l g enes is un d er b oth i ts a s p ect s— see a

st ru ggle b et w een s el f ev olu ti on a n d ra c e ev ol u ti on i n whi c h
- -
,
th e fi rs t for a tim e o v erco m e s th e l a s t a n d th e l a s t u lti m a t ely ,
o v erc om es th e fi r s t T h e c onti nu ed a ggr a ndi ze m ent of th e

.
p a rent i ndivid u a l m a k es a b or tiv e for tw o s ea s on s th e t en d ency

-
t o p r od u c e new i n divid u a l s ; a n d th e t end ency t o p r od u ce

new i ndivid u a l s be comi ng m or e d ecid ed s t op s a ny f u rth er
, ,
a gg ra n di zem en t o f t he p a r en t i ndivid u a l .
P a ra ll el ill u s t ra ti ons o ccu r i n th e a n im a l ki ngd om T he .
egg s l a id by a p u ll et a r e rel a tiv ely sm a ll a nd few S i m il a rly .

,
i t i s a ll ege d th a t a s a gener a l ru l e
, a bit ch h a s f ew er ,
“
”
p u ppi es a t fi r s t th a n a ft erwa rd s
, A ccordi ng t o B u r d a ch .
,
a s q u o t ed by D r D u nca n
“
. the el k the b ea r &c h a v e a t
, , ,
.
,
fi rs t only a si n gl e y ou n g one th en they c om e t o h a v e m os t ,
frequ en tly tw o a nd a t l a s t a ga i n on ly one

, T h e y ou n g .
h a m s t er p rod u c es only f rom th ree t o s i x y ou ng ones whil e ,
t h a t of a m ore a dv a n ced a g e p ro d u c es fro m eight t o s ixt een .
”
T he s a m e i s t ru e of th e pig I t i s rem a rked by B u ffon th a t
. . .
wh en a s ow of l es s th a n a yea r ol d h a s y ou ng th e nu m b er of ,
'
th e litt er is sm a ll a nd its m em bers a re f eebl e a nd ev en im

,
GROWT H A ND S E X UA L GE NE SIS .
9
p er fect .H ere we h a v e evid ence th a t i n a nim a l s g rowth '
c h ecks s ex u a l g enes i s A nd th en on th e oth er h a nd we

.
, ,
h a v e evid en ce th a t s exu a l genes is ch ecks growth I t i s w ell .
k nown t o b reeders th a t if a filly i s a ll owed t o bea r a f oa l ,
s h e i s th ereby p re v ent e d f r om r ea c hi ng h er p r op er s i z e A nd .
a lik e l os s o f p er f ecti on a s a n i ndivid u a l i s s u ffered by a ,
c ow whi c h b reed s t oo ea r ly I t m a y be a dded a s a c on v erse

.
,
f a ct th a t c a s t ra t ed a nim a l s a s c a p ons a n d not a bly c a t s

, , ,
o ft en b ec om e l a r g er th a n th ei r u n m u til a t ed a ss oci a t e s .
§ 342 . N ot with s t a ndi ng the w a y in which th e i nv er s e

v a ri a ti on of g r owth a nd s exu a l genes i s is c omplic a t e d with
o th er rel a ti ons i ts exi s t en c e i s I thi nk s uffi c i ently m a n i

, , ,
f es t I ndivid u a lly m a ny of the f oreg oi ng i ns t a nc es a re op en

.
,
t o c ritici s m a n d h a v e t o be t a k en w ith qu a lifi ca ti ons ; bu t

,
wh en l ook ed a t i n th e m a s s th ei r m ea ni ng is b ey on d d ou bt .
C om p a r i s on s b etween the l a rges t w ith the s m a ll es t typ es ,
wh eth er veget a l or a ni m a l yi eld resu lt s w hich a re u nm i s

,
t a k a bl e On the one h a nd rem em b eri ng th e f a c t th a t du r

.
,
i ng its c en t u r i es o f life a n Oa k d oes n ot p r odu c e a s m a ny

a c orn s a s a F u n gu s d oes s p ores i n a s i ngl e night we see th a t ,
th e F ungu s h a s a fertility exc eedi ng th a t of th e Oa k i n a

d egree lit era lly b ey ond ou r p owers of c a lc ul a ti on or im a gi na
ti on . On th e oth er h a nd wh en t a ki n g a mic roscopic p ro ,
t ophyt e whi c h h a s billi on s of d es c end a n t s i n a few d a y s ,
we a s k h ow m a n y s u ch w ou ld be requ i r ed t o b u ild u p th e
f ores t t ree whi c h i s yea rs b ef ore it d rop s a s eed we a re m et by ,
a p a r a ll el diffi cu lty i n conce ivi n g th e nu m b er if n ot i n s e tti ng ,
it d own S imil a rly if f rom the mi nu t e a n d p rodigi ou s ly

.
,
fer til e Roti fer we t urn t o the E l eph a n t whi c h a pp roa ch es ,
thi rt y yea rs b ef ore it b ea r s a s olit a ry y ou n g one we fi nd the ,
c onnexi on s b et w een s m a ll s i ze a nd g r ea t f ertility a n d b et ween
grea t size an d sm a ll fertility t oo i n t en sely m a r k ed t o be ,
m u ch di s g ui s ed by th e p ert u rbi n g rel a ti ons th a t h a v e b een

i ndi ca t ed F i na l l v a s thi s i nd u cti on r ea ch ed by a su rv ey of
.
, ,
org a nis m s i n genera l is v erifi ed by ob s er v a ti on s on th e rel a

,
t i on b et ween d ec rea s in g g rowt h a n d c om mencin g reprodu c

ti on in i ndi vid u a l or ga ni s m s we m ay I thi nk con sid er the , , ,
*
a ll eg ed a n t a g on i s m a s p r ov ed .
W h en , a ft er h av i ng h el d for s om e yea rs t he gene ra l doct rine el a bora ted

i n th es e c h a pte rs , I a greed ,
e arl y in 1 8 5 2 , t o p rep a re a n ou tl i ne of i t for t h e
Westm i ns ter Revi ew, I cons ul te d , am ong o th e r w ork s , the j us t -i ss u e d t h i rd
D r Ca rpe nte r s P r i ncip l es P hysi ol ogy, Gener a l
‘
ed i ti on of . o
f a nd Com p a r a
ti ve— s eek i ng
i n i t for fact s il l us t ra ti ng th e di fferent degrees of fe r til i ty of
di ffere nt organi s m s , I m e t wi th a p a ss a ge, q u ot ed ab o e i n v 3 3 9 , w hi c h

se em ed ta c i t l y to a ss ert t ha t i ndi v i dual a gg randi z e m ent is at v a ri a nc e wi th
t he p ro pa ga ti on o f t he ra ce ; b ut nowh ere found a di st i nct e nun c i a t i on of
th i s t ru th . I di d not then rea d th e Cha pte r ent i t l ed Genera l Vi e w of t h e
Functi ons, ”
whi ch h el d out n o p rom i s e of s u ch e v idence as I wa s l ooki ng for .
B ut on s i nce re fe rri ng to thi s ch ap te r, I di s cov e red in i t t h e de fi ni t e s ta te
m en t t h a t the re i s a ce rt ai n degree of an ta g oni s m b et we e n the N utri ti v e

and Rep rodu c ti ve funct i ons , the one bei ng e x e cuted at x
t h e e pens e o f the
othe r. The r ep roducti ve a pp a ra t u s d eri ve s t he m a te ri a l s of i ts op era ti ons
throug h th e nu tri ti ve sys te m ,

a nd is enti rel y de p e ndent up on i t for t h e co n
ti nua nce of i ts fu nc t i on . If, th ere fore, i t be i n a s ta te of e x ce s s i v e ac ti v i ty

,
i t wil l nece s sa ri l
y d ra w of
f from th e i ndi idua l v fa b ri c
p or ti on of t h e s om e
al im ent d es t i ned fo r i ts m a i nte nance It m a y b e uni v ers al l y ob s e rve d t ha t ,

.
wh e n the nu tri ti ve fun c ti ons a re p art icul a rl y a c ti v e i n s upporti ng th e i ndi

vidu a l , the rep rod ucti v e s ys te m i s in a corres pondi ng degree v
un de el op ed,
”
and vi ce versd. P 5 92
. .
C H A P TE R VI I .
THE O I
A N T A G N S M B E T W E E N D E VE L OP M E N T A N D GE N E S S, I
A SE X UA L AND SE X UA L .
BY D ev el opm ent , h er e t o be d ea lt w ith a p a rt

§
343 . as
f rom Growth is m ea nt i ncrea s e of s t ruc t u re a s di s ti ngu i s h ed

,
from i n crea s e o f m a s s A s wa s p oi nt ed ou t i n § 5 0 thi s is

.
,
t he bi ol ogic a l d efi niti on of th e w or d I n th e f oll ow i n g s ec

.
ti on s then we h a v e t o not e how c ompl exity of orga ni z a ti on

, ,
i s hin d ere d by rep ro d u ctiv e a ctivity a nd c on v er s ely,

.
T hi s r el a ti on p a rti a lly c oi ncid es with th a t w hich we h a v e

j u st c ont empl a t ed ; for a s wa s s h own i n § 4 4 d egree of
, ,
g rowth i s to a c ons id era bl e ext en t d ep end en t on d egree of

o r g a n i z a t i on B u t whil e the a nt a g on i sm to be ill u s t ra t ed i n
.
this c h a pt er i s mu ch en t a n gl ed with th a t ill u s t ra t e d i n the

l a s t ch a pt er it m ay be s o fa r s ep a ra t ed a s t o be id entifi ed a s
,
a n a dditi on a l a n t a g on i s m .
Bes id es the di rect opp ositi on b et ween th a t c on ti nu a l di s

i nt egra ti on whi ch ra pid genes i s impli e s a nd the fu lfil m ent of ,
t h a t p r e—r equ i s it e to e xt en s iv e org a ni za ti on— th e f orm a ti on of

a n ext en s iv e a gg reg a t e th er e i s a n i n di rect opp os i t i on w hich
,
we m a y recog ni ze u n d er s ev era l a s p ects T h e ch a n ge

.
fro m h om ogeneity t o h et erogeneity t a k e s tim e ; a nd tim e

t a k en i n t ra ns f orm i ng a rel a tively s t ru ct u rel es s m a ss i nt o a
-
d ev el op e d i ndivid u a l d el a ys the p eri o d of rep ro du cti on U su

, .
a lly thi s tim e i s m erged i n th a t t a k en for g r owth ; bu t c ert a i n
ca ses o f m et a m orph os is s h ow u s t h e one s ep a ra t e f rom t he
o t h er . A n ins ect p a s s i ng f rom its l owly orga ni zed c a t er

,
-
46 1
462 L A WS OF MU L T I P L I CA T I ON .
pill a r s t a ge i nt o th a t of ch ry sa li s i s a ft erwa r d s a week a

-
, ,
fortnight or a l onger p er i o d i n c om pl eti ng i t s st ru c t u re : th e

,
r e com m enc em ent of g en e s i s b ei n g by s o m u c h p os t p on e d

-
,
a n d th e r a t e o f m u ltipli c a ti on th erefore dimi n i s h e d F u rther .

,
th a t re a rra ngem ent of s u b s t a n ce whi ch d ev el op m en t i m pli es

-
,
en t a il s exp en dit u r e T he ch ry sa li s l oses weight i n th e c ou rs e

.
o f i t s t r a n s f or m a ti on ; a n d th a t i t s l o s s i s n ot l o s s o f w a t er
on ly m a y be i n ferre d f ro m th e f a ct th a t it res pi res a n d th a t

, ,
r es pi r a t i on i n di ca t es cons u m pti on C l ea r ly t he m a tt er c on
.
s u m ed i s oth er thi n g s equ a l a d ed u cti on f r o m the s u rpl u s

'
, ,
wh i ch m a y g o t o rep r od u c ti on Y et a g a i n th e m ore
.
,
wid ely a nd c om pl et ely a n org a ni c m a s s b ec om es di fferen ti a t ed ,
th e s m a ll er i s t h e p ort i on of it whi c h ret a i ns the r el a tiv ely

u n di fferen ti a t e d s t a t e th a t a d m it s o f b ei n g m ou ld e d i nt o new
i n dividu a l s or th e germ s o f th em
,
P rot opl a s m w hi ch h a s
.
b ec om e s p eci a li zed ti ssu e c a nnot be gen era li zed a fres h a n d ,
a ft erw a r d s t ra n s f or m ed i n t o s o m e thi ng e l s e ; a nd h en ce th e
p rog ress of s tru ct u re i n a n org a ni s m by di m i ni s hi ng the ,
u n s t ru ct u re d p a r t dimi ni s h es the a m ou n t a v a il a bl e for m a k

,
ing o ffs p ri ng .
I t i s t ru e th a t higher s t ru c tu re lik e grea t er growth m ay

, ,
i ns u re t o a s p eci es a dv a n t a g es whi ch ev ent u a lly f u r ther its

m u ltipli ca ti on— m a y giv e it a cces s t o l a rger su ppli es of f ood ,
or en a bl e it t o o b t a i n f oo d m or e ec on omi c a lly ; a n d w e s h a ll
h erea ft er s ee h ow th e i nv erse v a ri a ti on we a re c ons id eri ng i s

th u s q u a lifi ed Bu t h ere we a r e con cerned on ly with th e
.
neces s a ry a nd di rect ef fect s ; n ot w ith t h ose th a t a re c on

ti ng ent a nd rem ot e T h es e neces s a ry a nd di rect effect s we
.
will now l ook a t a s exemplified .
§ 3 44 . S p eaki n g
g en er a lly t h e s impl er pl a nt s p r op a g a t e
,
b oth s exu a lly a nd a sex u a lly ; a nd s p ea ki n g c omp a ra tiv ely , ,
th e compl ex pl a n t s p rop a ga t e on ly s exu a lly : th ei r a s exu a l

r p a g a ti on i s u s u a l ly i n c ompl et e —p rod u c es a u ni t ed a ggr e
p o
g a te of i ndivid u a l s i ns t ea d of nu m erou s di s ti nct i ndivid u a l s .
T he P rot ophyt es th a t p erp et u a lly s u bdi vi d e, the m erely

‘
.
D E VE L OP M E NT A ND GE NE S IS .
4 63
ce llu l a r A l gae th a t sh ed thei r t et ra s p ores the A rch eg oni a t es ,
t h a t s p on ta n eou s ly s epa r a te t h ei r f rond s or d rop th ei r g em

m ae s h o w u s a n e xt ra m o d e of mu ltipli ca ti on w hich a m ong
, ,
fl ow eri ng pl a nt s i s ex cepti ona l T his ext ra m od e of m ul tipli

,
.
c a ti on a m on g th es e s i m pl er pl an ts i s m a d e ea s y by th ei r l ow
,
d evel opm ent T etra s p ores a ri s e onl y wh ere the f ron d c on

.
s i s t s o f u nt r a n s for m ed cell s ; gem m e bu d ou t a nd d rop of f

'
c
only w h ere t he ti s s u e i s c o mp a ra tiv e ly h o m og eneou s .
S h ou ld it be s a id th a t this i s bu t a n oth er a s pe ct of the

a n t a g oni s m a l rea dy s et f o rth s i nce th es e u nd e v e l op ed f or m s
,
a re a ls o t h e s m a ll er form s ; the r eply i s th a t th ou gh i n p a r t
t ru e t hi s i s not w h olly t ru e Va ri ou s m a ri ne A l gee which

.
p r op a ga t e a sex u a lly a re l a rger th a n s om e P hae nog a m s whi c h

,
do n ot th u s p r op a g a t e T he obj e c t i on th a t diff erenc e of

.
m edi u m viti a t es thi s com p a ris on i s m et by the fa ct th a t it i s

,
the s a m e a m ong l a n d pl a nt s them s elv es

-
S und ry of the l ow .
l y or g a ni z ed L iv erw or t s whi ch a re h a bit u a lly gemmip a rou s

-
,
e xc eed i n s iz e m a ny fl oweri n g pl a nt s A nd th e F ern s s h ow

.
u s a g a mi c m u ltiplic a ti on occ u rr i n g i n pl a n t s w hich whi l e ,
th ey a re i nferi or i n c o m pl exi ty of s t ru ct u re a re s u p eri or i n ,
b u l k t o nu mb ers of a nnu a l Monoc otyled ons a n d D i cotyl ed ons .
345 . bility of th e l owly orga ni zed s u bst a nc e of a

I n th e a -
S p on ge t o t ra n s f orm it s elf i nt o mu ltit u d es of gemmu l es we ,
h a v e a n i ns t a nce of this s a m e di rect rel a ti on i n th e a nim a l

ki ngd om Moreov er the i ns t an ce yi eld s v ery di s ti nct p roof
.
,
o f a n a nt a g oni s m b et wee n d ev el op m ent a n d g enes i s i nd e ,
pend ent of the a nt a goni s m be t ween gr owth a nd genes i s ;

for th e S p onge w hich th u s m u ltipli es it self a s exu a lly a s well ,
a s s exu a lly i s fa r l a r g er th a n h os t s of m ore c o mpl ex a n i m a l s

,
whi ch do not mu l t iply a sexu a lly

’
Onc e a g a i n m a y be ci ted th e crea t u re s o oft en b rou ght i n

evid enc e th e H ydr a
,
a s s h ow i ng u s h ow r a pidity o f a g a mic
,
p r op a g a ti on i s a ss oci a t ed with i n feri or ity o f s t ruct u re I ts .
w t p o d u ce y ou n g o nes f r om n ea r ly a l l p a rt s o f i ts
p o er o r
b ody i s du e t o the c omp a ra tiv e h om ogeneity of its b ody I n

, .
4 64 L A WS OF MUL T I P L ICA T I ON .
ki n d re d bu t m ore orga n i zed typ es th e gemmip ar ity i s

-
,
gr ea tly res t r i ct ed or di sa pp ea r s A m ong th e free swi m m i ng

,
.
-
H y dr ozoa m u l t ipli ca ti on by bu ddi n g w h en it occu r s a t a l l

, , ,
o c cu r s only a t s p ec i a l pl a c es T h a t i ncrea s e o f s t ru ct u re
.
a p a rt f rom i n c r ea se o f s i ze i s here a c a u se o f d ec li ni n g ag a m o
,
gen es is we m a y s ee i n th e cont ra s t b etw een the s impl e H ydra

,
a nd th e c omp ou n d H yd ro id s T h ese l a s t a l on g with m ore

.
,
di fferen ti a t e d ti ssu es s h ow u s a gem m a ti on which d oes n ot

,
n a l l o v er t h e b ody of ea ch p olyp e a nd m u c h of it d oes

g o o ,
not en d i n s ep a ra ti on .
I t i s h owev er a m ong th e A nnu l os a th a t p ro gress i ng

, ,
org a ni z a ti on i s m os t cons pi cu ou s ly op era tiv e i n dimin i s hi n g

”
a ga m og enes i s T h e segm en ts or
. s om it es c omp os i ng a n
a nim a l b el on gi n g t o thi s c l a s s a re p ri m o rdi a lly a lik e ; a n d
, ,
a s b ef or e a r gu e d 2 05 a re p r ob a bly th e h om ol o gu es o f
wh a t were origi na lly i nd ep end en t i ndividu a l s T he p rogress .
f rom the l ower t o th e high er typ es of th e cl a s s i s a t on ce a ,
p rog res s t owa r d s typ es i n which the s t ri ng s o f s egm en t s c ea s e

t o u nd erg o s u bdivi s i on a n d t ow a r d s typ es i n whi c h th e s eg
,
m en t s no l onger a lik e i n th ei r s t ru ct u res a nd fu ncti ons h a v e

, ,
b ecom e phy s i ol ogi ca lly i nt egr a t e d or m u t u a lly d ep end en t .
A l rea dy thi s g rou p of ca ses h a s b een n a m ed a s ill u s t ra ti ng

t he a nt a g on i s m b et ween g rowth a n d a se xu a l genesis ; bu t it
i s p rop er a l so t o n a m e it h ere s i nce on th e on e h a n d the , , ,
g rea t er s i ze du e t o th e cea s i ng of fi s s i on is m a d e p os s ibl e ,
o nly by t h e s p ec i a li z a ti on of p a rt s a nd th e d ev el op m en t of a
c o or di n a ti ng a pp a r a t u s t o combi ne th ei r a c ti ons a n d si nc e
-
, ,
on th e oth er h a n d s p ec i a li z a ti on a nd c c or di na ti on c a n a d
,
-
v a nc e only i n p rop orti on a s fi ss i on c ea s es .
§ 3 46 . Th e
i nv ers e v a ri a ti on of d ev el opm ent a nd s exu a l
genes i s i s by no m ean s ea s y t o f oll ow One or two f a cts i ndi .
ca tiv e of it m a h o w e v er be n a m e d
y , , .
P haenog a ms th a t h a v e bu t littl e su pp orti n g ti s s u e m a y

f a i rly be cl a s s ed a s s t ru ct u ra lly i n feri or t o th os e h a vi ng s t ems
w ith a b u lky a nd c om pl ex w oody s y s t em ; for t h es e imply
"
D E V E L OP ME NT A N D GE NE SIS .
465
a dditi ona l diff erenti a ti ons a nd c on stitu t e wid er d ep a rt u res

,
f rom the p r i m itiv e typ e o f v eget a l ti s s u e T h a t the c on .
c om it a n t o f thi s high er o rg a ni z a ti on is a sl ower g a m o genes i s ,
s c a r cely n eed s p oi n ti n g ou t Whil e the h erb a c eou s a nnu a l

.
i s bl o s s omi n g a n d rip en i n g s eed th e y ou ng t ree i s t ra ns f or m ,
i ng i t s origi na lly su ccu l ent a xi s i nt o d en s e fib r ou s s u b s t a nc e ;

-
a n d y ea r by y ea r t h e y ou n g t ree e xp en d s i n d oi n g th e lik e ,
nu t r i m en t w hich s u cc es s iv e g enera ti on s o f th e a nnu a l exp en d
i n f ru it . H ere the i nv er s e r el a ti on i s b et wee n s exu a l rep ro

d u cti on a nd c om pl exity a nd not be t w een s exu a l r ep r od u cti on
,
a nd b u lk s eei ng th a t be sid es s ee di n g th e a nnu a l oft en grows

, ,
t o a s i ze g rea t er th a n th a t rea ch e d by the y ou ng i n f ertil e t ree

i n s ev era l yea rs .
P roo f of th e a nt a gon i s m b e t ween c ompl exity a nd ga m o

g enes i s i n a nim a l s is s till m ore difficu lt t o di sent a ngl e
,
.
P erh a p s the evid ence m os t t o th e p oin t i s f u rn i s h ed by the

c ont ra s t b et ween Ma n a nd c ert a i n oth er Ma mm a l s a pp roa ch
i ng him i n m a s s T o c om p a re h i m w ith the d om es tic
.
S h eep which th ou gh n ot v e ry u nlik e i n s i ze i s rel a tiv ely

, ,
p r olific is obj ec ti ona bl e bec au s e of the rel a tiv e i na ctivity of

,
S h eep ; a n d thi s t oo m a y be a ll ege d a s a rea s on why th e Ox

, , ,
th ou gh fa r m ore b u lky i s a l s o fa r m ore fertil e th a n Ma n

, ,
.
F u rther a ga in s t a com p a r i s on with the H ors e whi ch whil e

, ,
b oth l a rger a n d m ore p rolifi c i s t ol era bly a ctive it m a y be , ,
u r ged th a t i n hi s c a se a nd th e c a ses o f h erbiv or ou s crea t u r es

,
g enera lly the s m a ll ex erti on req u i red to p roc u re f ood j oi ned

, ,
w ith th e gr ea t ra ti o b orne by th e a li m ent a ry or g a n s t o th e

or g a n s th e y h a v e t o b u ild u p a nd r ep a i r viti a t es t h e res u lt ,
.
We m a y h owev er fa i r ly d ra w a p a ra ll el b et ween Man a nd a

, ,
l a rge ca rn iv ore The L i on s u p er i or i n s i ze a n d p erh a p s

.
, ,
equ a l i n a ctivity h a s a dig es tiv e s y s t em n ot p r o p orti o na t ely

,
g rea t er ; a nd yet h a s a high er r a t e of m u ltiplic a ti on th a n

Ma n H ere the on ly d ecid ed w a n t of p a r ity b es id es th a t of
.
,
o rg a n i za ti on i s t h a t of f oo d
,
P os s ibly a c a rniv ore g a i n s a n
.
a dv a n t a g e i n h a vi n g a s u rpl u s nu t r i m ent c on si s ti n g a lm os t
wh olly of th o s e nit rogenou s m a t eri a ls from which the bodi es

76
466 L A WS OF MUL TI P L ICA T ION .
’
of y ou ng one s a re m a i nly f orm ed B u t a ll o wi ng for a l l other

.
,
di fferenc es it a pp ea rs n ot i m p rob a bl e t h a t t he s m a ll ness o f

,
hu m a n fertility c om p a red wi t h the fert ility of l a rge feli n e

a nim a l s ,
i s du e t o t he g rea t er c om pl exi ty o f the h u m a n
o r g a n i z a ti on— m or e es p ec i a lly th e or g a n iz a t i on of th e n erv ou s
sy s t em T a ki n g d eg ree of nerv ou s orga ni za ti on a s the c hi ef

.
corre l a tiv e of m ent a l c a p a city ; a nd re m emb er in g th e phy s i o
l ogi ca l cos t of th a t sl ow ev olu ti on wh ereby high m ent a l

c a p a c i ty i s r ea c h ed ; w e m a y s u s p ect th a t n erv ou s org a n iz a
ti on i s v ery exp en s iv e : the i nf erenc e b ei ng th a t b ri ngi n g it

u p t o th e l ev el it r ea c h es i n Ma n w h o se dig es t iv e s y s t e m by
, ,
n o m ea ns l a rg e h a s a t th e s a m e tim e t o s u pply m a t eri a l s for

,
g enera l g rowth a nd d a ily wa s t e i nv olves a grea t ret a rd a ti on

,
o f m a t u r ity a n d s ex u a l g enesis .
C H A P T E R VI I I .
A N T A GON S M I BE T WE E N E X P E N D T I U RE A ND GE N E S S I .
347 . thi s h ea d we h a v e t o set d own n o evid ence

UN D E R
d eriv ed f rom th e v eg et a l ki ngd om P l a nt s a re not exp end ers
.
o f forc e i n s u ch d eg rees a s t o a f fect th e genera l rel a ti ons wi th

w hich we a re d ea li ng T h ey h a v e n ot t o m a i n t a in a h ea t
.
a b ov e th a t o f th ei r envi ron m en t n or h a v e th ey t o g enera t e

,
m oti on ; and h ence c ons u mpti on for th ese two p u rp os e s d oes

n ot dimi ni s h th e s t oc k o f m a t er i a l which s er v e s on the one
h a nd for grow th a nd on the oth er h a n d for p rop a ga ti on .
I t will be well t oo if we p a s s o v er th e l ower a n im a ls

, ,
e s p ec i a lly th os e a qu a tic one s which bei n g nea rly of the ,
s a m e t empe r a tu re a s the w a t er a n d nea rly of th e s a m e ,
s pec ific gra vity l os e bu t littl e i n ev olvin g m oti on s ens ibl e

, ,
a n d i nsen s ibl e A fu rth er rea s on for ex c l u di n g f rom con

.
sid era ti on t h es e i n feri or typ es i s th a t we do not kn ow enou gh

,
o f th ei r ra t es o f g enesis t o p ermit of ou r m a ki n g wi th a ny ,
s a ti s fa cti on th os e i nv olv ed c omp a ri s on s h ere to be entered

,
u pon .
T he fa cts on m u st m a i nly d ep end a re th os e t o be

which w e
th d f t t i a l a nim a l s a nd chi efly f rom th os e
‘
g a er e ro m er r es r ,
high er cl a s ses of th em whi ch a r e a t the s a m e tim e gr ea t

e xp en d er s a nd h a v e r a t es o f m u ltiplic a ti on a b ou t w hich ou r
kn ow l edge i s t ol era bly d efi nit e We will res t ri ct ou rs elv es

.
,
th en t o the evi d ence which B i r ds a nd Ma mm a ls s u pply

, .
§ 3 48 . S a ti s f a ct ory p roof th a t l os s of s ubst anc e i n the

467
4 68 L A WS OF MUL T IP L I CA TI ON .
m a i nt ena n c e of h ea t d1 m 1 n1 sh es th e ra pidity of p rop a g a ti on ,
i s di ffi cu lt t o obt a i n I t i s i n d eed obvi ou s th a t the w a r m

.
, ,
bl ood ed Ver tebra ta a re l es s p rolific th a n the cold bl ood ed ; -
bu t t h en t h ey a r e a t the s a m e ti m e m or e viv a ci ou s S im i .
l a rl y b et w een Ma m m a l s a n d B i r d s ( w hi c h a re th e wa rm er
,
bl ood ed of the two ) th ere i s oth er thi n gs equ a l a p a ra ll el , , ,
th ou gh m u c h s m a ll er differenc e ; bu t h ere t oo the u nlik e

, , ,
n esses of m u s cu l a r a cti on c om pli ca t e t h e evid en ce A ga in .

,
th e a nnu a l ret u rn of genera t iv e a c t ivi ty h a s a n a v era ge cor

res p on d en c e w ith t h e a nnu a l r et u r n o f a w a r m er sea son ,
’
w hi ch did it s t a n d a l on e might be t a k en a s evid enc e th a t a
, ,
dimi n i s h ed c os t of h ea t m a i n t en a n ce l ea ds t o s u ch a su r pl u s
-
a s m a k es rep ro d u cti on p oss ibl e B u t th en thi s p er i odi c ris e .

,
o f t em p er a t u r e is h a bit u a lly a ccom p a n i ed by a n i n c r ea s e i n
t h e qu a n tity of f ood— a fa ct or of equ a l or gr ea t er i m p ort a n ce .
We m u s t be cont ent th erefore with s u ch few s p eci a l fa cts a s

, ,
a dmit of b ei ng dis en t a n gl e d .
Cert a i n of th es e we a re i nt rod u c e d t o by th e g enera l r el a

ti on l a s t n a m ed— th e h a bit u a l recu rr enc e of genesis with the
r ecu rren c e of s p r i n g F or i n s om e c a ses a d om es tic a t ed crea
.
t u r e h a s i t s s u ppli es of f ood a lm os t equ a li zed ; a n d hence the

ef fec t of v a ryi n g n u t riti on m a y be i n g rea t p a r t eli m i n a t ed
f rom the c om p a ri s on T he com m on F owl yi eld s a n ill u s t ra
.
ti on I t i s fed th rou gh the c old m onth s bu t nev erth el es s i n

.
, ,
mid w i nt er it eith er wh olly l ea v es off l a yi ng or l a y s v ery

-
,
s p a r i ngl y A nd th en we h a v e the fu rth er evid ence th a t if it

.
l a y s s p a r i ngly it d oes s o on ly on c onditi on t h a t the h ea t a s

, ,
well a s th e foo d i s a r tifi ci a lly m a i nt a i ned

,
H ens l a y i n c old .
wea th er only w h en th ey a re k ept w a r m T o whi c h f a ct m a y .
“
be a dd ed the ki nd red on e th a t wh en pig eons r eceiv e a rti
fi ci a l h ea t th ey n ot only c onti nu e t o h a tch l on ger i n a u t u m n
, ,
bu t will r ec om m enc e i n sp r i n g s oon er th a n th ey w ou ld oth er

”
w i s e do . A n a na l og ou s pi ece of evid ence is th a t i n ,
wi nt er i n a d equ a t ely s h elt ered C ows eith er cea s e t o giv e m ilk

,
or giv e it i n dimi ni s h ed q u a n tity F or th ou gh givi n g milk .
i s n ot th e s a m e thi ng a s b ea ri ng a y oun g on e yet a s milk , ,

E X P E N D I T U RE A ND GE NE SIS .
46 9
is p a rt of the m a teri a l f rom w hich a you ng one i s bu ilt u p

’
'
i t i s p a rt of th e ou tl a y for rep rod u ctiv e p u rp os es a nd di m i nu ,
t i on of it i s a l oss of rep rod u c tiv e p ower I n d ee d t he ca s e .
a ptly ill u st ra t es u nd er a no th er a s p ec t
,
the st r u ggl e b et ween
,
s e l f p re s erv a ti on a n d ra c e p re serv a t i on
- -
Ma i nt ena nce of th e
.
’
c ow s li fe d ep end s on m a i nt ena nce of i ts h ea t ; an d m a i n
ten a n ce of i ts h ea t m a y en t a il s u ch red u cti on i n th e s u pply
o f milk a s t o c a u se th e d ea th of t h e ca lf .
E vid enc e d eriv ed from t he h a bit s of th e s a m e or a lli ed

genera i n di fferent clim a t es m a y na t u r a lly be l ooked for ; bu t
,
'
it i s diffi cu lt t o get a nd it c a n s c a rcely be ex pe ct ed th a t the

,
r em a i ni ng c on diti on s o f exi s t en ce will be so fa r s i m il a r a s t o
a ll ow o f a fa i r c om p a ri s on b ei n g m a d e T he on ly ill u s t ra tiv e
.
fa c ts I h a v e m et w ith whi ch s ee m not ew orthy a re s om e na m ed ,
by Mr Gou ld i n hi s work on Th e B i r ds of A us tra li a H e

. .
sa y s I m u s t n ot om it t o m enti on t oo the ext ra ordi n a ry, ,
fe cu ndi ty w hi ch p rev a il s i n A u s t ra li a m a ny of i ts s m a l l er ,
bi r ds b reedi ng th r ee or f ou r tim es i n a s ea s on ; bu t l a yin g

f ewer eggs i n t he ea rly s p ri ng wh en i nsec t li fe i s l es s de
vel op ed a n d a gr ea t er nu m be r l a t er i n the sea s on w h en th e
, ,
s u pply o f i n s ec t f oo d h a s bec om e m or e a b u n d a n t I h a ve .
a l s o s o m e r ea son t o b el iev e th a t th e y ou n g o f m a ny s p eci es
b ree d d u ri ng the fi rs t s ea s on for a m ong oth er s I frequ entl y

, ,
fou n d one s ec ti on o f the H oney ea t er s ( th e M el i th rep ti ) s i t

-
ti ng u p on egg s w hil e s till cl oth ed i n th e b rown d res s of i m m a

tu ri ty ; a nd we k now th a t su ch i s the c a s e w ith th e i n t ro
du ced Ga l l i na c ece ( or p ou lt ry ) th ree or f ou r gener a ti on s o f
”
w hich h a v e b een oft en p rod u c ed i n th e c ou rse of a y ea r .
T h ou gh h er e Mr Gou ld ref ers only t o v a ri a ti on i n the

.
u a n tity o f f oo d a s a c a u s e o f v a r i a ti on i n th e r a t e o f m u l t i
q
pli ca ti on m ay we n ot s u s p ec t th a t wa rm th i s a p a rt ca u s e
,
-
o f th e high ra et w hi c h h e d e s c r ib es a s g en er a l ?
§ 3 49 . Of th e inv ers e v a ri a ti on b et w een a ctivity a nd

.
‘
g enes s ,i w e g et cl ear p roof L et u s b egi n with th a t which

.
Bi rd s fu rni s h .
4 70 L A WS OF MU L TI P L ICA TI ON .
Fi rst we h a ve the a v era ge c ont ra s t a l rea dy hin t ed b et ween

, ,
t he f ertility of B i rds a nd th e f ertility o f Ma mm a l s C omp a r .
ing the l a r ge with the l a rge a nd the s m a ll with the s m a ll we ,
s ee th a t c rea t u res w hi c h c on ti nu a lly g o th r ou gh th e m u s cu l a r
e x erti on o f s u s t a i n i n g th e m se lv es i n th e a i r a nd p rop elli n g
th em s elves ra pidly th rou gh it a re l ess p rolific th a n c rea tu res

,
o f eq u a l w eight s w hich g o th rou gh t he s m a ll er ex ert i on of
m ovi ng a b ou t ov er s olid s u r fa ces P red a t ory B i rd s h a v e

.
fewer you ng ones th a n p red a t ory Ma mm a l s of a pp roxima t ely

t he s a m e si zes I f we c om p a re Ro oks with Ra t s or Fi nch es
.
,
with Mice we fi nd lik e diff erences A nd these di fferences a re

,
.
g rea t er th an a t fi rs t a pp ea rs F or wherea s a m ong Ma m m a ls

.
a m oth er i s a bl e u n a id ed t o b ea r a n d s u ckl e a nd rea r h a lf

, ,
wa y t o m a t u rity a b r oo d th a t p rob a bly w eigh s m ore i n p ro

,
p orti on th a n d oes the b roo d of a Bi r d ; a Bi rd or a t le a st a ,
B i r d th a t fli es m u ch is u na bl e t o do thi s
, B oth p a rent s h a v e
.
t o h elp ; a nd this i ndi ca t es th a t t he m a r gi n for rep rod u cti on

i n ea ch a d u lt i n divid u a l i s s m a ll er .
A m ong B i rd s th em s elv es oc cu r cont ra s t s whi ch m a y be

n ext c on s id ered I n th e Ra pt ori a l cl a s s v a ri ou s s p eci es of
.
,
which di fferi ng i n th ei r s i zes a re S imil a rly a ctiv e i n thei r

, ,
h a bit s w e s ee th a t the s m a ll a re m ore p r olifi c th a n the l a rg e

, .
T he Gold en E a gl e h a s u su a lly 2 egg s : s om etim es 3 s om e ,
tim es only 1 A s w e d escend to the K it es a nd F a l con s the

.
,
nu mb er i s 2 or 3 a nd 3 or 4 , A nd w h en w e com e t o th e
.
S p a rr ow H a w k 3 t o 5 i s the S p ec ifi e d nu m ber
-
, S imil a rly .
a m ong the Owl s : w hil e th e Grea t E a gl e Ow l h a s 2 or 3 -
e ggs th e c omp a r a t iv ely s m a ll C om m on Owl h a s 4 or 5

, As .
be f ore hi n t ed it is imp os s ibl e t o s a y wh a t p rop orti ons of th es e

,
differenc es a re du e t o u nlik eness es of bu lk m erely a nd wh a t ,
p rop orti on s a re due t o u nlik enes s es i n the c o s t s of l ocom o

ti on B ut we m a y fa i rly a ssu m e th a t the u nlik enes ses i n the
.
c os ts of l oc om oti on a re h ere the m ore imp or t a nt f a ct ors .
Weight s v a ryin g a s the cu bes of the di m ensi ons whil e the ,
s u rfa c es of diges tiv e sy s t ems v a ry a s the squ a res the expens e ,
of flight i n c rea s es m ore ra pidly th a n d oes the a bility t o t a k e

E X P E N D IT U RE A ND GE NE S IS .
4 71
in nu rt im en t ; a nd a s m oti on th rou gh the a i r requ i res m ore

eff o rt t h a n m oti on o n t he g rou n d thi s g eom et r ic a l p ro gres s i on
,
te ll s m ore ra pidly on B i rd s th a n on Ma m m a l s Be thi s a s it .
m a y h owe v er th ese c ont ra s t s s u pp ort t he a r gu m en t ; a s do

, ,
v a ri ous others which m a y be s et d own T he Fi nch fa m ily for .

-
,
ex a mpl e h a v e b r ood s a v era gi ng a b ou t 5 i n nu mb er a nd h a v e

, ,
c o m m only 2 b r ood s i n th e s ea s on ; w hil e in t he C row f a mily -
the nu mb er of t he b rood i s on the a v era g e l es s a nd th ere i s ,
bu t one b rood i n th e sea s on A nd th en on d escendi ng t o s u c h .
s m a ll bi r d s a s th e Wren s a n d t he T it s w e h a v e 8 1 0 1 2 to , , ,
1 5 egg s a nd s o m etim es two b roo ds i n the y ea r

, One o f th e .
be s t illu s t ra t i ons i s f u rni s h ed by the S wa ll ow t ribe th r ough -

,
ou t w hi c h th ere i s littl e or n o di f ference i n m od e of li fe or i n

f ood T he S a nd Ma rti n mu ch the l ea s t o f th em h a s 4 t o 6
.
-
, ,
egg s a n d t wo b ro od s ; th e S wa ll ow s o m ewh a t l a rg er h a s 4 or , ,
5 ; a n d the S wi ft ( s imil a r i n h a bit s th ou gh u n rel a t ed ) l a rger ,
s till
,
h a s bu t 2 H ere we s ee a l ower f ertility a s s oci a t ed
.
i n p a r t wi t h g rea t er s i ze bu t a s s oci a t e d s till m ore con

,
s i cu ou s l y w ith g rea t er exp en dit u re For the differenc e of

p .
fertility i s m ore th a n p rop orti ona t e t o the differenc e of b u lk ,
a s s h own i n oth er c a s es ; a nd fo r thi s g rea t er diff erenc e th ere
i s th e rea s on th a t th e S wi ft h a s t o s u pp ort not o nly the c os t

,
o f p rop elli n g its l a rger m a s s th rou gh the a i r bu t a l s o th e ,
c os t of p rop elli ng it a t a high er v el ocity .
Omitti ng m u ch e vid ence o f lik e n a t u re l et u s n ot e th a t ,
di scl osed by comp a ri s ons of c er t a i n grou p s of bi rd s with

o th er g rou p s . S ku l k ers i s th e d escr ip t iv e titl e a ppli e d t o
the Wa t er Ra il t he C orn C ra k e a nd th ei r a lli es which ev a d e
-
,
-
, ,
e nemi e s by c on c ea l m ent— { ons equ ent l y e xp en di n g bu t littl e

‘
i n l ocom oti on T h es e bi rd s h a v e r el a tiv ely l a rg e b roo ds— 6

.
t o 1 1 8 to 1 2 85 0 N ot l es s i ns t ru c tiv e a re th e cont ra s t s be
, , .
t ween the Ga lli na ceou s Bi r d s a nd oth er Bi rd s of like s i zes bu t

m or e a c tiv e h a bit s The P a rt ridge a nd the Woo d P igeon a re
.
-
a b ou t eq u a l i n b u lk a nd h a v e m u c h th e s a m e f ood Y et whil e .
the on e h a s f ro m 1 2 t o 2 0 y ou ng ones the oth er h a s bu t 2 ,
y oun g ones t wi ce a yea r : i ts a nnu a l rep rod u cti on i s l es s th a n

-
4 72 L A WS or MU L T I P L ICA T I ON .
o n e— thi rd I t m a y be s a id th a t the a bility of the P a rt ridge

.
t o b ri ng u p s o l a rge a b ro o d i s du e t o t h a t h a bit of i t s t r ib e
,
”
whi c h on e o f i ts na m es S cr a p ers d es crib es ; a nd t o t he
, ,
a cc om p a nyi n g h a bi t o f th e y ou n g w hi c h b egi n t o t th i
g e , e r
ow n livi n g a s s oon a s th ey a r e h a t c h ed : s o s a vi n g th e p a r ent s

’
l a b ou r . C on v ers ely it m a y be s a id th a t th e in a bili t y of

,
P ig eons t o rea r m or e t h a n 2 a t a ti m e i s ca u se d by the ne ce s ,
s i ty of fetc hi ng e v ery t hi n g t h e Bu t t he a ll eged r el a t i on

y ea t .
h old s neverthe l es s On the one h a n d a g rea t p a rt of th e f oo d

.
,
wh i c h the P a r t r idge chi c k s pi c k u p i s food whi ch i n th ei r , ,
a b s en ce the m o th er w ou ld h a v e pi c k e d u T h u gh h hi k
, p o e a c c c .
c os t s h er fa r l es s th a n a y ou n g P ig eon cos t s i ts p a re nt s
yet ,
t h e wh o l e o f h er c hi c k s cos t h er a g rea t d e a l i n t h e s h a p e of
a b s t i n en ce — a n a b s ti nen ce s h e c a n b ea r be ca u se s h e h a s t o fly
bu t littl e On th e ot h er h a n d the P igeon s h a bit of l a yi ng ’

.
,
a nd h a t c hi ng bu t tw o e gg s m u s t n ot be re ferre d t o a n f
, y or e
s een neces s ity o f g o i n g th ro u gh s o m u ch l a b ou r i n s u pp or ti ng
t h e y ou n g bu t t o a con s tit u ti on a l t en d en cy es t a bli s h ed by

,
su c h l a b ou r T hi s i s p rov ed by th e cu ri ou s f a c t th a t wh en
.
d om es ti ca t ed a n d s a v ed f rom su ch l a b ou r by a rtifi c i a l feeding

, ,
P igeons s a y s Ma c gilliv ra y ( qu oti ng A itki n )

,
a re freq u ently ,
s een S itti n g on e gg s l on g b efore th e f or m er b rood i s a bl e t o
l ea v e the nes t s o th a t the p a rent bi r d h a s a t the s a m e tim e

,
”
y ou ng bi rds and eggs t o t a k e c a re of .
350 . Ma d e to illu s t ra t e t he effect of a ctivity on fertility ,
m os t c omp a ri s ons a m ong M a m m a l s a r e o bj ec t i on a bl e : o th er
ci rcu mst a nc es a re n ot e qu a l . A few h ow ev er es ca p e thi s

, ,
c riticism .
One i s th a t bet ween the H a re a nd the Ra bbit T h es e a re .
c l os ely a lli ed s p eci es o f th e s a m e g enu s s imil a r i n th ei r di et

-
,
bu t u nlik e i n th ei r exp endit u res for l oc om oti on The rel a .
t i vel y i ner t Ra bbit h a s 6 y oung ones i n a litt er a n d four

-
,
litt ers a yea r ; whil e the rel a tiv ely a ctiv e H a re h a s bu t 2 t o

- -
5 i n a litt er T hi s is not a l l
. T he Ra bbit b egi ns t o b reed
.
a t S i x m onths ol d ; bu t a y ea r el a p s es b ef ore t he H a re be gi n s
E X P E ND IT U RE A ND GE NE SIS .
4 73
t o bree d
‘
. T h es e two f a ct ors c omp ou nd ed res u lt i n a dif ,
f erenc e of fertility fa r grea t er th a n ca n be a s crib ed to u n

lik enes s of the two c rea tu res i n S i ze .
P erh a ps the m o st s t r iki ng pi ec e of evid enc e which Ma m

m a ls fu rn i s h i s the ext rem e i nfertility of ou r c om m on B a t
,
.
T he C hei r op tera a n d th e Roden ti a a re n ot v ery di s s imil a r i n

thei r i nt ern a l s t ru ct u res D ivers ity of c ons tit u ti on th erefore
.
, ,
c a nnot viti a t e th e c omp a r i s on b et ween B a t s a nd Mi ce which ,
a re a b ou t th e s a m e i n s i z e T h ou gh th ei r di et s differ the
.
,
differenc e i s i n fa v ou r of th e Ba t : i t s foo d bei n g exclu s iv ely

a n im a l w hil e th a t of t he Mou s e is m a i n ly v eg e t a l Wh a t .
now a re th ei r res p ec tiv e r a t es of g enes i s ? T h e Mou s e h a s

s ev era l litt er s i n a y ea r of 5 t o 7 i n ea c h whil e th e B a t p ro
d u ces only one a t a tim e Wh eth er th e B a t rep ea t s i ts one
.
m ore f req u ently th a n the Mou s e rep ea ts i ts 7 i s not s t a t ed ;

bu t it i s qu it e c ert a i n th a t ev en if it d oes s o ( a n a b s u r d
s u pp os iti on ) ,th e m ore f requ en t rep etiti on c a nnot be su c h
a s t o r a is e i t s f er tility t o a nythi ng lik e th a t o f t h e Mou s e .
A nd this rel a tiv ely l ow ra t e of m u ltipli ca ti on we m a y f a i r ly

-
a s c rib e t o i t s r el a tiv ely high ra t e o f exp en dit u re

-
.
H ere l et u s n ot e i n p a s s i ng a n i nt e res ti n g ex a m pl e o f the

, ,
wa y i n whi c h a s p ec i es whi ch h a s n o sp eci a lly g rea t p ow er o f -
sel f p res erv a ti on

-
whi l e i t s p ower of m u ltiplic a ti on i s ex
, ‘
t rem el y s m a ll nev erth el es s a v oid s exti ncti on b ec a u s e it h a s

,
t o m eet a n u nu su a lly s m a ll t ot a l of ra c e d es t royi ng f orces

- -
.
L ea vi ng ou t p a ra s it es th e only enem y of th e B a t is the Owl ;

,
a nd th e Owl i s s p a ri n gly di s t r ib u t ed .
§ 3 5 1 T hes e genera l evid enc es m a y be enf orc ed by s om e

.
sp eci a l evid enc es We h a v e few opp ort uniti es of ob servi ng

.
h ow w ithi n th e s a m e S p eci es v a ri a ti ons of exp en dit u r e a re

, ,
r e l a t ed t o v a r i a ti ons of f ertility B u t a f a ct or two sh owi ng .
the c onn exi on m a y be n a m ed .
D oct or D unc a n qu ot es a st a t em ent t o the p oint res p ecti n g

the b reeding of d og s A l rea dy in 3 4 1 I h a v e ext ra ct ed a p a rt
.
o f thi s s t a t em ent , t o the effect th a t b ef ore her g rowth is com

4 74 L A WS O F MU L TI P L ICA TION .
pi ete a bit ch b ea rs a t a bi rth fewer p u ppi es th a n wh en s he

,
b ec om es fu ll grown A n a c com p a nyi ng a ll eg a t i on i s th a t

-
.
,
her d ec lini ng vig ou r i s s h own by a d ecrea se i n th e nu m ber o f

”
p u ppi es cont a i ned i n a lit ter en di n g i n one or two
, A nd.
th en it i s f u r th er a ll ege d t h a t a s r e g a r d s t h e a m ou n t o f
,
w ork a dog h a s t o p erform s o will th e d ecli ne be ra pid or

,
gra d u a l ; a nd h ence if a bit ch i s work ed h a rd y ea r a fter y ea r

, ,
s h e w ill f a il r a pidly a nd t h e dimin u ti on of h er p u ppi es w ill

,
be a cc or di ngly ; bu t i f w ork ed m o d er a t ely a n d well k ept sh e ,

”
will fa il gra du a lly a nd the dimi nu ti on wi ll be l es s ra pid
,
.
I n thi s pl a ce m ore fi tl y th a n e l sewh ere m a y be a dd ed a

, ,
f a ct of lik e impli ca ti on th ou gh of a di fferen t ord er

, Of .
c ou rs e wh eth er exc ess ive exp endit u re be i n the conti nu a l re

p a i rs of nerv o mu scu l a r ti ss u es or i n repl a ci n g oth er ti s su es
-
,
t he rea c tiv e effec t s if n ot q u it e th e s a m e wi ll be s i m il a r

, ,
th ere will be a d ec rea se of the s u rplu s a v a il a bl e for genes i s .
I f t h en i n a ny a n i m a ls th ere fro m ti m e t o tim e oc cu r u nu su a l

, ,
ou tl a y s for s elf m a i nt en a n ce w e m a y exp ect th e p eri od s o f

-
,
s u ch ou tl a y s t o be p eri ods of dimi ni s h ed or a rres t ed rep ro

d u cti on T h a t th ey a re s o th e m ou lti ng of bi rd s sh ow s u s
. .
Wh en h ens beg in t o m ou lt th ey c ea se t o l a y Whil e th ey .
a r e e xp en di n g s o m u c h i n p r o d u ci n g new cl o thi ng th ey h a v e
,
no thi ng t o expe n d for p rod u c i ng eggs .

C H A P TE R I X .
C OI N C I D E N CE BE T W E E N H GHI N UT RI T IO N A N D GE N E S S I .
§ 3 5 2 U N D E R thi s h ea d m a y be grou p ed v a ri ou s f a cts

.
whi ch i n a noth er wa y t e ll t he s a m e t a l e a s th os e cont a i ne d

, ,
i n the l a s t c h a pt er The evid en c e th ere pu t t o get h er wen t

.
t o S h o w th a t i nc rea se d c os t of self m a i nt ena nce ent a il ed de

- ~
c rea s e d p ow er o f p r op a g a ti on T he evid enc e t o be s et d own

.
h ere will g o to s h ow th a t p ower o f p rop a ga ti on i s a u gm ent e d

,
by m a ki ng self m a i n t ena nce u nu s u a lly ea s y F or i nt o this

-
.
m a y be t ra ns l a t ed th e effect of a b u n d a n t f ood .
T o pu t th e p rop os iti on m ore S peci fica l l y— we h a v e s een

th a t a ft er i ndivid u a l gr owth d ev el op m ent a nd d a ily con
, ,
s u mpti on h a ve b een p rovid ed for the s u rplu s nu t ri m ent

, ,
m ea s u res the r a t e o f mu ltipli ca ti on T his su r pl u s m a y be

.
ra i s e d i n a m ou n t by s u ch ch a ng es i n th e en vi r on m en t a s
b ring a l a r ger s u pply of the m a t eri a ls or f orc es on w hi c h

b oth p a rent a l li fe a nd the liv es o f off s p ri n g d ep end Be .
th ere or be there not a ny exp endit u re a high er nu t riti on

, , ,
w ill m a k e p o s s ib l e a g rea t er p rop a g a ti on We m a y exp ect .
thi s t o h old b o th of a ga m og enes i s a nd of g a m ogenesis a nd

we S h a ll fi nd th a t it d oes s o .
3 53 m u lti a xi a l pl a nt s the p rim a ry eff ect o f s u rpl u s

. On -
,
nu t ri m en t is a p ro d u cti on of l a rge a nd n u merou s l ea f s h o ot s -

.
H ow this a s exu a l m u ltipli ca ti on res u lt s f rom exc essiv e nu t ri

ti on i s well s h own wh en the l ea di ng a xi s or a chi ef b ra nch
, , ,
i s b r ok en off t ow a r d s i t s e xt remity Th e a xil l a ry b u ds b el ow

.
4 75
4 76 L A WS OF MUL T IP L I CA TI ON .
t he b rea k a g e q u i ck l y s well a nd bu rs t i n t o l a t era l sh oots ,
whi c h often pu t for th s econd a r y s h oot s : two g enera ti ons of

a g a mic i n divid u a l s a r i s e w h ere th ere p r ob a bly w ou ld h a v e
b een non e bu t for the l oc a l a bu nd a n ce of s a p no l ong er ,
d ra wn off I n lik e m a nner the a bnorm a l a ga m ogenes i s whi c h

.
we h a v e i n p r olif erou s fl ow ers is h a bit u a lly a cc omp a ni e d by ,
a g en era l l u x u ri a n ce implyi ng a n u nu s u a l pl eth or a

, .
N 0 l es s conc lu s iv e i s the evid ence fu r ni s h e d by a ga m o

.
g enes i s i n a ni m a l s S i r J ohn D a l yel l S p ea king of H ydra

.
,
tu ba a n d of the p eri o d b efore s t r obili za ti on comm ences s a y s

, ,
I t i s s i ngu l a r h ow m u c h p rop a ga ti on is p rom ot ed by

”
a b u n d a nt s u s t en a nce T hi s P o lyp e g oes on b u ddi ng ou t
.
-
y ou n g p oly pe s from i t s s id es with a ra pidity p rop or ti ona t e ,
t o th e s u pply of m a t er i a l s S o t oo i s i t w ith th e
.
, ,
ag a m ic re p r od u c ti on of th e A p hi s A s cit ed by P rof es s or .
H u xl ey Kyber s t a t es th a t h e ra is ed vivip a rou s b r ood s of

“
,
b oth thi s s p eci es ( A p hi s D i a n thi ) a nd A R os ce for fou r con .
s ecu t i ve y e a r s w ith ou t a ny i nt erv en ti on o f m a l es or ovi

,
p a rou s fem a l es a nd th a t the energy of the p ower of a ga m ic

,
rep rod u cti on wa s a t t h e en d o f th a t p eri o d u ndimi ni s h e d .
T he ra pidity of th e a g a m ic p r olifer a ti on th rou gh ou t t he

wh ol e p eri od w a s di rect ly p rop orti ona l t o th e a m ou nt o f
w a rm th a nd f oo d s u ppli ed .
I n th ese c a ses th e rel a ti on is n ot a pp reci a bly c ompli ca t ed

by exp en dit u re T he pa rent h a vi ng rea c h ed its li m i t of
.
g row th th e a b s orb ed f oo d g oes t o a s exu a l m u ltipli ca ti on

,
s c a rc ely a ny b ei n g d ed u ct ed for th e m a i nt ena n c e o f p a renta l
life .
§ 354 . The
s exu a l m u ltiplic a ti on of org a nism s u nd er
c h a ng ed c onditi ons u nd e rg oes v a ri a ti ons c on f o rmi n g t o a
,
p a ra ll el l aw C ultiv a t ed pl a nts a nd d om es tic a t ed a nim a ls

.
yi eld u s p r oof of thi s .
F a ct s s h owi ng t h a t in c ultiv a t ed pl a nts s exu a l genes i s i n

c rea s es with nu t riti on a re obscured by f a ct s sh owing th a t a
,
l es s r a pid a sexu a l g en esis , a nd a n incipi ent s ex u a l g enes is ,

N UT RIT ION A ND GE NE S IS .
4 77
ac oc mp a ny the f a ll from a high to a m od era t e nu t riti on The .
c on f ou n di n g o f th e se t wo re l a t i ons h a s l ed t o mi s t a k en i n fer
e nces . Wh en t rea ti ng o f Gen esi s i nd u c tively we r ea ch ed th e ,
g en era liz a ti on th a t the p rodu cts of a f ertili zed g er m g o

“
on a ccu m u l a ti n g by s i m pl e g rowth s o l o n g a s t h e f or c e s
,
wh ence g r owth resu lt s a re grea tly i n exc ess of th e a nt a g oni s t

f orces ; bu t th a t wh en dimi nu ti on of the one s et of f orces or ,
i ncrea s e o f th e oth er ca u s es a con s id er a bl e d ecli n e i n thi s ex

,
c es s a nd a n a pp r oa ch t owa r d s equ ilib ri u m f ertili zed germ s

, ,
a re a g a i n p r o d u c e d .
’
I t wa s p oi nt ed ou t th a t thi s
h old s of or ga ni s m s whi ch m u ltiply by h et erog enes is a s well ,
a s th ose w hi c h m u ltiply by h o m og enes i s A nd pl a nt s w ere

.
r efer re d t o a s ill u s t ra ti ng b oth genera lly a n d l oca lly th e

, ,
d ecli ne of a ga m ic m u ltiplic a ti on a n d c om m encem ent of g a mic

m u ltiplic a ti on a l on g with a l esseni ng ra te of nu t riti on N ow
,
.
th e m a n y c a s es w hi ch a re giv en of f ru itfu l ness ca u s ed i n t rees

by depl eti on a re rea lly c a s es of thi s ch a nge f ro m a ga m o
,
genes i s t o g a m ogenes is ; a nd s imply g o t o p rov e th a t wh a t

w ou ld n a t u ra lly a ris e wh en d ec rea s e d p eriph era l g rowt h h a d
f oll owed i nc rea s ed s i ze m a y be b rou ght a b ou t a rtifici a lly by
,
dimi ni s hi ng th e su pply of m a t eri a l s for g rowth Cra m pi ng .
its r oots i n a p ot or cu tti ng th em or r i ngi ng i ts b ra nch es

, , ,
will m a k e a t ree b ea r v ery ea rly : b ri n gi n g a b ou t a p rem a

t u re es t a bli s h m en t of th a t rel a tiv e i nnu t riti on whi ch w ou ld
h a v e s p on t a neou s ly a ri sen i n c ou rs e of tim e S u c h f a cts .
by n o m ea ns S h ow th a t i n pl a nt s s exu a l genes is i ncrea ses a s

nu t r iti on di m i ni s h es When it h a s once s et i n s exu a l
.
,
g enes i s i s s ca n ty or imp erfect u nl es s nu t riti on i s g ood .
T h ou gh th e s ta rv e d pl a nt m a y bl os som yet m a ny o f i t s ,
bl os s om s will fa il ; a nd su ch s eed s a s it p r odu ces will be ill

fu rni s h ed with th os e env el opi n g s t ru ctu res a nd th a t st ore of
a lb u m en &c
,
. n eed ed t o giv e g o o d c h a n ces o f s u cce s s f u l g er
,
mi n a ti on— the nu mber of su r vivi n g offs p ri ng w ill be dimin

i s h ed Were it oth erwi s e t he m a nu ri n g of fi eld s w hi c h a r e t o
.
,
b e a r s eed c rops wou ld be not S imply u s el es s bu t i n j u ri ou s

-
, .
Were it oth erwi se d ungi ng the roots of a fru it t ree w ou ld i n

,
-
al l c a s es be imp olitic ; i nst ea d of be ing imp olitic only wh ere

the g rowth of s exl es s a xes is s till l u x u ri a nt Were it other .
wi s e a t ree which h a s b orne a h ea vy c rop s h ou ld by the

, ,
c on s eq u ent d epl eti on be l ed t o b ea r a s till hea vi er c rop next

,
y ea r ; wh erea s it i s a pt t o be wh olly or p a rti a lly b a rren next

y ea r— h a s t o rec ov er a s t a t e of t ol era bly high nu t riti on b efore-
i ts se x u a l genesis a g a i n b ec om es l a rge .
B u t th e b es t ill u s t ra ti ons a re yi eld ed by a nim a l s— th os e

a n i m a l s a t l ea s t i n w hi c h we h a v e b es id es a n i n c rea s ed su
, p
ply o f nu t rim ent a dimi ni s hed exp endit u re T wo cl a ss es of
, .
c om p a ris ons a lik e i n thei r impli ca ti ons m a y be m a d e

, ,
c om p a ri s on s b et ween t a m e a nd w ild a ni m a l s of the s a m e

s p eci es or g enu s a n d c omp a ri son s b et ween t a m e a n im a ls of
,
the s a m e s p eci es differently t rea t ed .
T o b egin w ith B i r d s l et u s fi rs t c ont ra s t the f a rm y a rd

,
-
Ga l l i na cece w ith th ei r ki nd red of th e fi elds a nd w oods N ot .
with s t a ndin g thei r g rea t er s i ze w hich oth er thi ng s equ a l

, , ,
sh ou ld be a c comp a n i ed by s m a ller f ertility the d om es ti ca t ed ,
ki n d s h a ve m ore nu m erou s off s p ri ng th a n the wild ki nd s A .
T u rk ey h a s a d ozen i n a b r ood w hil e a P h ea s a nt h a s f rom 6

,
t o 1 0 T wic e or th r ic e i n a s ea s on a H en r ea rs a s m a ny
.
,
c hi ck en s a s a P a rt ridg e rea r s onc e i n a s ea son A n s eri ne .
bi rd s S h ow u s p a ra ll el differences T he T a m e Goos e sit s on

.
1 3 t o 1 8 egg s a n d oft en sits a s ec ond ti m e ; bu t the Wild

Goose s its on 5 6 or 7 a nd th es e a re n ot e d a s c ons id era bly
, , ,
sm a ll er I t is the s a m e with D u ck s
. T h e d om estic a t ed
.
v a ri ety l a y s a nd h a tch es t wic e a s m a ny egg s a s th e wild

v a r iety A nd the lik e h old s of P igeons A ft er rem a rki ng
. .
o f the C ol u m ba l i vi a th a t i n S p ri n g wh en th ey h a v e pl enty
o f c orn t o pick f r o m t h e newly sown fi elds th ey begi n t o
-
,
t f t a n d p a i r ; a n d a g a i n i n h a rv est wh en th e corn i s c u t
g e a ,
d own Ma cgilliv ra y g oes on t o s a y th a t the s a me p a i r wh en

”
,
t a m ed g enera lly b reed f ou r tim es i n the y ea r T ha t .
b et ween di ff eren t p ou lt ry y a rd s i neq u a liti es of f ertility a re

-
c a u s ed by i nequ a liti es i n the s u ppli es of f ood is a f a mili a r ,
t ru th H igh feedi ng s h ows i ts effect s not only i n the c on

.
NU TRITION A ND GE NE S IS .
4 79
ti nu ou sl a yi ng bu t a ls o i n the s i zes of the eggs

,
A m on g .
di recti ons giv en for obt a i n i ng egg s fro m p u ll et s l a t e i n t he

y ea r it i s es p eci a lly i n s i s t ed on th a t th ey s h a ll h a v e a
,
g enerou s di et Res p ecti ng P igeons Ma cgilliv ra y wri tes

.
th a t th ei r b reedi ng d ep en d s m u ch on th ei r h a vi ng pl enty of
“
f ood t o fa tt en t h em seem s I thi nk evid en t from the c i rcu m

, , ,
s t a nc e t h a t when t a m ed whi ch th ey ea s ily a re t hey a re

, , ,
o b s erv e d t o b r eed i n ev ery m on th of t he y e a r I d o not m ea n .
th a t the s a m e p a i r will b reed ev ery m onth bu t s ome i n the

fl oc k i f we ll fed will b reed a t a ny sea s on
, ,
T here m a y .
be a dd ed a f a ct of lik e m ea n i ng whi ch p a rti a lly d om es tic a t e d -
bi rd s yi eld T he S p a rrow i s one of the Fi nch t ribe th a t ha s

.
t a ken t o the neighb ou rh ood of h ou s es ; a n d by its b oldn es s

s ecu res food not a v a il a bl e to its c ongeners T he re s u lt i s .
th a t it h a s sev era l b rood s i n a s ea son whil e its fi el d h a un ti ng ,

-
ki nd red h a ve n one of t hem m ore th a n two b roo ds a nd s om e ,
h a v e on ly one .
E qu a lly c l ea r p roof th a t a b u n d a n t nu t rim en t ra i ses the

r a t e of m u ltipli ca t i o n occu r s a m ong Ma m m a ls
, C omp a re the .
litt ers o f the D og with th e litt ers of the Wolf a nd the F ox .
Wh erea s th os e of the one r a nge i n nu mb er f rom 6 t o 1 4 th os e ,
o f the oth er s c ont a i n re s p ectiv ely 5 or 6 or occ a s i on a lly 7 a nd

,
4 or 5 or r a rely 6 A ga in, the Wild C a t h a s 4 or 5 kitt ens ;

.
bu t th e t a m e C a t h a s 5 or 6 kitt ens 2 or 3 ti m es a y ea r -
.
S o too i s it w ith th e Wea s el t rib e Th e S toa t h a s 5 y ou n g

'
, , .
ones on c e a y ea r -
T h e F erret h a s 2 litt ers yea r ly ea ch con
.
,
t a i ni ng fr om 6 t o 9 ; a nd thi s not w ith s t a ndi ng th a t it i s the

l a rger of the two P erh a ps the m os t s t riki n g c ont ra st is th a t
.
b et ween the wild a nd t a me v a ri eti es of the P i g Whil e the .
o ne p r od u c es a cc or di ng t o i t s a e fro m 4 t o 8 or 1 0 y ou n g
, g ,
o nes onc e a y ea r th e o th er p r o d u ces s om etim es a s m a ny a s 1 7

,
i n a litt er ; or i n oth er c a s es will b ri ng u p 5 litt ers of 1 0 ea c h

, ,
i n two y ea rs— a ra t e of rep rod u ct i on whi ch is u np a ra ll el e d

'
i n a ni m a l s of a s l a rge a si ze * A n d l et u s n ot om it t o n ote
.
tha t thi s exc ess iv e fertility occu rs wher e th ere i s the

The cli m a te, the l ocali ty, and the ki nd o f food, a re of cours e a l l fa ct ors ;
L A WS OF MU L T I P L ICA T ION .
g rea t est i na ctivity— wh ere th ere i s pl enty to ea t a nd nothi ng

t o do . T h ere i s n o l es s di s ti nc t e vid en ce t h a t a m ong
d om es ti ca t ed Ma m m a l s th em s elv es th e well fed in dividu a l s ,
-
a re m ore p ro lifi c t h a n th e - ill fed i n dividu a l s On the high -

.
a n d c om p a r a tiv ely i n fertil e C ot s wold s it i s u nu s u a l fo r

-
,
e w e s t o h a v e t w i n s ; bu t th ey v er y com m on ly h a v e t w i ns
i n the a dj a c en t r i ch v a ll ey of th e S ev ern S imil a rly a m ong .

,
t he b a rren hill s of th e w es t of S cotl a n d t wo l a m b s will be ,
b orne by a b ou t one ewe i n t wenty ; wh erea s i n E ngl a n d ,
s om ethi n g like one ewe i n th ree w ill bea r two l a mb s N a y .

,
i n r ich p a st u res twi ns a re m ore f requ en t th a n si ngl e bi rth s ;

,
a nd it occa s i ona lly h a pp en s th a t a ft er a g en i a l a u t u m n a n d ,
c on s eq u en t g oo d g ra z i ng a fl o ck o f ew e s will n ext sp ri ng
,
yi eld d ou bl e th ei r nu mb er of l a mb s— the t ripl et s b a l a nci ng

t he u n ip a rae S o di r ec t i s thi s rel a ti on th a t I h a v e h ea rd a
.
,
f a rm er a s sert hi s a bili ty t o f oret ell f rom th e high m edi u m , , ,
o r l ow c on diti on of a n ew e i n th e a u t u m n wh eth er S h e will

, ,
next sp ri n g b ea r t wo or on e or n one , , .
355 bj ecti on m u s t h ere be m et Many f a cts m a y

. An o .
be b rou ght t o p r o v e th a t f a t nes s i s n ot a cc om p a ni ed by f erti

l i ty bu t by b a rrennes s a nd th e i nferenc e d ra w n i s th a t high
f eedi n g i s u n fa v ou ra bl e t o genes i s T he p remi s s m a y be .
a d m itt ed w hil e th e c on cl u s i on is d en i ed .
T h er e i s a di s ti nc ti on b et ween wh a t m a y be c a ll ed norm a l
p l eth ora a nd a n a bnorm a l pl eth ora li a bl e to be con fou nd ed
, ,
w ith it T he one i s a m a rk o f con s tit u ti ona l w ea lth ; bu t th e

.
o th er i s a m a r k o f con st it u ti on a l p ov erty N orm a l pl eth ora .
i s a su p erfl u i ty o f m a t er i a l s b oth for th e b u ildi ng u p of
a nd h ence, p rob abl y, t he difference s b et wee n th e s t ate m ent s of different eu
t hori t i es co nc e rni ng th ese s ev e ra l ca ses P rof MacBri de wri t es . .
A c cordi ng to Fl ow er ( Ma m m a l s L i vi ng a nd E xti nct) th e Ferre t i s

“
,
a
dom es t i ca te d v a ri ety of t he com m on pol eca t, w hi ch has 3 to 8 young . Da r

wi n (A n i m a l s a nd P l a nts ) sa ys th a t th e wil d s ow o fte n b reeds t wi ce yea r
a
a nd produce s a l i tt e r of 4 to 8 , a nd som e t i m e s e v en 12 . Th e dom es t i c sow
b re e ds t wi ce a nd woul d breed oftene r if perm i tted, and if any good a t a ll p ro

”
duces 8 in l i tte r .
N UTRITI ON A ND GE NE SIS .
481
ti s s ue a nd the ev olu ti on of f orc e ; a nd thi s is the pl eth ora

which we h a ve f oun d to be a s s oci a t ed with u nu s u a l f ecun di ty .
A b norm a l pl eth ora w hich a s t ru ly a ll eged is a cc o mp a ni ed

, ,
by i nfecu ndity i s a s u perfl u i ty of f orce ev olvi ng m a t eri a l s

,
-
j oi ned with eith er a p os itive or a rel a tiv e d efi c i ency of ti ssu e

-
f orm i ng m a t eri a l s : the i nc rea s ed b u lk i ndic a ti ng thi s st a t e ,
be i ng rea lly th e b u lk o f s o mu c h i nert or d ea d m a t t er N ot e .

,
fi rs t a few o f the f a cts which S h ow u s th a t obes ity i m pl i e

,
phy s i ol ogic a l i m p overi s hm ent .
N eith er i n b ru t es n or m en d oes it or di na r ily occ u r eith er

in y ou th or i n th a t ea r ly m a t u rity d u ri ng which the vig ou r
i s th e g rea t es t a nd t he dig es ti on the b es t : it d oes not
h a bitu a lly a cc omp a ny the high es t p ower of t a ki n g u p nu t ri
tive m a t eri a ls Wh en f a t ness a ri s es i n th e p ri m e of li fe
.
,
w h eth er f rom p ecu li a r ity of f ood or o th er ci rc u m s t a nc e it is ,
n ot t h e S ign of a n i nc rea s e d t ot a l vit a lity On th e c on t ra ry

.
,
if grea t mu s cu l a r a cti on h a s to be g one th rou gh the fa t ,
mu s t be g ot ri d of either a s i n a m a n by t ra i ni ng or a s i n
, , ,
a h or s e th a t h a s gr own b u lky w hil e ou t a t g ra ss by p u tti ng ,
him on s u ch m ore nu t ritiv e di et a s c a t s . T h e f req u ency

o f s en il e f a t nes s b oth i n d o m e s ti ca t ed c r ea t u res a n d i n ou r
,
s elv es h a s a s imil a r i m plic a ti on

,
Wh ether we c ons id er the
.
s m a ll er a bility of th os e who di s pl a y it t o with s t a n d l a rge

d em a nd s on thei r p ower s or wh et h er we c ons id er the com
,
a ra ti v el y i nfer i or dig es ti on c om m on a m on g th em we s ee
p
-
,
th a t the i nc rea s ed S i ze i ndi ca t es not a n a bu nd a nce of m a t e

,
ri a l s whi c h th e org a n i s m requ i res bu t a n a bu nd a nc e o f

,
m a t eri a ls which it d oes n ot req ui re . Of lik e m ea ni n g

i s the f a c t th a t w om en wh o h a v e h a d s ev era l child ren a nd ,
a n i m a ls a ft er th ey h a v e g on e on hea r i ng y ou n g for s om e
tim e f requ ently b ec om e fat ; a nd l os e th ei r fecu ndity a s

,
they do this I n s u ch c a s es the f a t nes s i s not to be t a k en

.
a s th e c a u s e of the i n f ec u n dity ; bu t the c on s titu ti ona l

“
exh a u s ti on w hich t he p revi ou s p r o d u cti on of offsp r i n g h a s
l eft sh ow s it s elf a t once i n the f a ili ng fecu ndity a nd the

,
c om menci ng f a t ness . T h ere is yet a noth er ki nd of evi

77
L A WS OF MUL TI P L ICA TION .
den ce . Obes ity n ot u n o c mm only s et s i n a fter th e syst em h a s

b een s u bj ec t t o d ebili t a t ing i n flu enc es Oft en a s eri ous ill nes s
.
i s foll owed by a c orp u l ence t o whi c h th ere wa s p revi ou sly

n o t en d enc y . An d th e p r ol onged a dmi ni s t r a ti on of m erc u ry ,
cons tit u ti ona lly i nj u ri ou s a s it i s s o m et im es p ro d u ces a lik e

,
eff ect .
C l os er i nqu i ry v erifi es th e c onclu s i on t o whi ch th es e f a cts

p oi nt T he mic r os c ope S h ow s th a t a l ong wi th th e i ncrea s e of
.
bu lk c omm on i n a dv a nced lif e th ere g oes on wh a t i s c a ll ed

,
f a tty d egenera ti on oi l gl ob u l es a r e d ep osit ed w h ere th ere

-
s h ou ld be p a rti c l e s o f fl es h— or ra t h er w e m a y s a y t he
, ,
hvdroca rbon a c eou s m ol ecu l es l oca lly p ro d u c ed by dec om

p os iti on of the n it rogen ou s m ol ecu l es h a v e not b een repl a ced
,
by other ni trogenou s m ol ec u l es a s th ey sh ou ld h a ve been

, .
T his f a tty d egen era ti on i s i n d ee d a ki n d of l oc a l d ea th

, ,
.
For so reg a r di ng it we h a v e n ot si m ply th e rea s on th a t a n

a c tiv e s u b s t a nc e h a s i ts pl a c e occu pi e d by a n inert s u b s t a n ce ;
bu t we h a v e th e fu rth er rea s on t h a t t he fl es h of d ea d bodi es ,
u n d er ce rt a i n c ondi ti ons i s t ra n sf or m ed i nt o a f a tty m a tt er

,
c a ll ed a dip oc ere .
Th e in fertility th a t a cc omp a n i es f a t nes s i n d om e stic

a nim a ls h a s h ow e v er o th er c a u ses th a n th a t d ecli ni n g c on
, ,
s ti tu t i on a l vig ou r which the f a t ness c om m only i ndic a t es .
B ei ng a rtifi ci a lly fed th es e a nim a l s ca nnot a lw a y s obt a i n

,
wh a t th ei r s y s t em s n eed . T h a t w hich is giv en t o th em i s

giv en exp res s ly b ec a u s e of its f a tt ening qu a lity A nd s i nc e .
the c a p a c ity of th e diges tiv e a pp a r a t u s rem a i n s t he s a m e ,
the a b s orpti on of fa t p rod u c i ng m a t eri a ls i n exce s s impli e s

-
,
d efect i n the a b s orpti on of m a t eri a l s f rom which the

ti ss u es a re f orm ed a nd ou t of which y ou ng ones a re bu ilt
,
up . Moreov er this S p eci a l fee di ng with a vi ew

,
t o ra pid a nd ea rly f a tt eni n g c on ti nued a s it is th rou gh

,
genera t i on s a nd a c c omp a ni ed a s it is by a s el ec ti on of i n di
,
vi du a l s a nd v a ri eti e s which f a tt en m os t r ea dily t end s to,
es t a bli s h a m odifi e d c on s tit u ti on m ore fitt ed for p rod u ci n g

‘
fa t a nd c orres p on di ngly l ess fitt ed for p ro d u cing fl esh— a

-
N UTRITION A ND GE NE S IS .
483
c onstitu ti on which f rom thi s rel a tiv ely defi ci ent a bs orp ti on
,
-
o f nit rog en ou s m a tt ers is lik ely t o b ec o m e i n f ertil e ; a s i n

, ,
d eed th es e v a ri eti es oft en d o b ecom e

,
H enc e n o con .
,
e lu s i on s res p ecti ng the eff ect s of high nu t r iti on p rop er ly s o ,
c a ll ed ca n be d ra wn f rom c a s es of thi s ki nd T he c a s es a re
,
. .
i n t ru th of a ki n d whi ch c ou ld not exi st bu t for hu m a n

,
a g enc y Und er na t u ra l c on diti on s no a nim a l w ou ld di et

.
it self i n the wa y requ i red t o p rod u c e Su ch res u lts A nd if it .
*
did i ts ra c e w ou ld qu ickly di s a pp ea r .
T h ere is yet a n oth er m od e i n whi ch a ccu mu l a ti on of fa t

dimi ni s h es f ertili ty E v en s u pp os i ng it u na cc omp a ni ed by
.
a s m a ll er a b s orpti on of nit rog enou s m a t eri a ls it i s still a ,
c a u s e of l ess en i ng th e s u r pl u s of nit r ogen ou s m a t eri a ls F or .
the rep a i r of the m ot or tis su es b ec om es m ore c ostly Fa t .
st ored u p i s weight t o be c a rr i ed A c rea t u re l oa d ed with

-
.
i nert m a tt er mu st oth er thi n gs eq u a l c on s u m e a grea t er

, ,
a m oun t of ti ss u e f or mi ng s u b s t a nces for k ee pi n g its l oco

-
m otiv e a pp a ra t u s i n ord er ; a nd thu s exp endi ng m ore for Self

m a i n t en a nce ca n exp e nd l es s for ra c e m a i n t ena nc e A bn or m a l -
.
pl eth ora i s thu s a nt a g oni s tic t o rep rodu cti on i n a d ou bl e way .
I t ordi na rily impli es a sm a ll er a bs orpti on of ti s s u e f ormi n g -
m a tt er s a nd a n i ncrea s ed d em an d on the dimi ni s h ed s u pply

, .
H enc e fert ility d ec rea s es i n a geom et ric a l p rog ress i on .
T he c ou n t er c on cl u s i on d ra wn f rom fa cts of thi s cl a s s i s

-
,
th en du e t o a misc onc epti on of thei r na t ure— a mi s c onc epti on

,
I t i s worth whil e i nq ui ri ng wh eth er unfi tne s s o f th e food gi v en to them ,
is not th e chi ef ca u s e of th a t steril ity wh i ch , as Mr D a rwin

. sa ys ,
“
i s th e
”
great b a r to th e dom es ti cati on of a ni m a l s He rem ark s th at wh en an i m al s
.
a nd pl ant s a re rem ov ed from th ei r nat ural condi ti ons th ey a re ex trem el y ,
l i abl e to h a ve thei r rep od ct v e y

”
r u i s s t e m s s e ri ou s l y a fl ’
ect ed P os s ib l y th e .
rel a ti ve or a b s ol ut e a rre s t of genesi s, i s l e ss due to a di rect effect on t h e

rep roduct i ve s ys tem ,
th a n to a ch a ng e d nu tri t i on of w hi ch t he re prod uct i v e
s ys te m m os t cl ea rl y sh o w s th e re sul t s . Th e m a tte rs re q uired for form i ng an
em b ryo great er prop ort i on ni troge nou s tha n are the m a t te rs re q u i red
a re in a
for m a i nt ai ni n g an a dul t Hence , an a ni m al for ced to l i ve on i nsuffi ci entl v

.
ni t rog e ni zed food m ay h a v e i t s s urp l us for r e p r od uct ion cut off, but s til l h a v e
.
a su f
fi ci ency t o k eep i t s own t i s su es i n re pa i r, a nd a pp ea r to b e i n good h eal th
— m eanwhi l e i ncreas ing i n bulk from x
e ce s s of the non ni trogenous m a tters
~
it ea t s .
4 84 L A WS OF MUL TIP L I CA TI ON .
ar isi ng p a rtly f rom the c i rcu mst ance th a t the in c rea s e of bu lk

p r odu ced by fa t is s om ewh a t lik e the i ncrea se of bu lk whi c h
g rowth of ti ssu es c a u s es a nd p a rtly from the ci rcu m s t ance
,
th a t a bu nd a nce of g oo d f ood norm a lly p r odu c es a c ert a i n

q u a n ti ty of fa t whi ch w ithi n na rrow li m i t s is a v a l ua bl e
, , ,
s t ore o f f or c e e v olvi n g m a t er i a l
-
Wh en h owev er we limit
.
, ,
t h e ph r a s e high nu t r iti on t o its p rop er m ea ni ng — a n a b un

d a nc e of a nd du e p rop orti on a m ong a l l the su b s t a nces whi ch
, ,
th e org a nism need s — we fi n d th a t oth er thi n gs equ a l f ertili ty

, ,
a l wa ys i nc rea s es a s nu t riti on i ncr ea s es An d we s ee th a t th ese

.
a pp a rently e xc epti ona l c a ses a r e c a s es whi c h rea lly s h ow u s

-
,
the s a m e thi ng ; S ince th ey a re c a s es o f rel a tiv e i nnu t riti on .
N — B
[ O TE y st ra nge overs ight wh en w riti ng this ch a pt er
a
i n th e fi rs t e diti on— a n ov ers ight I wa s on the eve of rep ea t

ing i n t hi s p resen t e diti on— I o mitt ed to b ri ng f orw a rd the
fa mili a r a n d a l l imp ort a n t evid ence fu rn i s h ed by the v a ri a
-
ti on s of genes i s w hi c h ordi na rily a cc omp a ny th e a lt ern a ti ons

o f the s ea s on s T h es e v a ri a ti on s i n m u lti tu di nou s c rea t u res
.
,
o f a l l typ es S h ow u n mi s t a k a bly th a t r ep rod u cti on b egi ns a t

,
th os e tim es of the y ea r wh en grea t er wa rm th a nd l a rger

s u ppli es of f ood ren d er m a i nt ena nce of i ndividu a l life
rel a tiv ely ea s y a nd wh en th ere i s th eref ore a s u rpl u s a v a il
,
a bl e for p ro d u c i ng n ew i ndivi d u a ls C onv ers ely a l ong

.
,
w ith the d ec rea s e of h ea t a n d the rel a tiv e d efi ci ency of f ood

whi c h m a k e it c omp a r a tiv ely di ffic u lt i n wi nt er to m a i nt a i n
i ndivid u a l life th ere cea s es t o be the p ower of p r odu c i ng
,
ot h er liv es : th e re p rod u ctiv e or g a n s be c om e qu i es cent a nd
o ft en d w i ndl e With this genera l f a ct is a ss oci a t ed a sp eci a l

.
fa ct T oh u gh a m o n g wild a n im a l s— bi rds m a mm a ls a nd
.
, ,
o th ers — b reedi n g c ea s es wh en N a t u re no l onger su ppli e s

a b u n d a n t foo d a nd w a r mth ; i n d om es tic a t e d m a m m a l s a nd
bi rd s a rtifici a lly s u ppli ed with f ood a nd wa r mth the breed

, ,
i ng s ea s on i s grea tly ext en d e d a n d o ft en m a d e c onti nu ou s ,
a s u n d er th e s a m e c on diti ons it i s i n Ma n him s elf

, ,
.
E vid en c e yi eld ed by the v eget a l w orld is l ess c onspicu ou s ,

NU TRI TION A ND GE NE SIS .
4 85
for the s
r ea on th a t the c old which a rr ests rep rod uctiv e
a ctivity a l s o a r re s t s i ndivid u a l a ctivity : growth o f th e i n di
vid ua l a nd m u ltiplic a ti on of the ra c e v a ry S imu lt a neou s ly

with va ri a ti on s i n the s ea s on s S till th ere a re s om e f a mili a r
.
f a cts sh owi ng th a t the ext erna l c onditi ons whi ch f a v ou r

nu t riti on a l s o b ri ng a b ou t rep ro du cti on E a rly in the y ea r
.
we a re s u ppl i ed with fl owers f ro m regi on s w a r mer th a n ou r

own a n d by a nd by th ere c om e t o ou r m a rk et s f ru it s a n d
,
v eget a bl es f rom the s ou th of F ra nce the C ha nn el I s l a nd s

, ,
a nd e v e n f rom th e S cilly I s l es ,
which a re mu ch in a dv a n ce
of th ose f u rn i s h ed by the g a rd en s o f ou r own c o ld er re gi on s :
r ep ro d u cti on c o mm en c es ea r li er wh ere the light a nd h ea t
f u rth eri ng nu t riti on a re g rea t er . A nd th en th er e is a

ki n d red mea ni ng i n the n ot u n freq u ent occu rrence of a s ec
ond fl ow eri ng a n d e v en of a s econ d f ru itin g i n w a r m b right
,
a nd p rol on g e d a u t u m n s
. H ere th e a b norm a l rec omm enc e
m ent of rep r odu cti on is d et ermined by a n a bnorm a l i nc rea s e
of n u t r iti on ]
CH A P T E R X .
I I I
S P E C AL T E S OF T H E SE RE L A T ON S I .
356 . T E ST S genera l d oct ri nes set f orth i n p rec ed

of t he
i ng ch a pt ers a re a ffor d ed by org a ni s m s h a vi ng m o d es of l ife
,
which di v erge w id ely from ordi na ry m od es H ere a s el s e .

,
wh ere a b erra nt c a s es yi eld c ru ci a l p roof s

,
.
I f c ert a i n org a ni s ms a re s o ci rcu m s t anc ed th a t highly

nu t ritiv e m a tt er i s s u ppli e d t o th em with ou t s ti n t a n d th ey ,
h a ve n othi ng to do bu t a bs orb it we m a y i nfer th a t th ei r

,
p ower s of p rop a g a ti on will be enorm ou s .
I f th ere a re cl a s s es of c rea t u res which exp en d v ery littl e

for s elf su pp ort i n c om p a ri s on w ith a lli ed cr ea t u res a rel a
-
,
t i vel y ext rem e p r ol i fi cnes s m a y be exp ect ed of th em

-
.
Or if a g a i n we fi n d s p ec i es p resen ti ng th e p ecu li a r ity

, ,
th a t whi l e s om e of th ei r i n divid u a ls h a ve mu ch t o do a nd
littl e to ea t oth er s of th ei r i ndivid u a ls h a v e mu ch t o ea t a n d
,
l ittl e t o do we m ay l ook for g rea t fertility i n th es e l a s t a nd

,
c omp a ra tiv e i nf ertility or b a rr ennes s i n the fi rs t .
T h es e s ev era l a nticip a ti ons we S h a ll fi nd c ompl et ely

v erifi ed .
§ 357 . whi ch lik e the Ra fii es i a cece c a rry th ei r p a ra

P l a n ts , ,
s i ti s m t o the ext ent o f livi ng o n th e j u i ce s th ey a b s or b f rom
o th er pl a nts e xhibit one of th es e rel a ti on s i n th e v eg et a l

,
ki ngd om .I n th em th e orga n s for s elf su pp ort b ei ng n eed

-
l es s a re ru dim en t a ry ; a nd th e p a r ts di rectl v or i ndi rect l y

,
c oncerned i n th e p rod u cti on a nd di s t r ib u ti on of g er m s c on ,
s ti tu te th e m a ss o f th e org a ni s m T h a t s m a ll ra ti o which
.
S P E CIA L I TIE S OF T H E SE RE L A TI ONS .
487
the ra e-c p res ervin g st ru ct u res bea r t o the s elf p res ervi n g -
st ru ct u res i n or din a ry t e nog a m s i s i n th es e P h aenog a m s , , ,
i n v ert ed A lik e rel a t i on occu rs i n the c omm on D odd er

. .
T h ere m a y be a dd ed a kin d red pi ec e of evid ence w hich the

F u ngi p res en t T h ose of th em whi ch g row on livi ng pl a n t s
.
,
rep ea t th e a b o v e connec ti on c o m pl e t ely ; a nd th os e of th e m
which th ou gh n ot p a ra s itic nev erthel ess s u b s ist on org a nized

, ,
m a t eri a ls p revi ou s ly el a b or a t ed by oth er pl a n ts s u bst a nti a lly ,
r ep eat it T he S p ore p rodu ci n g p a rt i s rel a tiv ely en or m ou s ;

.
-
a nd t he f ertility i s fa r gr ea t er th a n th a t of C rypt og a m s o f
lik e s iz es whi c h h a ve t o form for th em selv es the or ga ni c

,
c o m p ou n ds o f whi c h th ey a n d th ei r g er ms c on sist .
3 58 . Th e
m e l es s on i s t au ght u s by a nim a l p a ra sit es
sa -
.
A l ong wi t h th e d ecrea s ed c o s t o f I n dividu a ti on th ey simil a rly ,
S h ow u s a n i n c r ea s ed exp en dit u re for Gen es is ; a nd th ey sh ow
u s thi s i n the m os t s t r iki ng m a nn er w h er e t he d evi a ti on f r om
or di n a ry c on diti on s o f lif e i s the g r ea t es t .
T a k e a m ong th e E pi zoa s u c h a n i n s t a n ce a s Ch on dra ca n

, ,
thus gi bbos u s B el on gi n g t o the E n tom os tra ca b o th m a l es

.
,
a nd fem a l es o f thi s s pe ci es a r e i n th ei r ea rly d a ys s i m il a r t o

, ,
th ei r a lli es ; a nd th e m a l es p ra cti ca lly p a ra s iti c th ou gh th ey

, ,
bec om e g rea tly d egra d ed c onti nu e th r ou gh ou t li fe to S h ow

,
by th ei r s egm ent a ti on a nd oth er ext ern a l t ra its th ei r origi na l

n a t u re . T h e fem a l e h owev er h a vi ng fix e d h er s elf wh ere
, ,
s h e c a n s u c k th e j u i c es o f h er h os t the L op hi us grows t o , ,
t welve ti m es the l ength o f the m a l e a nd p rob a bly a th ous a nd

tim es i ts bu lk a n d b ecom es u tt erly t ra ns f orm ed by l os s of
,
th e orga ns of a ni m a l li fe a nd en orm ou s d ev el op m ent of th e

o r g a n s o f r ep r od u cti on N o h ea rt i s di s c ov era bl e a n d the
.
,
nerv ou s sy s t em a n d o r g a n s of s en s e ( if a ny ) a r e eq u a lly u n
di s ti ngu i s h a bl e T he i nt ers p a c e bet ween th e a lim ent a ry

.
ca n a l a nd th e w a lls o f th e b ody i s a l m os t wh olly o cc u pi ed

”
by the ov a ri u m A n d th en b ey on d thi s th ere a re a p
.
pend ed ovi s a cs t wic e th e l ength of the b ody S o th a t th e

-
.
Huxl ey A na tom y of I nver tebr a ted A ni m a l s p 2 74

, ,
. .
488 L A WS OF MUL TIP L I CA TI ON .
g erm p rod u ci ng organs a nd thei r c ont ents event u a lly

-
,
a cqu i r e a t ot a l b u lk m a ny ti m es th a t o f a l l th e oth er org a ns
p u t t og eth er .N u m erou s s p eci es of this type a nd h a bit ,
rep ea t thi s rel a ti on b et ween a life of i na c ti on with high
f ee di ng a nd a n enorm ou s ra t e of g enes i s P a ra s it es bel ong

,
.
i ng t o a n oth er grea t divi s i on of the a nim a l ki ngd om th e ,
P l a tyhel m i n thes s u pply a n ex a mpl e of a n epi zoon i n whi ch

,
t he ra t e of mu ltiplic a ti on is m a d e grea t n ot s o mu ch by
immens e d evel opm ent of the egg p rodu ci ng orga n s a s by -
the r a pidity w ith which genera ti on s s u cc eed one a n oth er— a

r a pidity s u c h th a t ea c h g e nera ti on p a rti a lly d ev el op s th e
n ext b ef ore it i s it self a nythi n g l i k e rea dy for ind epe n d ent
li fe T hi s i s the Gyroda c tyl us el ega ns of w hich it i s s a id

.
,
“
th a t i ts m os t rem a rk a bl e f ea tu re i s th a t it i s vivip a rou s ,
a nd its em b ry os b e f ore th ey l ea v e th e b ody o f th e i r m oth er
h a ve a l rea dy d ev el op ed th ei r emb ry os i ns id e th em ; a nd the

l a tt er m ay c ont a i n th ei r em b ry os s o th a t f ou r genera ti on s
,
m a y be i ncl u d ed un d er the c u ticl e of the s exu a lly m a t u re

”
a ni m a l .
E n tozoa yi eld u s m a ny ex a mpl es of thi s c a u s a l rel a ti on ,
ra is e d t o a still high er d eg ree T h e Gor di u s or H a i r w orm

.
,
-
,
i s a crea t u re which fi ndin g its w a y wh en y ou n g i nt o th e

,
bo dy o f a n i ns ect which i s a ft erwa rds s w a ll owed by a fi sh,

there gr ows r a pidly a nd th en em er gi n g t o b reed l a ys a s
, ,
m a ny a s egg s i n l es s th a n a da S imil a r ly with

y .
th os e l a rger type s i nfes ti ng the hi gh er a ni m a l s I t h a s been .
c a lcu l a t ed by D r E s chri ch t a s qu ot ed by P rofes s or Owen

.
, ,
th a t th ere a re o f ova i n th e m a tu re fe m a l e
A s ca r i s l u m bri c oi des Very m a ny of th e E n tozoa b el ong t o

.
t he P l a tyh el m i n thes a nd a m ong th em occu r ex a mpl es of fer

,
t i l i ty c a u s ed not only by g rea t nu m b er s of ova bu t by r a pid ,
s u c c es s i on of p a rti a lly d ev el op ed i n divid u a l s a nd a l s o ex a m

-
pl es of fertility c au s ed by p rodu cti on of ova a lm ost exc eedi ng

nu m er a ti on . A m ong the fi rst the L iv er fl u ke m a y be na m ed
-
.
Of the h a lf milli on eggs it p rod u c es ea ch yi elds a f ree swim

- -
Shi pl ey Z ool ogy

, of I nvertebrata , p 112
. .
S P E CIA L I TIE S OF TH E S E RE L A TI ONS . 4 89
m in g cili a t ed em b ry o a nd a ny one of th es e which fi nds i ts

, ,
wa y int o a w a t er s na il bec om es a sp or ocy s t— a ba g p res ently

-
, ,
o cc u pi ed e xcl u s iv ely by m a ss es o f c ell s : ea c h m a ss by a nd
by bec omi n g a Redia whi ch m a k es i ts wa y ou t L ik e a l l

,
.
its fell ows whi ch d ev el op i n su cc ess i on thi s with the excep , ,
ti on of a sm a ll s p a ce occ u pi ed by the s t om a ch d ev ot es the ,
wh ol e of its i nteri or p a rtly t o th e f or m a ti on of other Redi ce

hich p tly s p d b s i m il a r ly t ra ns f or m ed )
( w res en e c a e a n ec o m e ,
a nd p a r tly t o th e d ev el op m ent o f C er ea ri ce i n t o which th e ,
i nterna l s u b s t a nc e of a l l the Redi ce i s event u a lly t rans form ed

C eroa ri ce which esca pi ng from th e h o s t bec om e a gen ts for
, ,
i nf ecting oth er c rea t u res S o th a t ea c h o vu m th u s giv es

.
ri s e t o a nu m be r o f f orm s w hich s ev er a lly s u b s erv e m u lti
plic a ti on i n di fferent wa y s Of the oth er divi s i on of P l a ty

.
hel m i nthes referre d t o a s ca rryi n g on i ts mu ltiplic a ti on by

p rod u cti on of ova only the c om m ones t of the Ces toi dea
,
f u rn i s h es the b es t ex a m pl e I m m ers ed a s a T a p e w or m i s i n
.
-
nu t ritiv e liqu id w hi c h it a b s orbs th r ou gh its i n t egu m ent it

, ,
req u i res no diges tiv e a pp a ra tu s The roo m whi c h one w ou ld

.
oc c u py a nd the m a t er i a l s it w ou ld u s e u p a re th er ef ore a v a il
, ,
a bl e for g er m p ro d u c i ng org a n s w hich n ea r ly fill ea ch seg

-
,
m ent : ea ch s egm ent s exu a lly c om pl et e i n it s elf is littl e el s e

, ,
th an a n enorm ou s rep rod u ctiv e s y s t em w ith j u s t enou gh of ,
o th er st ru ct u re s t o bi n d it t og eth er Rem em beri ng th a t the

.
T a pe worm ret a i ni n g its h old c onti nu es t o bu d ou t s u c h

-
, ,
-
s egm ent s a s fa s t a s the fully d ev el op e d ones a re c a s t off a nd

-
,
g oes on d oing this a s l ong a s the i n fes t ed i n dividu a l lives ;

we s ee th a t h er e wh ere th ere i s n o e xp en dit u re w h ere the
, ,
c os t of i ndivid u a ti on i s redu ced t o the grea tes t ext en t whil e

th e nu t riti on i s t he high es t p oss ibl e the d egr ee o f f ert ility ,
rea ch es i t s ext r em e . T h e s e E n tozoa yi eld u s f u rth er

i nt eres ti ng evid enc e Of th ei r v a r i ou s s p ec i es m os t if n ot
.
,
a l l u nd er g o p a s s iv e m igr a ti on fr om a n i m a l t o a ni m a l be fore
th ey b ec om e m a t u re U s u a lly the f orm a ssu m ed i n th e b ody

.
,
o f th e fi r s t h o s t i s d ev oid o f a l l th a t p a rt i n w hi ch th e rep r o
duc ti ve st ru ct u res t a k e th ei r ri s e ; a nd this p a rt gr ows a nd

490 L A WS OF M UL T I P L I CA TION .
d ev el op s rep rod u ctive st ru ctu res only i n s om e p red a t ory

,
a ni m a l t o w hi ch its fi rs t h os t f a lls a s a crific e Occ a si on a lly .

,
h owever the egg giv es origin t o the s ex u a l f or m i n t he

,
a ni m a l th a t origi na lly s wa ll ow e d it bu t th e d ev el op m ent

,
rem a i n s i n c ompl et e— th er e i s n o s exu a l g enes is n o f or m a ti on ,
o f eggs i n t he ru di men t a ry s egm en t s T h a t thes e m a y .
becom e fertil e it i s nee d ful a s b efore for th e cont a i ni ng

, ,
a n im a l t o be d ev ou r e d ; s o th a t t h e i m p er f ec t T a p e w or m -
m a y fin d its wa y i n t o th e i n t es ti ne of a high er a n i m a l .
T hu s th e B othri ocep ha l us s oli dus f ou n d i n the a bd omi n a l

,
c a vity of th e S ti ckl eb a ck i s b a rren whil e it rem a i ns there ;

,
bu t if t he S ti c kl eb a c k be ea t en by a Wa t er f ow l the re -
,
p rod u ctive s ys t em of the t ra ns ferred B othri ocep ha l us ( th en

k nown a s B n odos us ) b ec om es d ev el op ed a nd a ctiv e S o t oo
. .
, ,
a ki n d o f T a p e w or m w hich rem a i ns in fertil e whil e i n th e

-
i nt es ti ne of a M ou s e b ecom es fertil e i n th e i nt es ti ne of a Ca t
,
th a t d ev ou rs the m ou se Ma y we n ot reg a r d thes e f a ct s a s

.
a g a i n s h owi ng th e d e p en d en ce of f ertility on nu t riti on ?
B a rrenn es s h ere a c com p a n i es c ondi ti on s u nf a v ou r a bl e t o th e

a b s orpti on of nu t ri m ent ; a n d it g iv es w a y t o fe cu ndity wh ere
nu t rim ent is l a r g e i n qu a ntity a nd s u p er i or i n qu a lity .
o s e c a s es o f p a rt i a l
§3 59 . xtEr e m ely s ignific a n t a r e th
rev ers i on t o p r i m itiv e f or m s o f g en es i s w hi ch oc cu r u n d er,
s p eci a l c on diti on s i n s om e of the hi gh er A n nu l os a I refer .
t o th e p s eu d o p a rth en ogenes i s a nd m et a genes i s i n I ns ect s

-
.
Un d er wh a t con diti on s do th e A p hi des e xhibit thi s s t ra nge

d evi a t i on f rom th e h a bit s of th ei r ord er ? Why a m ong th em
s h ou ld i m p er fect f em a l es p r od u c e a g a m i c a lly
,
o th er s lik e ,
th em s elv es g enera ti on a ft er genera ti on wi th g rea t ra pidi ty ?

, ,
T h er e i s the obvi ou s expl a na ti on th a t th ey g et pl en ty of

ea s ily a ss imil a t e d f oo d w ith ou t ex erti on
- P i erci ng th e t end er
.
c oa ts of y ou ng s h oot s th ey s i t a nd s u ck— a pp r op ri a ti ng the

,
n it r og en ou s el em en ts o f t h e s a p a n d ej ec ti ng i t s s a c ch a
”
ri ne m a tt er a s h oney dew
“
A l ong w ith a s l u ggi s h nes s
.
s t rongly c ont ra s t e d with the a ctivity of m ost i ns ects— a l on g

S P E CI A L IT IE S OF T HE SE RE L A T IONS .
491
with a v ery l ow ra t e of c ons u mpti on a nd a c orrel a tiv e d eg ra

d a ti on of s t ru ct u re ; w e h a v e h ere a ret rogres s i on to a s ex u a l
genes i s a nd a g rea tly i ncrea s ed ra t e of mu ltipli ca ti on
,
-
.
T h e rec ently di s c ov ered i ns t a nc e of i nt ern a l m et a genes is

i n the m a gg ots o f c ert a i n Fli es h a s a lik e m ea ni ng In .
c redible a s i t a t fi rs t s ee m ed to na t u ra li s ts it is n ow p rov ed ,
tha t th e C ecydom ia l a rv a d ev el ops i n its i nt eri or a b rood of

-
l a rvae of lik e st ru ct ur e wi th it s elf I n thi s c a s e a s i n the .

,
l a s t a b un d a n t f oo d i s c ombi ned with l ow exp endit u re T h es e

,
.
l a rvae a r e fou n d i n s u ch ha bit a ts a s the refu s e of beet root -
s u ga r f a ct ori es — m a s s es of nit rogenou s debri s rem a ini ng a fter

the e xt r a c ti on o f th e s a c ch a ri ne m a tt er E a ch l a rv a h a s a .
p ra ctic a lly u nlimi t ed s u pply of s u st enan c e imbeddi ng it on

-
*
a l l s id es .
I t i s t ru e th a t s om e oth er m a gg ot s a s th os e of th e Fl es h ,
fl y a re simil a rly or s till b ett er c i rc u m s t a nced a nd it m a y

, , , ,
be s a id ou ght th eref ore t o h a v e th e s a m e h a bit

, B ut t h is .
d oes not nec es sa rily f oll ow S u r viv a l o f th e fitt es t will .
d et erm i ne wh eth er su ch s p eci a lly f a v ou r a bl e c on diti on s res u lt -
i n a ggr a n diz em ent o f the i ndivid u a l or i n m u ltiplic a ti on

o f t h e ra c e A n d i n th e c a s e of th e Fl es h fl y th ere is a
.
-
r ea s on why gr e a t er individ u a t i on r a th er th a n m ore ra pid
genes i s will oc cu r F or a d ecomp os i ng a ni m a l b ody l a s t s so

.
s h ort a ti m e th a t w ere Fl es h fl
y l a rv ae t o mu ltiply a g a m ic a lly
-
, ,
the s ec on d g enera ti on w ou ld di e f rom the di s a pp ea r a nc e of

th ei r food H en c e i ndivi d u a ls i n which th e exc es s iv e
.
nu t r iti on l ed t o i nt ern a l m et a genes is w ou ld l ea v e no

p os ,
t eri ty a nd n a t u ra l s el ecti on w ou ld es t a bli s h t he v a ri ety i n

,
I am t ol d th a t “
Wa gner ,
wh o des cri b ed t he l a r va , foun d th a t i t b ore d
i nto th e b a rk of t re es . It a tta ck s al so th e whea t p l a nt , a nd is a m os t
de s tructi v e p aras i te .
”
A p p a rentl y thi s s t ate m e nt is at v a ria nce wi th th e
foregoi ng i nference . It i s cl ea r, h owe v er, th a t s i nc e the s e h ea p s of ni t ro
genou s re fu s e i n whi ch i t h a s b e en found a re a rtifi ci a l and re cent , t hey ca n

no t b e i ts na tu ral h ab i t a ts ; a nd i t s eem s n ot i m ro b a bl e th a t th e s e l a rv ae
p ,
s uddenl y s uppl i ed wi th a m ore nutr iti ve foo d i n unl i m i ted a m oun t, m a

y ha v e
as a con seq u ence ac q ui red th i s h a b i t o f a ga m o ene ti c m ul t i l i ca ti on
g p wh i ch
di d not ch a ra cteri ze th e s p ec i es under i ts na tu ral condi ti ons and rel a ti el
y
v l ow
nut ri t i on .
49 2 L A WS OF MUL TI P L ICA TI ON .
which grea t er gr owth resu lt ed A l l whi c h the a r gu m ent

.
r equ i res is th a t w h en s u ch rev er s i on t o a g a m o g en es i s does
t a k e pl a ce it s h a ll be wh ere the f oo d i s un u s u a lly a bu nd a nt

,
a n d th e e xp en dit u re u nu s u a lly sm a ll ; a nd thi s th e c a s es
i ns t a nc ed go to S h ow .
§ 3 6 0 T he physi ol ogic a l l ess on t au ght u s by Bees a nd

.
A nt s not q u it e h a r m oni zi ng w ith th e m ora l l ess on th ey a re

,
s u pp osed to t ea ch i s th a t highly fed idleness is f a v ou ra bl e t o

,
-
f er tility a nd th a t exces s ive i n dus t ry has b a rrenn es s for its

,
c oncomit a nt .
The egg of a B ee d ev el op s i nt o a sm a ll b a rren fem a l e or

i nt o a l a r ge f ertil e f em a l e a cc ordi ng t o the s u pply of f ood
,
giv en t o the l a rv a h a t ched f rom it We h ere s ee th a t the .
germ p rod u ci ng a cti on is an ov erfl ow of th e s u rplu s rem a in

-
i ng a ft er c ompl eti on of th e i ndivid u a l ; a n d th a t the l ower

f ee di ng which the l a r va of a w ork ing Bee h a s resu lts i n a ,
dwa rfing of th e a d u lt a nd a n a rres t ed d ev el opm ent o f the

g enera tiv e orga n s F u rth er we h a ve the fa ct th a t the c on
.
,
diti on u nd er whi ch the p er fect f em a l e or m oth er Bee g oes ,

-
,
on ,
u n lik e i n s ec ts i n g ener a l l a yi ng eggs c ontinu ou s ly is
, ,
th a t She h a s pl enty of f ood b r ou ght t o h er is k ept wa rm a nd , ,
g oes th rou gh no c ons id era bl e ex er ti on Whil e c ont ra riwi s e .

, ,
it i s t o be not ed th a t the i n fertility of the w ork er s i s a s s o

c i a ted wi th th e c ea s el ess l a b ou r of b ri n gi n g m a t er i a l s for th e
c ombs a nd bu ildi ng th em a s well a s the l a b ou r of feeding

,
.
t he qu een the l a rv ae a n d th em selv es

, , .
A nts a l s o S h ow u s th es e rel a ti ons a n d th ey a r e S h own i n a

,
gr ea tly ex a ggera t ed f orm by w h a t a re c a ll ed whit e a nt s

i nsect s b el ongi ng t o a qu it e di fferent ord er T he c on t ra st .
i n b u lk b et w een th e f ecu n d a n d i nfecu n d fem a l es i s h ere

imm en s ely g rea t er T h e m oth er A nt h a s the rep rod u ctiv e
. .
-
s y s t e m s o en orm ou s ly d ev el op ed th a t the r em a i n d er of h er
,
b ody i s rel a tiv ely i ns ignific a nt E nti rely i nc a p a bl e of l oc o

.
m oti on she is u na bl e t o d ep os it h er eggs i n the pl a ces wh ere

,
th ey a re to be h a tch ed ; s o th a t th ey h a v e to be c arri ed a wa y
S P E CIA L IT I E S OF T H E S E R E L A TI ONS .
493
by the w ork er s a s fa s t a s th ey a re e xt ru d ed H er lif e i s thu s .
re d u c e d s u b s t a nti a lly t o th a t o f a p a ra s it e
— a n a bs orpti on of
a b u n d a nt f oo d s u ppli e d g ra ti s a t ot a l a b se nc e of ex pen di
,
t u re a nd a c ons equ ent exc essiv e ra t e of g enes i s

,
T he .
qu een a nt of the A f ric a n T er m i tes l a ys

-
eggs i n twen ty
four h ou r s .
§ 3 6 1 I t m ay be needfu l to sa y th a t th ese exc epti ona l

.
rel a ti ons c a nn ot be a s c r ib ed t o th e a s s ign ed c a u s es a c ti n g
a l one . The e xt re m e fert ility w hich a m ong p a ra s it es a nd ,
s oci a l i ns ect s a ccom p a n i es ext rem ely high fee di ng a nd a n

,
exp en dit u re red u c e d nea r ly t o z ero p res u pp oses typic a l s t ru e

,
t u res a n d t end enc i es o f s u it a bl e ki nd s ; a nd thes e a re not

di rectly a ccou nt e d for On c rea t u res oth erwi s e org a ni zed
.
,
un li m it e d s u ppli es o f f oo d a n d t ot a l i na c tivity a re n ot fol
l owed by s u ch res u lt s T h ere of c ou rs e requ i res a c ons ti

.
t u ti on fitt e d t o the s p ec i a l c on diti on s a nd th e ev ol u ti on o f ,
thi s ca nnot be du e s i m ply to pl eth ora j oi n ed w ith res t T h es e .
c a s es a re given a s illu s t ra ti ng the c onditi ons u nd er whi c h

ext rem e ex a lt a ti ons o f f ertility b eco m e p os s ibl e T h ei r m ea n .
i ng s th u s li m it ed a re c l ea r a nd c om pl et ely t o the p oin t We

, , , .
s ee i n th e m th a t t h e d ev oti on of nu t r im en t t o ra c e p res er v a -
ti on i s ca rri ed fu rth es t w h ere the c os t of s elf p res erv a ti on

,
-
i s red u c ed t o a mi n imu m ; a nd c on v er sely th a t n othi n g i s

, ,
d ev ot ed di rectly t o ra ce p reserv a ti on by i ndivid ua l s on w hi c h

-
f a ll s a n exc e ssiv e exp endit u re for s elf p res erv a ti on a nd p re -
’
s erv a ti on of other s off s p ri ng .
— sp eci a liti es of th es e rel a ti ons m ay be fitl y

[ N O TE A m ong
a dd ed h ere a v ery s t ra n ge o ne for a d es c r ipti on of w hi ch I
,
a m i nd ebt ed t o M Ch a r l e s J u l m
. P ro fes s or of C omp a ra tiv e
A n a t omy i n th e Univ er s ity of L i ege I n th e Revu e Gén éra l e
.
des S ci ences for 3 0th A u gu st 1 8 9 4 i n a n a cc ou nt of c ert a i n

, ,
i nv estig a ti ons of M Gi a r d he d es c rib es wh a t he c a lls l a

.
,
“
c a st ra ti on p ara sit a i re —a c a st ra ti on n ot of a lit era l ki nd

4 94 L A WS OF MU L TIP L I CA TION .
bu t one ef
fect e d by th e a rrest of d ev el opm ent whi ch f oll ow s
f rom the d epl eti on c a us ed by a p a ra s it e T he S a ccu l i na i s .
a n a m a z in gly t ra n s f or m e d typ e be l on gi ng t o t h e Ci r rhi p edi a
— a typ e wi th ou t s eg m ents or a pp end a ges a nd with ou t m ou th

a nd a lim ent a ry c a n a l Fixi ng it s el f du ri ng its ea rly l oco
.
,
m otiv e st a g e un d er the a bd omen of a d ec a p od ou s c ru s t a c ea n

, ,
an d l ea vi n g b ehin d i t s exo s k el et on it m a k es its wa y i nt o -

,
th e i n t eri or a n d th ere bec om i ng a m ere ba g c ont a i ni ng the

,
rep rod u ctiv e org a ns o bt a i n s th e need fu l nu t rim ent by de

,
ve l op i ng w h a t a re p r a ctic a lly r oot s a n d roo tl et s w hi c h run
e v erywh ere a m on g the vi s c er a a nd a b sorb n u t r im en t f ro m
th e s u rrou n di n g ti ssu es H ere we a re c oncerne d m erely.
w ith the efi ect p rod u c e d u p on th e h os t by thi s phy s i ol ogi ca l

r obb ery T hi s effec t i s t o a rres t th e d ev el opm en t n ot on ly
.
o f th e p r i m a ry s ex u a l or g a n s d ev ot e d t o t he p ro d u cti on of
ger m s bu t a l s o of th os e s ec ond a ry s exu a l org a n s which ch a r

,
a cteri z e th e m a l e M J u lin wr it es . .
11 c on vi ent c e p e nd a n t d e d i re , p ou r etre p l us e x act , qu e , d ans

l es cas d e s C ra b es i nfest és pa r d es S a c cu l i nes , i l n y
’
a p as , e n r éa l i t é ,
a pp a ri t i o n de cara c t er es fe m e l l es c h ez le se xe m Al e, m ai s p l u tbt
a b se nce d e d é v el opp em e nt d es ca ract eres m Al es . E n fai t , l a ni m a l
’
res t e a n u s ta d e jeu ne ,
n on d i ffé re nc i é se x u el l em ent , to ut en p re na nt
u ne t a i l l e pl u s c ons i d era bl e C el a nou s p orte a a ttri bu er
. l e s m odi
fi c at i ons d ont nou s a v on s p a rl a un s i m p l e arret d e d v e l oppe m
é é e nt ,
se ns i bl e le
Al e , p a rc e qu e c h e z l u i l es c arac tere s
qu i e st p l us c h ez m
se x u el s se c o nd a i res sont a l eta t norm a l

’
pl us d eve l Opp és qu e c h ez l a
fe m el l e .
D
’
u ne m a ni é re
g é néra l e , n ou s c royons , a e c v M Gi ard .
,
qu i l
’
fa ut
as s i m i l er l es m odi fi ca ti on s d ue s a l a cast ra t i onp a ra si tai re a c el l es q ui
le ré s ul t at d e l a p rog e nes e
s ont ou qu i e ng e nd rent l e d im orp h i sm e
sa 1 s onn1 er .
1 1 y a p r ogenese l ors q u e , c h ez u n a ni m al , l a re prod u cti on se x u ée

s op are d u ne fa co n pl u s ou m oi ns p r éc oc e , c est a-di re l ors u e l es
’ ’ ’
q
-
rod ui ts se x u e l s ( ce n ts on s e rm a t oz oi d e s
p p ) s e torm e nt e t m fi ri s sent
a v a nt qu e l etre
’
n a i t a tte i nt s on c om
’
p l et d é v el oppe m ent On pe u t .
c i te r c om m e e x em pl es l es A x ol ot l s e t l es l a r v es d e T ri to ns
qu i , l e s u ns
n orm al e m ent , l es au t re s ac c i d e nt e l l e m e nt,
p ond e nt en aya nt e nc ore
l eu rs b ra nch i es .
Tres s ou ent v 1 a proge nese n af

fecte ’
l xe T anté t,
'
’
qu u n seu se . c est
S P E CIA L IT I E S OF T H E S E R E L A TI ONS .
495
le se xe fe m el l e q u i m uri t a l eta t l arv a i re

’
c om m e c h ez l e s p uc eron s,
l es Beg/lop e,
’
c e st le se xe m el e , com m e c h ez l a B onel l i e , l es
m al es co m p l ém ent ai re s d e Ci rri p ed e s , l es m el es pyg m é es d es Rotifere s ,
foi s , e nfi n, l a ni m al p r ése nte
’ ’
al e d e l A ng u i l l e , e tc
’
le m D a u t res .
s u c ce s si v em e nt l e s d eux s e x es a v e c p rog e nes e p o u r l un d e nt re e ux

’ ’
.
C e st a i ns i q u i l y a p rogenese p r ota nd r i qu e ch e z l es C rus tac é s cym o

’ ’
t h oa d i e ns , et, pa rm i l es Ve rt é brés , ch ez l es My xi ne s, q ui , m el es d a ns
le j e u ne ag e , d e v i ennent fe m el l es e n vi eil l i s sant et e n ac he v ant de
p rendre l e u r d é v el o pp em e nt . Le cas d es vi eil l es fem el l es d e Ga l li

n ac é s a p l u m a g e e t a i n s t i nct s m a sc u l i ns se m bl e é tre , a u contrai re , n u
e x em p l e
i m p ar fai t d e p r ogenése p r otoggni gu e, p ui squ e c es fem el l es o nt
p ond u l ors q e e a v e t e i e l a l v é e d es je
’
u l l s a n n c o r i r u n
’
e s e t q u e l l es o nt
c o nt i nu é pl us t ard l eu r d é v e l oppe m e nt , e t p r é se nt e nt l s ca rac t ere d e s
m el es sa ns q ue, cep enda nt , l

’
on ai t c onsta t é 1 a p rod u c ti on d e sp e r
m a t oz o i d es .
D an s l es cas e x trem es d e p rog enese fe m el l e , l a reprod u cti on se fai t

de l el é m ent m al e , re v e na nt ai ns i a l a form e
’
m em e sa ns le c o nc ours
a g am i qu e p ri m ord i a l e Ces . cas sont c onnu s d epu i s l ong t em p s sou s l o

n om d e p édogenése On l es
. 9. obs e rv e c h ez l es l arv e s d e M i a stor , d e
C hi ronom us e t c h ez ce rt ai ns pu c erons .
qu il y a
C h a qu e foi s
’
.
p rog e nese d a ns
nu typ e d ete rm i ne on c ons ,
ta te s oi t m om e nt a n ém e nt soi t d u ne fa co n d efi ni ti v e un a rret d e
’
, ,
c ro i ss a nc e e t d e dé v e l o ppe m e nt : l a ni m a l p rog é né ti qu e a p ar s u i te
’
, ,
l as p e ct d u ne l arv e s e x u ée l ors qu on l e c om p a re soi t a l a ut re s e x e

‘ ’ ’ ’
, ,
s oi t a u x form e s v oi s i ne s qu i ne p ré s ent ent pa s l e p h é nom é ne d e l a

,
p ro g enes e .
C el a e st enp arfa i te h a rm oni e a v ec l e p ri nc i p e , Si bi en m i s e n

’
l u m i ere p a r H e rbe rt S pe nc er, d e l a nta goni sm e ent/re la genése et l a
croi ssa nce et entre genése
la et l e développ em ent . C et a nta g oni s m e
qu e l es m at é ri au x e m pl oy é s p ou r
’
s e x pl i qu e fa ci l em ent s i l o n
’
s ong e
p v i ’
la re
prod u ct i on ne e u e nt se r v r a l a c c ro i sse e t d e l i nd i vi d u
m n
’
.
S il
’
es t a v a nt ag eu x p ou r nu org a ni s m e de s e r e p rod u i re sa ns a c u
q é ri r
d es org a nes i nu t il es , l a s él ec ti on n at urel l e d é term i nera bi entbt u ne
p rog e n ese d e pl u s en pl u s c om pl ete L es a ni m au x p ar as i tes , ou t re.
t i rent d e l eur h 6te u ne n ou rri tu re abond a nte , n ont g u ere b es oi n

’ ’
qu i l s
d u ne fou l e d org a nes qu i se r v e nt a l eu rs c ong é nere s l i bres d a ns l a
’ ’
A us s i v oyo
’
v i e d e rel a ti o n q nn tres g ra nd nom bre d a ni
’
n s.n ou s u -
m au x p aras i tes sont p rog é n é ti qu es L e s m fil es p rog é né t i qu es d e l a

.
Bo nel l i a et d es C i rri p ed es v i v e nt e n p arasi te s d ans l eu rs fe m el l es .
C h ez c e rt ai n s typ e s , l es p u cer ons , l a p rog en ese c es s e d é s qu e , l a

n ou rri ture d e v enant m oi ns a bond ante , nu d é pl ac em ent pourra é tre
496 L A WS OF M UL TI P L IC A TI ON .
En ré s um é , l a rr et
’
d e d é v el op p em d d 51 l a p rog enese r és ul te
e nt
no u rri c i ers a u d etri m e nt d e l a ni m s l

’
d ’
unederi vati on d es p ri nc i pe s
ro é né t i u e D a ns l e s e x e m p l es d e p ar as i t ai re qu e n ou s
cas t ra ti o n
p g q .
a v o ns l o p a ras i t e jou e , p a r
e x a m i n es , ra p p or t a s o n h ot e, a b sol um e nt
l s m em e rol e qu e l a g l a nd e g é ni ta l e d un t ype p rog é né ti q u e

’
11 .
d ét o ur ne , p our se propre su bs i s ta nce , u ne p a rti e d es pri n ci p es qu i

a urai e nt servi a u d é v e l opp e m e nt d e l ani m al
’
A us si l es efi et s p rod ui ts
.
”
s ont -il s t out a fai t d e m em e ordr e .
Aphenom enon so a n om a l ou s a s this explica bl e u p on the ,
hyp oth esis set f ort h bu t not oth erwi s e explic a bl e, fu rni sh es
st riki ng v erific a ti on ]
CH AP T E R X I .
I N TE RP RE T A T IO N A ND ! UA L I F I CA T I ON .
C O N S I D E RI N G th e nd u ctiv e veri fi ca
di fficu lti es
i of
§ 3 62 .
ti on we h a v e I thi n k a s c l ea r a corr es p on den c e between the

, , ,
a p r i or i a nd a p os ter i ori c on cl u s i on s a s c a n be exp ec t ed ,

T he .
m a ny fa ct or s c o op era ting to b ri ng a b ou t the res u lt i n every

-
c a s e a re s o v a ri a bl e i n th ei r a b s olu t e a nd r el a tiv e a m ou nts

, ,
th a t we ca n ra rely di s ent a ngl e the effect of ea c h one a nd

h ave u su a lly t o be c on t ent with qu a lifi ed i nferences T h ou gh .
i n the m a s s org a n i s m s S h ow u s a n u nmi s t a k a bl e rel a ti on

be t ween g rea t si ze a n d s m a ll fertili ty yet s p eci a l c omp a ri ,
s on s a m on g th em a re nea rly a l wa ys p a rti a lly viti a t ed by
di fferen c es of s t ru c tu re di fferences of nu t riti on differenc es of

, ,
exp en ditu r e T h ou gh it i s b ey on d qu es ti on th a t th e m ore

.
c om pl ex org a nisms a re th e les s p rolific yet a s c om pl exity ,
ha s a c ert a i n genera l connexi on w ith b u lk a nd in a n im a ls ,
with exp endit u re we ca nnot oft en identify i ts res u lts a s i nd e

,
p end ent o f th es e A nd S imil a rly th ou gh the c rea t u res whi ch

.
, ,
w a s t e mu ch m a t ter i n p r od u c i ng m oti on s en sibl e a nd i n sen ,
sibl e h a v e l ower ra t es of m u ltiplic a ti on th a n th ose w hich

,
wa st e l ess yet a s the c rea t u res which w a s t e mu ch a re

, ,
g enera lly l a rger a n d m ore c ompl ex we a re a ga i n m et by a n ,
ob s t a cl e which limits ou r c omp a ri s on s a n d c omp els u s t o ,
a ccept c onc l u s i on s l e ss d efi nit e th a n a r e d es i r a bl e .
S u ch di ffic u lti es a ri s e h owe v er on ly wh en we end ea v ou r

, , ,
a s i n f or eg oi n g ch a pt er s to p rove th e i nv ers e v a ri a ti on
,
78
498 L A WS OF MU L TI P L I CA TION .
b etween Genes i s a nd ea ch s ep a ra t e el ement of I ndivid u a ti on

— g rowth
,
d evel op m ent a cti vity We a re s c a rcely a t a l l
,
.
h a m p ered by qu a l ifi c a ti ons when f ro m c ont empl a t i ng th ese ,
s pe c i a l rel a ti ons we ret u rn t o th e g enera l rel a ti on

,
T he .
a nt a g on i s m b etween I n di vidu a t i on a n d Gen es i s i s s h own by

a l l the f a c t s w hi c h h a v e been grou p ed u n d er ea ch h ea d We .
h a v e s een th a t i n a s c endin g from th e l owes t t o th e high es t

type s th ere i s a dec r ea s e of f ertili ty s o gr ea t a s t o be a bs o
,
l u tel y i n c onceiv a bl e a n d ev en i nexp res s ibl e by figur es ; a nd

,
w h eth er th e s u p er i ori ty of typ e c on s i s t s i n rel a tiv e l a rg eness ,
i n grea t er c ompl exity i n high er a ctivity or in s om e or a l l of

, ,
th es e c om bined m a t ters n ot t o the u ltim a t e i nferen ce T he

,
.
b roa d fa ct enou gh for u s h ere i s th a t org a ni s ms i n whi ch

, ,
th e i nt eg ra ti on a nd di fferenti a ti on o f m a tt er a n d m oti on h a v e
b een c a rri ed f u rth es t a re th ose i n whi ch the ra t e of mu ltipli
,
ca ti on h a s f a ll en l ow es t H ow mu ch of the d ec l i ne of rep ro
.
du cti ve p ower i s du e t o th e gr ea t er i n t egr a ti on of m a tt er ,
h ow m u ch t o i t s grea t er differen ti a ti on h ow m u ch t o th e ,
l a rger a m ou nt s o f i nt eg ra t ed a nd di fferenti a t ed m oti ons gene

r a t ed it m a y be imp os s ibl e t o s a y ; a nd it is n ot needfu l t o
,
s ay. T h es e a re a l l el em ent s of a high er d e gree of li fe a n ,
a u gm en t ed a bility t o m a in t a in the org a n ic eq u ilib ri u m a m id
envi r on i n g a cti on s a n i n c rea se d p ow er of s el f p reserv a ti on ;

,
-
a nd we fi nd th ei r i nv a ri a bl e a cc omp a ni m ent t o be a di mi ,
ni sh ed exp en dit u re o f m a tt er or m oti on or b oth i n ra c e

, , ,
p res er v a ti on .
I n b ri ef th en ex a m i na ti on o f the evid en ce sh ows th a t

, ,
th ere does exi s t th a t rel a ti on whi c h we i nferred m us t exi s t .
A r gu i ng f ro m genera l d a t a we s a w th a t for th e m a i nt ena n ce

,
o f a s p eci es th e a bi li ty t o p r o d u c e of
,
fs p ri ng m u s t be grea t ,
i n p rop orti on a s th e a bili ty of th e i ndividu a l s t o c ont en d w ith

d es t royi ng f orc es i s sm a ll ; a n d conv ersely A rg ui ng f rom .
oth er g en er a l d a t a w e s a w th a t d er iv ed as the s elf s u s t a in

, ,
-
i ng a nd ra ce sus t a i ni ng f orc es a re f rom a c om m on s t ock o f

-
f orce it nec es s a r il y h a ppen s th a t oth er thi ngs equ a l i ncrea se

, , ,
of one i nv olv es d ec rea s e of the oth er A nd th en t urni ng.

,
IN T E RP RE TA T I ON A ND ! U A L IF ICA TION .
49 9
to speci a l f a cts we h a ve f ound th a t thi s i nv ers e v a ri a ti on i s

,
cl ea rly t ra c ea bl e th rou gh ou t b oth the a nim a l a nd v eget a l

k i ng d om s . We m a y therefore s et it d own a s a l a w th a t ,
ev ery hi gh er d egree o f or ga n ic ev ol u ti on h a s for its c onc om i

,
t a nt a l ower d egree of th a t p ecu li a r or ga n ic diss olu ti on whi ch

is s een i n th e p r od u cti on of new org a ni s ms .
§
3 63 . S om ethi n g m a ins
ret o be s a id in reply t o t h e i n
qui ry— h ow i s the ra ti o b et ween I ndividu a ti on a nd Genes is

es t a blish ed i n ea c h c a s e ? T his i n qu i ry h a s b een bu t p a r ti a lly
.
a ns were d i n the c ou rs e of th e f oreg o i ng a rg u m en t .
M a ny s p eci a liti es of the rep rod u ct iv e p roces s a re m a n i

fes tl y du e t o th e na t u ra l s el ecti on of f a v ou r a bl e v a r i a ti ons .
Wh eth er a crea tu re l a y s a few l a rge eggs or m an y sm a ll ones

eq u a l i n w e ight t o th e few l a r g e i s n ot d et er mi ned by a ny
,
phys i ol ogic a l nec ess ity : h ere th e only a ss igna bl e c a u s e is th e

s u rviv a l o f v a r i eti es i n whi c h th e m a tt e r d ev o t e d t o rep r o
du cti on h a pp ens t o be divid ed i nt o p or ti on s of su ch S i ze a nd

num b er a s m o st t o f a v ou r m u ltipli c a ti on Wh ether i n a ny
.
c a s e th ere a r e fre qu en t s m a ll b r oo ds or l a rg er b r ood s a t
l onger in t er v a l s d ep end s wh olly on th e c ons tit u ti ona l p ecu

,
l i a ri ty th a t h a s a ri sen fro m th e dyi n g ou t of fa mili es i n

whi ch th e s i zes a nd i nt erv a l s of th e b rood s were l ea st s u it ed
t o th e c on diti on s of lif e Wh eth er a sp eci es of a nim a l pro
.
d u c es m a ny off s p r i ng o f whi ch it t a k es n o c a re or a few of

whi ch it t a k es m u ch ca re— th a t i s wh eth er its r ep rodu ctive
,
s u rpl u s i s l a id ou t wh olly i n germs or p a rtly i n ger m s a nd

p a r tly i n l a b ou r on thei r b eh a l f— m u s t h a v e b een d ecid ed by
th a t m ou ldi ng of c on stit u ti on t o conditi ons sl owly effect ed
th rou gh the m ore f requ ent p res er va ti on of d es c end a nt s f rom
th os e wh os e rep rodu ctiv e h a bit s were b es t a d a pt ed t o th e
ci rcu m st a nce s of the S p eci es Giv en a c ert a i n s u rpl u s a v a il
.
a bl e for ra c e p r es erv a ti on
- a nd it i s cl ea r th a t by i n di rect
,
e qui lib ra ti on only ca n th er e be es t a bli s h ed th e m or e or

,
l es s p ecul i ar dist ribu ti on of this s u rplu s which we s ee i n

ea ch c a s e . Obvi ou sly t oo s u rviv a l of the fitt est
, ,
5 00 L A WS OF MU L TI P L ICA TI ON .
h as ash a re i n d et ermi ni ng the p rop orti on between the

a m ou nt Of m a tt er th a t g oes t o I ndivid u a ti on a n d the a m ou n t
th a t g oes t o Genes i s Wh eth er the i nt erests of the s peci es

.
a r e m o s t s u b s erv ed by a high er ev o l u ti on o f t h e i ndi vid u a l
j oi ned with a dimi ni s h ed fertili ty or by a l ower ev olu ti on of

,
the i n dividu a l j oi ned with a n i nc rea s ed f ertility a re qu es ,
ti on s ev er b ei ng exp erim ent a lly a ns wered I f the m ore .
d ev el op ed a nd less prol i fic v a ri ety ha s a grea t er nu mber of

-
s u rvi v ors it bec om es e s t a bli s h ed a nd p red o mi na n t

,
I f con .
,
tra ri wi s e the c on diti ons of life b ei n g S impl e the l a r ger or

, ,
m ore org a ni zed i ndividu a ls g a i n nothin g by thei r grea t er siz e

-
or be tt er org a ni za ti on ; th en th e grea t er f ertility o f th e l ess
ev olv ed o nes wi ll i ns ur e t o th ei r d escen d a nts a n i nc rea si ng

,
p red om i nan ce .
Bu t di rec t equ ilib ra ti on a l l a l on g m a i nt a ins the limits

withi n which i nd irect eq u ilib ra ti on thu s w ork s Th e .
nece ss a ry a n t a g onism w e h a v e t ra c e d rigidly res t ri c ts t he,
ch a nges th a t na t u ra l s el ecti on c a n p rodu c e u nd er given c on ,
di ti on s i n eith er di rec ti on A grea t er d em a n d for I ndivid ua

, .
ti on be it a d em a nd c a u sed by s om e s p ont a neou s v a ri a ti on or

,
by a n a d a ptive i nc rea s e of st ruct u re a nd fu ncti on i nevit a bly ,
dimi n ishes the s u pply for Genes i s ; a nd na t u ra l s el ecti on

c a nnot oth er thi ng s re m a in ing the s a m e res t ore the ra t e of
, ,
Genes is whil e th e hi gh er I ndivid u a ti on i s m a i nt a in ed C on .
versel y s u r viv a l of th e fitt es t a cti ng on a sp eci es th a t h a s

, , ,
by S p ont a neou s v a ri a ti on or oth erwi s e bec om e m ore p rolific, ,
c a nn ot a g a i n r a is e its l owered I n dividu a ti on S O l ong a s ev ery ,
thin g els e c onti nues c onst a nt .
§ 3 6 4 H ere h owever a qu a lific a ti on mu st be m a d e I t

.
, , .
wa s p arenth etic a lly rem a rk ed i n 3 2 7 th a t th e i nv ers e v a ri a

,
ti on betw een I ndi vidu a ti on a nd Genesis is not ex a ct ; a nd it

wa s hint ed th a t a Sli ght m odific a ti on of st a t em ent w ou ld be
req u i s it e a t a m ore a dv a nc ed st a g e o f the a r gu m en t We .
h a v e now rea ch ed the p rop er pl a c e for S pecifying thi s m odi

fi ca ti on .
I NTE RP RE T A TI ON A ND ! U A L I FI CA T I ON .
5 01
E a ch i ncrem ent of evol uti on ent a ils a d ec rem en t of rep ro

d u cti on whi c h i s not a ccura t ely p r op or ti ona t e bu t s om ewh a t ,
l es s t h a n p rop orti on a t e T he ga i n i n the one di recti on is not

.
wh olly c a n c ell ed by a l oss i n the other di recti on bu t only ,
p a rti a lly c a n c ell ed : l ea vi ng a m a rgi n of p rofit to the s pe ci es .
T h ou gh a u g m ent ed p ower of s elf m a i nt ena nce h a bit u a lly -
n ec ess it a t es dimi ni sh ed p ower o f ra c e p rop a g a ti on yet the -

,
p rod u ct of the two f a ct or s i s grea t er th a n be f ore ; s o th a t the

f orces p res erv a tiv e of ra c e b ec om e th erea ft er i n exc ess of the
, ,
f orc es d es t ru ctiv e of ra ce a nd the ra c e s p rea ds We S h a ll

, .
s oon s ee why this h a pp ens .
E v ery a dv a nc e i n ev ol u ti on impli es a n ec on omy T h a t a ny .
i nc rea s e i n b ul k or st ru ctu re or a ctivity m a y bec ome es t a b

, , ,
l i sh ed the life of t he org a nism m u s t be t o s om e ext ent

,
f a cilit a t ed by the ch ange— th e c ost of s elf s u pp ort mu st be -

,
o n th e a v era g e re d u c ed ,
I f the g rea t er c ompl exity or the
.
,
l a rger S i ze or the m ore a gil e m ov ement ent a il s on the i n

, .
dividu a l a n ou tl a y th a t i s not rep a id i n f ood m ore ea sily -
o bt a i ne d or d a ng er m ore e a sily esc a p ed ; th en the i ndivid u a l

,
-
will be a t a rel a tive di s a dv a nt a ge a nd i ts dimi nish ed p ost erity

,
wi ll di s a pp ea r I f the ext ra ou tl a y is but j u s t m a d e g ood

.
by th e ext ra a dv ant a ge the m odifie d i ndividu a l will n ot su r

,
vive l onger or l ea v e m ore d esc end a nt s th a n the u nm odifi ed

, ,
i n dividu a ls Cons equ ently it is only wh en th e exp ens e of

.
,
grea t er i ndividu a ti on is ou t b a l a nced by a s u b s equ ent s a vi ng

-
,
th a t it ca n t end t o s u b serve the p res erv a ti on of th e i ndi

vi d ua l a nd by implic a ti on the p res erv a ti on of the ra c e
, , , .
T h e vit a l c a pit a l i n v est ed i n the a lt era ti on m u st b ri n g a

m ore th a n equ iv a l ent ret u rn A few i ns t a n c es
.
will S h ow th a t wh eth er the ch a n ge res u lt s f rom di rect

,
equ ilib ra ti on or f rom i n di rect equ ilib ra ti on thi s m u st h a pp en , .
S u pp os e a c rea t u re t a k es t o p erf orm i ng s om e a ct i n a n u n

u s u a l w a y— l ea p s wh ere o r di na r ily its ki n d red c ra wl el u d es ,
pu rsu it by divi ng i ns t ea d of like oth ers of its kind by s wim

, ,
mi ng a l ong the s u r fa c e es c a p es by d ou bli ng i ns t ea d of bv

,
S p ee d
. C l ea rly p ers ev era nc e in the m odifi ed h a bit will oth er
, ,
5 02 L A WS OF MUL TI P L I CA TION .
’
thi ngs equ a l imply th a t it t a k es l ess eff ort T he c rea t ure s
,
.
s ens a ti on s will ev er p r o mpt d es i s t a n ce fr om th e m ore l a b or i
ou s c ou rs e ; a nd h ence a c on genit a l h a bit is n ot lik ely t o be
diverg ed f rom u nl ess a n ec onomy of for ce is a chi ev ed by the

div erg enc e A ss u min g then th a t th e new m eth od h a s n o
.
, ,
a dv a n t a g e ov er t he ol d in di rec t ly dimi ni s hin g t h e ch a nces
of d ea t h ,th e es t a bli s h m en t of it a nd of th e s t ru ct u r a l
,
c ompli ca ti ons i nv olved nev erth el ess impli es a phys i ol ogic a l

,
gai n . S u pp os e a g a i n th a t a n a nim a l t a k es t o s om e
, ,
a b u nd a nt f ood p revi ou s ly r e fu s ed by it s ki n d I t i s l i k ely t o .
p ers i s t onl y if the c omp a ra tiv e ea s e i n obt a i ni n g thi s f ood ,
m ore th a n c omp ens a t es for any wa nt of a d a pt a ti on t o i t s

diges tiv e orga n s ; s o th a t s u perp os ed m odifi c a ti ons of the
'
diges tiv e org a ns a re lik ely t o a ri se only wh en a n a v era g e

ec on omy res u lts . Wh a t now m u s t be th e i nfl u ence
’
on th e c rea t u re s sy s t em a s a w h ol e ? D imi n i s h ed exp endit u re
i n a ny di recti on or i n crea s ed nu t riti on h owev er effect ed
, ,
will l ea v e a grea ter s u rpl u s of m at eri a ls T he a ni m a l w ill be .
phy s i ol ogic a l rich er P a rt of its a u gm ent ed w ea lth will g o

.
t ow a r d s its own g rea ter i ndividu a ti on— its s i ze or its ,
s t ren gth , or b oth will i nc rea s e ; whil e a n oth er p a rt will g o

,
t owa rds m ore a ctiv e g enes i s Ju st a s a st a t e of pl eth ora

.
di rectly p rodu ced enh an c es f ertility ; s o will s u ch a st a t e

i ndi rectly p roduc ed .
I n a noth er wa y the s a m e thi ng mu s t res u lt f rom th os e

,
a dditi on s t o b u lk or c om pl exity or a ctivity th a t a re du e t o
s u rviv a l of the fitt es t Any ch a nge whi ch p rol ong s i ndivid u a l

.
li fe will oth er thi n gs rem a i ni ng the s a m e f urth er th e pr o

, ,
d u cti on of offs p ri ng E v en wh en it i s not lik e th e f oreg oi ng

.
, ,
a m ea n s o f ec onomi z i ng the f orc es of the i n divid u a l still if it , ,
i nc rea ses th e ch a nces of es ca pi ng d es t ru cti on it i nc rea se s the ,
ch a nc es of l ea vi ng p os t erity A ny furth er d egree of evol u

.
ti on th erefore will be es t a bli s h ed on ly wh ere the cost of

, ,
it is m ore th a n r ep a id : p a rt o f the g a i n b ei ng s h own i n the

l ength ened li fe of the i ndividu a l a nd p a rt i n the grea t er ,
p rod u cti on o f oth er i n divid u a l s .

INT E RP RE T A T ION A ND ! UA L I FICA TI ON .
5 03
We h a ve h ere the
s ol uti on of v a ri ou s mi nor a nom a li es by
whi ch the i nv erse v a ri a ti on of I ndivid u a ti on a nd Genes i s is
o b s c u red .T a k e a s a n i nst a nc e the f ertility of the B l a c kbi r d
a s c omp a r e d with th a t of th e L i nnet B oth bi r ds l a y fi ve eggs
.
,
a n d b oth u s u a lly h a v e two b ro ods Y et the B l a ckbi r d i s fa r

.
t he l a rger of the two a nd ou ght a c c or di n g t o th e g enera l

, ,
l a w t o be mu ch l es s p rolific
, Wh a t c au s es this noncon
.
f or mity ? We S h a ll fi nd a n a ns wer i n th ei r res p ective f oods

a n d h a bit s E x cept d u ri ng the ti m e th a t it i s rea ri ng its
.
you ng th e L i nnet c oll ect s only v eget a l f ood— liv es du r i ng

,
t he w i nt er on th e s ee d s it fi nds i n th e fi eld s or wh en h a r d , ,
p res s ed pi ck s u p a roun d f a r m s ; a nd t o obt a i n thi s S p a re

,
di et i s conti nu a lly flyi ng a b ou t T he resu l t if i t s u rviv es th e

.
,
f ros t a nd s now i s a cqns i dera bl e d epl eti on ; a nd it rec ov er s

,
i t s c onditi on only a ft er s om e l ength of s p r i ng wea th er Th e .
B l a c kbi rd on the oth er h a nd i s o mniv orous

, , Whil e it ea ts .
g ra i n a nd f ru it wh en th ey com e i n its way it d ep ends l a rgely ,
on a ni m a l f ood I t cu t s t o pi eces a n d d ev ou rs th e dew w orms

.
-
which m orn i n g a nd eveni ng it fi nds on the s u r f a ce of a l a wn

, , ,
a nd ,
e v en di s c ov eri ng w h ere th ey a re u n ea rth s th em ; it ,
sw a ll ow s S l u g s a n d b rea kin g s na il s h ell s eith er with its b ea k

,
-
,
or byh a mm eri n g th e m a g a i ns t s t ones t ea rs ou t th ei r t ena n t s ;

,
'
a n d it ea ts b eetl e s a nd l a rv ae T hu s th e s t rength of the

.
B l a ckbi r d Op en s t o it a st ore of g ood f ood mu ch of whi ch is ,
i na c ces s ibl e t o s o sm a ll a nd wea k a bi r d a s a L i nnet— a st ore

es p ec i a lly h elpfu l t o it d u r i n g t he c old m on th s when th e ,
hybe rn a t i ng s na ils i n h edge b ott om s yi eld it a b u n d a nt pr o

-
vi s i on The resu lt i s th a t th e B l a ckbi r d is rea dy t o b ree d

.
v ery ea rly in S p ri ng a nd is a bl e du ri ng the su mm er to rea r

,
a s ec ond a n d s om etim e s e v en a thi rd b r oo d

,
H ere th en a , .
, ,
high er d egree of I ndivid u a ti on s ecu res a dv a nt a ges S O g rea t ,
a s t o m u ch m ore th a n co mp en s a t e i t s c os t I t i s n ot th a t th e .
d ecli ne of Genes i s i s l es s th a n p r op orti ona t e to the i nc rea s e of

I ndi vidu a ti on bu t th ere i s n o d ecli ne a t a l l
,
C om
p a ri s on of the Ra t with the Mou s e yi eld s a p a ra ll el resu lt .
T h ou gh t h ey differ g rea tly i n S ize yet the one is a s p rolific,

5 04: L A WS OF -
'
MUL TI P L I CA T I ON .
as th e th er T hi s a b s ence of di fferenc e c annot be a scribe d

o .
t o th ei r u nlik e d egrees of a ctivity We mu s t s eek i ts c a u s e .
i n s om e f a cili ty of li vin g s ec u re d t o th e Ra t by its grea t er

i nt elligenc e gr ea t er p ower a nd c ou ra g e grea t er a bility t o
, ,
u tili ze wh a t it fi n ds T he Ra t i s not ori ou sl y cu nni ng ; a nd i ts

.
c u nning giv es s u cces s t o its f ora gi ng exp editi ons I t is n ot .

,
lik e the Mou s e limit ed m a i nly t o v eget a l f ood ; bu t whi l e it

,
ea t s g ra i n a nd be a n s lik e th e Mou s e it a l s o ea ts fl es h a n d ,
c a rri on d ev ou r s y ou ng p oul t ry a nd eggs T he resu lt i s th a t

,
.
,
with ou t a p rop or ti ona t e i n crea s e of exp endi t u re it gets a far ,
l a rger s u pply of nou ri s hm ent th a n the Mou s e ; a nd rel a tiv e

e xc es s of nou ri s h m ent m a k es p ossibl e a l a rg er S i ze wi th ou t
a s m a ll er r a t e of m u ltipli ca ti on H ow cl early thi s i s th e .
ca u s e w e see i n th e c ont r a s t b et ween th e c omm on Ra t a n d

,
th e Wa t er Ra t Whil e the c omm on Ra t h a s ordi na rily

-
.
s evera l b rood s a y ea r of f ro m 1 0 t o 1 2 ea ch the Wa t er Ra t

-
,
-
,
th ou gh s om ewh a t s ma ll er h a s bu t 5 or 6 i n a b rood a nd bu t
, ,
B u t th e Wa t er
'
one b rood or som et i m es t wo b r oods a y ea r

, ,
-
.
Ra t liv es on v eg et a l f oo d a nd it l a cks a l l th a t its b old s a g a

, ,
ci ou s om n iv orou s c on g ener g a i ns f rom th e w a r mth a s w ell a s

,
’
th e a bu nd a nc e which m en s h a bit a ti on s yi eld .
T he i nv ers e v a ri a ti on of I ndivi d u a t i on a nd Genes is is ,
th eref ore bu t a pp roxim a t e

, Rec ogn iz i ng th e t ru th th a t .
e v ery i n crem en t of ev o l u ti on which is a pp rop ri a t e t o t h e
ci rcu m s t a nces of a n or ga n ism b ri ng s a n a dv ant a ge s om ewh a t ,
i n exc es s of its c ost ; we s ee the genera l l a w a s m ore st rictly ,
s t a t ed t o be th a t Genesis d ec rea s es not quit e S O f a st a s

,
I ndivid u a ti on i nc rea s es Wh eth er the gr ea t er I ndivid u a ti on

.
t a k es the f orm of a l a rger b u lk a nd a cc omp a nyi ng a ccess of

st rength ; wh eth er it be sh own in high er s p ee d or a gility ;
wh et h er it c onsi s t s in a m odific a ti on of s t ru ct u re which fa cili
t a t es s om e h a bit u a l m ov em ent or i n a vis cera l ch a nge th a t ,
h elps t o u tili ze be tt er the a b s orbe d a li m ent ; the u ltim a t e

eff ec t is id entic a l T h ere i s eith er a m ore ec onomic a l per
.
form an ce of the s a m e a cti ons i nt erna l or ext erna l or th ere , ,
is a s ec u ri ng of g rea t er a dva nt a ges by m odifi ed a cti on s which ,

INT E RP RE TA TI ON A ND ! U A L I FI CA TI ON .
5 05
c os t no m ore or h a v e a n i ncrea s ed c ost l ess th an the i n

,
c rea s ed g a i n I n a ny c a s e the resu lt i s a g rea t er s u rplu s of

.
vit a l c a pit a l p a rt of which g oes to the a ggra ndi zem ent of

,
th e i ndi vidu a l a nd p a rt t o the f or m a ti on of n ew i ndi vid u a l s

,
.
Whil e the high er tid e of nu t ritiv e m a tt ers ev erywh ere filli n g

,
t he p a rent orga nis m a dds t o its p ower of s elf m a i nt ena nc e i t

-
,
-
,
a l s o c a u s es a rep r od u ctiv e o v er fl o w l a rg er th a n b ef ore .
H enc e ev ery typ e whi c h is be st a d a pt ed t o its c onditi ons ,
( a nd this on the a v era ge m ea ns ev ery higher typ e ) h a s a ra t e

,
o f m u ltiplic a ti on th a t ins u re s a t en d ency t o p red o mi na t e .
S u rviv a l of th e fitt es t a cti n g a l one is ev er repl a c i ng i n

, ,
feri or sp eci es by s u p eri or sp eci e s . B ut b ey on d the l onger

s u rviv a l a nd theref ore grea t er c h a nc e of l ea vi n g off s p ri ng
, ,
whi ch s u p eri ority giv es we see h ere a noth er wa y i n which

,
the S p rea d of the s u p eri or is i nsu red . T h ou gh th e m ore

e v olv ed o rg a ni sm is the l ess f ertil e a bs ol u t ely, it is the m ore
f ert il e rel a tiv ely .

C H A P T E R XI I .
M U L T I P L I CA T IO N OF TH E H UM AN RA CE .
§ 3 65 . l tiv
TH E e f r tility
re a
e o f M a n c on s id ere d a s a
S p ec i es a n d th os e ch a n ges i n Ma n s f ert ility w hich occ u r u nd er

’
,
ch a nged c onditi on s mu s t c onform t o the l a ws whi ch we h a v e

,
t ra ce d thu s fa r A s a m a tt er of c ou rs e th e i nv ers e v a ri a ti on
.
,
b et ween I ndivid u a ti on a nd Genes i s h old s of him a s of a l l

o th er org a n i ze d b ei ngs H i s ext rem ely l ow ra t e o f m u ltipli
.
c a ti on— fa r b el ow th a t of a l l t erres t ri a l Ma m m a l s except the

E l eph an t ( whi c h th ou gh oth erwi s e l es s e v olv ed is i n ext ent
, ,
of i n t eg ra ti on m ore ev olv ed ) — we Sh a ll rec ogni ze a s the

,
n ecess a ry c onc o mit a n t of h i s m u ch high er ev ol u ti on A nd .
the c a u s es of i nc rea s e or d ec rea s e i n hi s fertility sp eci a l or ,
genera l t emp ora ry or p erm a nent we S h a ll exp ect t o fi nd i n

, ,
th os e ch a nges of b u lk of s t ru ct u re or of exp endit u re which

, . ,
we h a v e i n a l l oth er ca ses s een a ss oci a t ed wi th s u ch eff ect s .
I n the a b s ence of d et a il ed p roof th a t th ese p a r a ll el i s ms

e xist it might s u f
, fi c e t o con t empl a t e the s ev era l c omm u n iti e s
b et ween the rep rod u ctiv e fu ncti on i n h u m a n b ei ngs a nd other
b ei ngs I d o not refer S imply t o th e f a ct th a t genes is pro
.
ceeds i n a S i m il a r m a nner ; bu t I r efer t o t he si m il a r ity of
t h e rel a ti on b et ween th e genera tiv e fun cti on a n d th e fu nc

ti ons whi ch h a v e for th ei r j oi nt end the p reserv a ti on of the
i ndivid u a l I n Ma n a s i n oth er c rea tu res th a t exp end mu ch
.
, ,
g enes i s c omm enc es on ly wh en gr owt h a n d d ev el opm ent a re

d ecli ni ng i n ra pidity a nd a pp roa chi ng thei r t ermi na ti on .
A m ong the high er org anisms i n g enera l the rep rod u ctiv e ,
5 06
MUL T I P L ICA TI ON OF T HE H U MA N RA CE .
5 07
a ctivity c onti nu i ng d u ri ng the p rim e of life c ea s es wh en the

, ,
vig ou r d eclines l ea vi n g a cl os i ng p eri od of i nfertility ; a nd i n

,
lik e m a nner a m ong ou rselves b a rrenness s u p er v enes wh en ,
middl e a ge b ri ng s the s u rpl u s vit a lity t o a n end S o t oo .

, ,
it is f ou n d th a t i n Ma n a s i n bei ng s of l ower or d er s th ere i s

, ,
a p eri o d a t w hi c h f ecu ndity cu lmin a t e s I n 3 4 1 fa c t s were .

,
c it ed s h ow i n g th a t a t th e c om m en cem en t of th e rep rod u ctiv e
p eri od a ni m a l s b ea r f ewer ofl s pri ng th a n a ft erw a r ds ; a nd

,
th a t t ow a rd s the cl os e of t he rep r od u ctiv e p eri od th ere i s a ,
d ecrea s e i n the nu mb er p rodu c ed I n lik e m a nner it is sh own .
by the t a bl es of D r D u nca n s recent w ork th a t the fec un dity

.
’
,
o f w om en inc rea s es u t o t h e a ge of a b ou t 2 5 y ea r s a nd
p ,
c o nti nu i n g high with bu t S light dimi nu ti on till a ft er 3 0 ,
th en g ra du a lly w a nes I t is the s a m e w ith the s i zes a nd

.
w eight s of off s p r i n g I n fa n t s b orn of w om en from 2 5 t o 2 9

.
y ea r s of a g e a re b oth l ong er a n d h ea vi er th a n i nfa nts b orn

,
o f y ou ng er or Old er w om en ; a n d this di fference h a s the s a m e

i m plic a ti on a s th e grea t er t ot a l weight of the offs p r i ng pro
du ced a t a bi rth d u r i ng th e m ost fecu n d a ge of a pl u r ip a r ou s
,
a n im a l . Onc e m ore th ere i s th e fa ct th a t a t o o ea rly b ea ri ng

,
-
o f y ou n g p r o d u c es on a w o m a n th e s a m e inj u r i ou s ef fec t s a s
on a n i n f eri or crea tu re a n a rres t of growth a nd a n en feebl e
g
m ent of cons tit u ti on .
C ons id eri ng th es e genera l a n d S pe ci a l p a ra ll elisms we ,
might s a fely i nfer th a t v a ri a ti ons of hu m a n fertility c onfor m

t o the s a m e l a ws a s d o v a ri a ti on s of f ertility in genera l .
B u t it is n ot n ee dfu l t o c on t ent ou rs elv es w ith a n impli c a ti on .
E vid ence i s a s sign a bl e th a t wh a t c a u ses i ncrea s e or d ecrea s e

o f g enes i s i n o th er c r ea t u r es c a u s es i nc rea s e or d ecrea s e of
,
genes i s i n Ma n I t i s t ru e th a t ev en m ore th a n hith ert o ou r

.
, ,
rea s on i n gs a re b eset by diffi cu lti es S o nu m erou s a re t he .
i nequ a liti es i n th e conditi on s th a t bu t few u n obj ecti on a bl e

,
co mp a r i son s ca n be m a d e T he h u m a n ra c e s diff er c on sid er

.
a bly i n th ei r s i ze s a n d n ot a bly i n th ei r d egr ee s of c ereb r a l

,
d evel op m ent T he c ou nt ri es th ey i nh a bit ent a il on th em

.
wid ely different c ons u mpti ons o f m a tt er for m a i nt ena nce of

5 08 L A WS OF MUL TI P L I CA T ION .
t emp era t ure B oth i n th ei r qu a liti es a nd qu a ntiti es the

.
f ood s they live on a re u nlik e ; a nd the su pply is here regul a r

a nd th ere v ery i rr egu l a r T h ei r exp en dit u res in b odily a cti on
.
a r e e xt rem ely u n equ a l ; a n d ev en stil l m ore u nequ a l a re

th ei r e xp en ditu res i n m ent a l a cti on H en c e the f a ct ors .
,
v a ryi ng s o mu ch i n thei r a m ou nts a nd c ombi na ti ons ca n ,
s c a rcely ev er h a v e th e i r r es p ec tiv e efi ect s id en tifi e d N ev er .
thel es s th ere a re a few c omp a ris ons the res u lts of whi ch m a y
with s t and c ritici s m .
3 66 . i ncrea s e of fertility c aus ed by a nu t riti on th a t

The
is grea tly i n ex ces s of the exp en dit u re is t o be d et ec t ed by ,
c ont ra sting p opu l a ti ons of the s a m e r a ce or a lli ed ra c es , ,
one o f which obt a i ns g ood a n d a b u n d a nt s u st en a nc e m u c h
m ore ea sily th a n the oth er T h ree c a s es m a y h ere be set

.
d own .
T he t ra v ell er B a rrow d es c ribing the C a p e B oers s a ys

,
-
,
” “
Unwilli ng t o w ork a n d u na bl e t o thi nk i ndu lgi ng ,
t o exc ess i n the gra tific a ti on of e v ery s ens u a l a pp etit e the ,
A f ric a n p ea s a nt g rows t o a n u n wi eldy s i ze ; a nd r es p ecti ng
t he oth er s ex h e a dd s ,
th e w om en o f th e A f rica n p ea s a nt ry
”
l ea d a life of th e m ost li s tl es s in a ctivity T h en a ft er ill u s .
,
t ra ti ng th es e s t a t em ents he g oes on t o n ot e
“
, the p rolific
t end en cy of a l l the A f ric an p ea sa nt ry S i x or s ev en child ren
.
i n a f a mily a re c ons id ered a s v ery few ; f rom a d oz en to

tw enty a re not u nc om m on . T h e na tiv e ra c es of
thi s regi on yi eld evid enc e t o the s a m e effect S p ea king of .
th e c ru elly u s ed H ott ent ot s ( h e i s w riti ng a c ent u ry a g o )

-
,
wh o whil e th ey a re p oor a nd ill fed h a v e t o do a l l t he w or k

,
-
,
for th e idl e Boer s B a r row s a ys th a t th ey s eld om h a v e m ore

,
th a n two or th ree child ren ; a nd m a ny of the wom en a re

”
b a rren T hi s u nu su a l i nfertility st a n ds i n rem a rk a bl e con
.
t ra st with the unu s u a l fertility of the K a ffi rs of wh om he ,
a ft erw a r ds giv e s a n a cc oun t Rich i n c a ttl e l ea di ng ea sy

.
,
lives a nd livi ng a lm os t excl u siv ely on a nim a l f o od ( chi efly

,
milk with oc ca s i ona l fl es h ) thes e p eopl e were th en repu t ed

,
MU L T I P L ICA TI ON OF T HE H U MA N RA CE .
5 09
t o ha v e v ery high ra t e o f m ultiplic a ti on B a rrow w rit es

a .
T h ey a re s a id t o be exc ee di ngly p r olific ; th a t twi n s a re

“
a l m os t a s f req u en t a s si ngl e bi rths a nd th a t it is n o u n ,

”
c om m on thi ng for a w om an t o h a v e th ree a t a ti m e P ro .
b a bly b oth th es e st a t em ents a re in exces s of the t ru th ; but

th ere is room for l arge di s c ou n ts wi th ou t d es t royi ng the
ext re m e differenc e A thi rd i ns t a nce i s th a t of th e
.
F rench Ca n a di a n s N ous s om m es terri bl es p ou r l es en

.
f a n ts ! obs erv e d one of th em t o P r of J oh n st on wh o t ell s u s .

,
th a t the m a n wh o s a id thi s wa s one of f ou rt een child ren

wa s hims elf t he fa th er of f ou rt ee n a n d a ssu red m e th a t f ro m ,
’
eight t o s i xt een w a s the u s u a l nu m b er of t he f arm ers
f a mi l i es . H e ev en n a m ed one or two w o m en wh o h a d
b rou ght th ei r hu s b a nds fi ve a nd t wenty a nd th rea t ened Z o - -
,
vi ngt si xiém e p ou r l s
-
F ro m th es e l a rge f a mili es ,
j oin ed with the ea rly m a rri a ges a nd l ow ra t e of m ort a lity it ,
res u lts th a t by na t ura l i n c rea s e

,
th ere a re a dd e d to the ,
F rench Ca na di a n p op u l a ti on of L ower Ca na d a fou r p ers ons

-
”
for ev ery one th a t is a dd ed t o th e p op u l a ti on of E ngl a nd .
N ow th es e F rench Ca na di a n s a re d es c rib ed by P rof J ohn s t on

-
.
a s h o m e l ovi ng c ont en t ed u nent er p r i s i n g ; a nd a s livi n g i n

-
, ,
“ ”
a regi on wh ere l a nd a n d s ubsis tenc e a re ea s ily obt a i ned .
Very m od era t e i nd u s t ry b ri ngs t o th em lib era l su ppli es of

n ec es s a ri es ; a nd th ey p a s s a c on sid era bl e p orti on of the yea r
in idl enes s H enc e the c ost of I ndivid u a t i on being mu ch
. .
re d u c ed , the ra t e of Ge nes is i s mu ch i n c rea s ed T h a t this .
u n c omm on f ertility is n ot du e t o a ny di r ec t i nfl u enc e o f t h e
l oc a lity i s impli ed by the fa ct th a t a l on g w ith th e res tl ess

, ,
disc ont ent ed s t r ivi ng bu rni n g en ergy of th ei r S a x on neigh

, ,
”
b ou rs no s u ch r a t e of mul tipli ca ti on i s ob s erv ed ; whil e
,
fu rth er s ou th wh ere the physi ca l ci rcu mst an ces a re m ore

,
f a v ou ra bl e if a nythi ng th e A ngl o S a x ons l ea di ng liv e s of

,
-
,
e xc es s iv e a ctivity h a v e a f ertility b el ow the a v era ge A nd

, .
th a t the p ec u li a rity i s not a di rect effect of ra c e is p rov ed by ,
the f a ct th a t i n E u rop e the ru ra l F rench a re c ert a inly not

,
m ore p rolific th an the ru ra l E ngli sh .

51 0 L A WS OF MU L T I P L I C A T I ON .
To v y rea d er there will p rob a bly occu r the s eem i ngly

e er
a dv ers e e vid enc e fu rn i s h ed by t h e I r i s h ; wh o th ou gh not ,

.
well fed m u l tiply f a s t

,
P a rt of t h i s m ore ra pid i ncrea se is
.
due t o the ea rli er m a rri a ges c om m on a m on g th em a n d con ,
s equ ent qu ick er s u c c es s i on of g en era ti ons — a fa ct or whi ch ,
a s w e h a v e s een h a s a l a r ger effec t th a n a ny ot her on t h e

,
ra t e o f m u l tiplic a ti on P a rt of it i s du e t o the g rea t er

.
genera li ty of m a rri a ge — t o the c omp a ra tiv e sm a lln ess of the

nu mber wh o di e w ith ou t h a vi ng h a d t h e opp ort u nity o f pro
du ci ng offs p r i ng T h e e ff ect s o f th es e c au s es h a vi ng been

.
d ed u ct ed we m a y d ou bt wh eth er the I ri s h individ u a lly c on

, ,
s idered w ou ld be f ou n d m ore p r olific th a n th e E nglish

, .
P erh a p s h owev er it will be s a id th a t c ons id eri ng th ei r di et

, , , ,
th ey ou ght t o be l ess p rolific T hi s i s by no m ea ns obvi ou s . .
I t i s n ot s i m ply a qu es ti on of nu t ri m en t a b s orb ed I t i s a .
qu es ti on o f h ow m u ch rem a i ns a fter the exp endi tu re i n s elf

m a i n t en a n c e N ow a not or i ou s p ecu li a r ity in the life of the
.
I ri s h p ea s a nt i s th a t h e obt a i n s a ret u rn o f foo d w hich i s

,
l a rge i n p r op orti on t o hi s ou tl a y i n l a b ou r T he cu ltiv a ti on .
of h i s p ot a t oe g r ou n d o c c u pi es ea c h c ott a g er bu t a s m a ll p a r t
-
of the y ea r ; a n d the d o m es tic econ o m y o f his w if e is n ot of a
ki nd t o ent a il on her m u c h d a ily ex erti on C ons equ ently the .
c rop t ol era bly a b u nd a nt i n qu a ntity thou gh i nnu tri ti ve i n

,
q u a lity p os s ibly s u ffi ce s t o m ee t th e c omp a ra tiv ely l ow ex

,
-
en d i tu re a nd t o l ea v e a g ood s u rpl us for g enes i s— p erh a p s

p ,
a gr ea t er s u rpl u s th a n r em a i ns t o th e m a l es a n d f em a l es o f
the E n gli s h p ea s a nt ry wh o th ou gh fed on b ett er f ood a re , , ,
/
h a r d er w ork e d .
We c oncl u d e th en th a t i n the hu m a n ra ce a s i n a l l oth er

, , ,
ra c es ,s u ch a b s ol u t e or rel a tive a bu n d a nc e of nu t rim ent a s

l ea v es a l a rge exc ess a ft er d efra yi n g the c ost of c a rryi ng on
p a rent a l life is a cc omp a ni ed by a high ra t e of genes i s *
, .
§ 3 67 . E vid ence of the c onv ers e t ru th th a t , re a l tiv e in
Th i s i s e x actl y th e re v erse of Mr D oub l eda y s doc tri ne ; whi ch i s tha t

.
’
th rough ou t b ot h th e a ni m a l a nd v egeta bl e k i ngd om s ,

“
v
o e r-feed i ng che ck s
i nc re ase ; w hi l st , on th e oth e r h and, a. li m i t ed or defici ent nut ri m e nt st i m u

MU L T I P L ICA T I ON OF T HE H U MA N R A C E .
51 1
c rea s e of exp endit u re l ea vi ng a di m i ni s h ed s u rplu s r edu c es

, ,
t he d eg ree o f f ertili ty i s n ot w a nti ng S om e of it h a s b een

,
.
s et d own for th e s a k e o f a n tith es i s i n th e fo r eg oi n g s ecti on .
H ere m a y be g r ou p ed a few fa c ts of a m ore s p eci a l ki nd

h a vi ng the s a m e i m pli ca ti on .
T o p r ov e th a t m u c h b odily l a b ou r rend er s w om en l es s p ro
l i fi c requ i res m ore e vid en ce th a n h a s a t p resent b een coll ect ed
, .
Neve rth el es s it m a y be n ot ed th a t D e B ois m ont i n F ra n c e a n d

D r Szu ki t s i n A u st ri a h a v e s h own by ext en siv e st a ti s tic a l
.
,
c om p a r i s on s th a t the rep rod u c tive a ge i s rea ch ed a yea r

,
l a t er by w om en of the l a b ou ri ng c l a s s th a n by middl e cl a s s -
w om en ; a nd whil e a s c ribi n g thi s d el a y i n p a rt t o i nf er i or

”
l at os a nd adds to i t . Or, as he e l s e wh ere s a ys Be th e ra ng e of th e
na t u r a lp owe r to i ncrea s e i n a ny sp eci es wh a t i t m ay, t h e p l ethori c s ta te i n a v .
r i a bl y ch e ck s i t , a nd t h e dep l ethori c s ta t e i n ari a bl v

y de el op s i t a nd th i s ha p v
x
p en s l n th e e a ct ra t i o of th e in t e nsi ty a nd com pl et e nes s of e a ch s ta t e, u nti l
e ach s tat e be c a rri e d s o fa r as to b ri ng a b out t he a ct u a l dea th of th e a ni m a l
”
or p l a nt i t se l f .
I ha v e s p a ce here onl y to i ndi ca t e th e m i s i nterpr et ati ons on whi ch Mr .
Do ubl ed a y h a s b a s ed h i s a rgum e nt .
I n th e fi rs t p l a ce , b e h a s co nfounded norm a l pl eth ora with wha t I ha v e, i n

3 5 5 , di s ti ngui sh ed as a bnorm a l pl eth ora Th e cas es of infert i li ty a ec om
.
p a nyi ng fa tne s s , wh i ch h e c i t es i n p roof th a t o v e r-fe e di ng c he ck s i nc re as e , a re
not ca se s o f high nu tri ti on p rop e rl y s o ca l l e d ; but ca s e s of s uch defe cti ve ah
s orp t i on or as s i m i l ati on a s cons titu t e s l ow nutri ti on. In Cha p I X, . a bun da nt
p roof wa s gi v en th a t a t ru l y pl eth ori c st a te i s a n unu s ua ll y ferti l e s ta te It .
m ay b e a d de d th a t m uch of th e e v i dence by wh i ch Mr D oubl eday seek s t o .
s h ow th a t a m ong m en, h i ghl y-fe d cl as s es a re i nfert il e cl a s ses , m ay be ou t
b a l ance d by c ount e r -e vi dence . Ma ny ye a r s ag o Mr G H L e wes p oi nt ed t hi s

. . .
out : e xtra ct ing from a book on t he p ee ra ge, th e na m es of 1 6 p eers who h a d,
a t th a t t i m e , gi v i ng a n a v e ra ge of 1 1 C) i n a fa m i l y
1 8 6 ch i l dren
°
Mr D oubl eda y i ns i st s m u ch on th e s up port gi v en to hi s th eory by th e

.
b a rrennes s of v ery l u xu ri a nt p l ant s , a nd th e frui tful ne ss p rodu ce d i n pl ant s

by de pl eti on Ha d h e b een a wa re th at th e cha nge from ba rrenn es s to fr ui t
.
ful nes s i n pl a nt s , i s a ch a nge from a gam ogenes i s to g a m ogene s i s— h a d i t b een

a s w el l k nown a t t h e t i m e wh en h e w rote a s i t i s now , t h a t a t re e whi ch
g oe s
on pu tt i ng ou t s e xl e ss sh oot s, i s th us p roduc i ng new i ndi vi du a l s ; a nd th a t
wh en i t b egi ns to b e a r frui t , i t s i m pl y b egi ns t o p r oduce n e w i ndi v idua l s a ft er
anoth er m a nner— h e woul d h a v e p ercei v e d th at fa ct s of th is cl as s do not t el l
i n hi s fa v ou r .
In the l a w wh i ch Mr . D oubl eda y a l l eges , he s ees a gu ara ntee for th e

m a i nte na nc e of s p e c i es . He a rg ue s th a t th e pl ethori c s ta t e of t he i ndi vi
51 2 L A WS OF MUL TIP L ICA T I ON .
nu t ri ti on, we m a y s u s p ect th a t it is in p a rt du e t o grea t er

m u s cu l a r exp endi t ure A ki nd red fa ct a dmi tti ng o f a
.
,
ki nd red i nt erp ret a t i on m a y be a dd ed T h ou gh the com

,
.
a r a ti vel y l ow ra t e o f i nc re a s e i n F ra n c e is a tt r ib u t ed t o
-
p
o th er c a u s es e t v ery p os s ibly one o f its c a u s es i s t he
y , , ,
grea t er p rop orti on of h a r d wor k ent a il ed on F rench wom en ,
by the exc ess iv e a b s t ra cti on of m en for n on p rod u ctiv e -
o c cu p a ti ons milit a ry a nd c ivil

,
T he high er ra t e of m u ltipli .
c a t i on i n E ngl a nd th a n i n c onti nent a l c ount ri es g enera lly is ,
not i m p r ob a bly fu r th ered by the ea s i er liv es which E ngli s h
w om en l ea d .
T h a t a bs o lu t e or rel a tiv e i nf ertility i s c omm only p rod u c ed

i n w o m en by m ent a l l a b ou r c a rri ed t o exc es s is m ore cl ea r ly ,
sh own T h ou gh the regim en o f u pp er c l a ss gi rls i s not wh a t

.
-
it sh ou ld be yet c on s id eri ng th a t th ei r f eedi ng i s b ett er th a n

, ,
th a t of gi rls b el ongi ng t o the p oorer cl a s s es whil e i n m os t , ,
o th er r esp ect s th ei r phy s ic a l t rea tm ent i s n ot w ors e the

, ,
d ua l s con s ti tu ti ng a ny race of orga ni s m s, pre su pp oses condi tions s o fa v ou r

a bl e t o l i fe tha t th e ra ce ca n be i n no da nger ; a nd th a t ra p idi ty of m ul ti
p l ica tion be com es ne edl e s s Conv e rs e l y, h e a rgue s th a t

. a de pl ethori c s ta t e
i m p l i es unfa voura bl e co ndi ti on s — i m pli es , conse q u entl y, unus ua l m orta l i ty ;

th a t is — i m pl i es a nece s s i ty for i ncreas ed fe rt il i ty to p r e ent the ra ce fro m v
dyi ng out It . m ay be rea dil y sh ow n, howe v er, th a t s uch an arra ngem ent
w oul d be the re ve rse j

of s el f-a d us t i ng . S up po se a s peci e s , too nu m erou s
for i ts food, to be i n th e r es ul ti ng depl ethori c s ta te . It wi l l , a ccordi ng to
Mr . D oubl eday, b ecom e unu s u a l l y fert i l e ; a nd th e ne t x ge nera ti on wil l be
m or e num erous ra th e r t h a n l es s num erous . For, b y th e hyp othes i s, t h e un
fert il ity due t o th e depl eth ori c s t at e,
u s ua l i s t he ca us e of undu e i ncreas e of
p opul a ti on Bu t i f th e nex t gene ra ti on i s m ore

. num e ro us wh i l e t he s uppl y
of food ha s not i ncre a s ed i n p rop ort i on, th en thi s ne x t ge nera ti on wi l l be i n

a s t i ll m ore d epl eth ori c s ta te , a nd wi l l be s till m ore fe rt il e Th u s t here wil l .
g o on an e v er i ncrea si ng ra te
~
o f m ul t i pl i ca ti on, a nd a n e e r-decrea s i n v g sh a re
of food, for e ach p ers on, unt il th e s pe c ie s di sa pp ea rs S up pose , on the .
o th er h a nd, the m em b e rs o f a s p eci es t o b e i n a n un u s ual l y pl eth ori c s t a t e .
The i r ra te o f m ul ti pli ca ti on, ordi na ril y s u f

fi ci ent to m ai nt ai n the i r num b ers,
wi l l becom e i nsu ffi ci ent to I n t he ne x t ge nera tion,
m a i nta i n th ei r nu m b ers .
th erefore, th ere w i ll b e fewer t o eat th e a l rea dy a b unda nt foo d, wh ich be

c om i ng rel a ti v el y st i l l m ore ab undant, wi l l render t h e fewe r m em b ers o f th e
s p ec i e s s t i l l m o re p l ethori c, a nd s ti ll l e s s fe rt i l e , th a n th e i r p a rent s A nd the .
a c t i ons a nd re a cti ons conti nui ng, the s peci es wi l l prese ntl
y di e out from a bs o
l ute ba rre nness .
MU L T I P L I CA TI ON OF T H E H U MA N RA CE .
51 3
d efici ency of r ep rod u ctiv e p ower a m ong th em m a y be r ea s on

a bly a t t rib u t e d t o th e o v e rt a xi n g o f th ei r b r a i n s— a n o v er
t a xi ng whi ch p rod u c es a s eri ou s r ea cti on on the phy s iqu e .
T hi s di m i nu ti on o f rep r o du ctiv e p ow er i s n ot s h own only by

t he gr ea t er f requ enc y of a b s ol u t e s t er ili ty ; n or is it s h own
o nly i n th e e a r li er ces s a ti on of chi ld b ea ri n g ; bu t it i s a l s o -
s h ow n i n t he v e ry f req u en t i n a bility o f s u c h w om en t o s u ckl e
th ei r i nf a nts I n i t s fu ll s ens e th e rep rod u ctiv e p ow er m ea n s

.
,
th e p ower t o b ea r a w ell d ev el op ed i nf a nt a nd t o s u pply th a t

-
i n fa n t with th e na t u ra l foo d for th e na t u ra l p eri od Mos t o f .
t he fl a t ch es t e d gi r l s wh o s u rviv e th ei r high p r es su r e ed u c a
- -
ti on a re i nc ompe t ent t o do thi s Were th ei r f ertility m ea

,
.
s u re d by t h e nu m b er o f c hild r en th ey cou ld rea r w ith ou t
a rtifi c i a l a i d th ey w ou ld p r ov e rel a tiv ely v er y i n fer til e

,
.
T h e cos t o f rep r o du cti on t o m a l es b ei n g s o m u ch l es s

th a n it is t o f em a l es th e a nt a g oni s m be t ween Genes i s a nd
,
I n divid u a ti on i s n ot oft en s h own i n m en by s u pp res s i on of

g enera tiv e p ower c ons equ ent on u nu su a l exp en dit u re i n
b o dily a cti on N ev erth el es s th ere a re i n dic a ti on s th a t thi s
.
,
r es u lt s i n ext rem e c a ses We rea d th a t the a n ci ent a thl etes

.
ra rely h a d c hild ren ; a n d a m on g s u c h o f th ei r m od ern repre
s ent a ti ves a s a c r ob a t s a n a lli ed r el a ti on of c au s e a nd effect

,
i s a ll eged I ndi rectly thi s t ru th or r a th er i ts c on v ers e

.
, ,
a pp ea rs t o h a v e b ee n a s c ert a i ned by th ose wh o t ra in m en
for fea t s o f s t r ength— th ey fi n d it n ee d ful t o i ns i s t on c on

'
ti nen ce .
S p ec i a lp roo f s th a t i n m en gr ea t c ere b ra l exp endit ure

dimini s h es or des t roy s gener a t iv e p ower a re di fficu lt t o ,
obt a i n. I t i s i n d eed a ss erted th a t i nt en se a pplic a ti on t o

, ,
m a th em a tic s r equ i ri ng a s it d oes ext rem e concent r a ti on of

,
t h ou ght i s a pt t o h a v e thi s res u lt ; a nd it i s a s s ert e d t oo

, , ,
th a t thi s r es u lt i s p rodu c ed by th e ex ces s iv e em oti on a l ex

ci tem en t o f g a mbli ng T h en a ga i n it i s a m a tt er of comm on
.
, ,
r em a r k h ow fre u ent l y m en of u n u s u a l m en t a l a ctivity l ea v e

q
'
n o offs p ri n g Bu t fa c t s of thi s ki n d a dmi t of a noth er i n t er

.
p ret a ti on T he rea cti on o f the b ra i n on the b ody i s s o vi ol ent

’
79
514 L A WS OF MU L TI P L ICA TI ON .
— th e v t xi ng of th e nerv ou s syst em i s s o a pt to p rost ra te

o er a
t he h ea rt a n d d era n ge th e diges t i on ; th a t th e i nc a p a citi es

c a u s e d i n t h es e c a s es a re p r ob a bly o ft en d u e m ore t o c on
,
s t i t u ti on a l di s t u rb a n ce t h a n t o th e di r ec t d ed u cti on w hi c h
e x c es s iv e a cti on en t a il s S u c h i ns t a n c es h a r m oni ze with th e

.
hyp o th es i s ; bu t h ow fa r th ey yi eld it p os itiv e s u pp or t we

c a nnot s a y .
3 68 . Anbj ecti on mu s t h ere be gu a r d ed a g a i nst I t is

o .
lik ely t o be u r ged th a t s i nce the civili zed r a c es a re on the ,
a v er a g e l a rg er th a n m a ny of t he u ncivili zed ra c es ; a n d s i nc e
,
they a re a l s o s om ewh a t m ore c om pl ex a s well a s m ore a ctiv e ;

th ey ou ght i n c on form ity w ith the a ll ege d genera l l a w t o
, ,
be l es s p rol ific T h er e i s h owev er n o evid en ce t o p rov e th a t

.
, ,
th ey a re s o : on the wh ol e th ey s eem ra th er the reverse , .
T he reply i s th a t were a l l oth er t h i ng s eq u a l th es e ,
s u p eri or v a r i eti es of m en sh ou ld h a ve i nferi or ra t es of i n

c rea s e. B u t oth er thi ng s a re n ot equ a l ; a n d it i s t o the
i neq u a lity of other t hi ngs th a t thi s a pp a rent a n om a ly i s
a tt r ib u t a bl e A l rea dy we h a v e s een h ow m u c h m ore f ertil e
.
d om es tic a t ed a n i m a l s a re th a n th ei r wi l d ki ndred ; a nd the '
ca u s es of this gr ea t er fert ility a re a l s o the ca u s es o f the

g rea t er fertility rel a tiv e or a b s ol u t e whi ch civili zed m en
, ,
exhibit wh en co m p a r e d w ith s a v a ge s .
T h ere is th e differen c e in a m ou n t of f ood A u s t ra li a ns .

,
F u egi a n s a nd s u nd ry ra ces th a t might be na m ed a s h a vi n g

,
l ow r a t es o f m u ltiplic a ti on a re obvi ou s ly u nd er fed , T he .
s k etch es o f na tiv es con t a i ned i n th e v ol u m es o f L ivi ng s t one ,
B a k er a nd oth ers yi eld cl ea r p roofs of th e ext rem e d epl eti on

, ,
c om m on a m on g th e u ncivili z ed I n q u a lity a s well .
a s i n q u a n tity th ei r feedi n g i s ba d Wild fru it s i nsect s

,
.
, ,
l a rv ae v ermi n &c which we ref u s e with di sg us t o ft en ent er

, ,
.
, ,
l a rgely i nt o th ei r di et a ry Mu ch of thi s i n feri or f ood th ey

.
ea t u nc ook ed ; a n d th ey h a v e not ou r el a b ora t e a ppli a n ce s
for m ech a ni ca lly p rep a ri ng it a nd rej ec ti ng its u sel ess p a rts

-
, .
S o th a t th ey l iv e on m a tt ers of l es s nu t ritiv e v a l u e which ,

MU L T I P L I CA TI ON OF T HE H U MA N RA CE .
51 5
c os t m ore b ot h t o m a s tic a t e a nd t o diges t Fu rth er

.
,
t o u nc iviliz e d m en s u ppli es o f f oo d c om e v ery i rr egu l a r ly .
L on g p eri od s of s ca rc ity a r e divid ed by sh ort p er i od s o f

a b un d a n c e . A nd th ou gh by g orgi ng w h en opp ort un ity
occu rs s om ethi n g i s d on e t owa rd s c om p en sa ti n g for p revi ou s
,
fa s ti ng yet th e eff ect s of p r ol ong ed s t a r v a ti on c a nnot be

,
neu t ra liz ed by oc c a si on a l en or m ou s m ea l s Bea ri ng i n mi n d

.
,
t oo th a t i m p r ovid en t a s th ey a re s a v a ges o ft en b es ti r th em
, ,
s elv es only u nd er p res su re of h unger we m a y f a i rly c on sid er

,
'
t h em a s h a bit u a lly ill nou ri s hed— m a y s ee th a t ev en the

-
p oorer cl a s s es of civili ze d m en m a ki n g regu l ar m ea l s on f ood

,
s ep a r a t ed f ro m i nnu t r itiv e m a tt er s e a s y t o m a s tic a t e a nd

,
diges t t o ler a bly g ood i n qu a lity a n d a d equ a t e if not a b u nd a nt

,
i n qu a ntity a re m u ch b et t er n ou r i s h ed
,
.
T h en a ga i n th ou gh a g rea t er c on
, ,
su mpti on i n m u s c u l ar
a c ti on a pp e a r s t o be u nd erg one by c ivili z e d m en th a n

by s a v a g es ; a n d th ou gh it i s p rob a bly t ru e th a t a m ong ou r
l a b ou ri n g p eopl e the d a ily r ep a i rs c ost m ore ; yet i n m a ny
c ases th ere d oes n ot exi s t s o m u c h differen c e a s w e a r e a t
p
t o s u pp os e . T h e ch a se i s v ery l a b ori ou s ; a n d g rea t a m ou nts
o f e x erti on a r e g one th r ou gh by t h e l ow es t ra c e s i n s eeki ng
a n d s ecu ri ng th e odd s a nd en d s o f w ild f o o d on whi c h th ey
l a r gely d ep end . We n a tu ra lly a ss u m e th a t b ec a u s e ba r

ba ri a ns a re a v er s e t o r egu l a r l a b ou r th ei r m u s cu l a r a cti on
,
-
i s l ess th a n ou r own Bu t thi s i s n ot n ec es s a r ily t ru e

.
The.
m on ot onou s t oil i s wh a t th ey ca nnot t ol er a t e ; a nd th ey m ay

be rea dy t o g o th r ou gh a s mu c h or m ore ex erti on wh en
it i s j oi ned w ith excit em ent I f we rem emb er t h a t th e
‘
s p ort s m a n wh o gl a dly s cr a mbl es u

p a n d d own rou gh hill
s id es a l l da y a ft er gr ou s e or d ee r w ou ld thi nk hi m s el f h a r dly
,
u se d h a d h e t o s p en d a s mu ch ef for t a nd ti m e i n diggi ng ; we
s h a ll see th a t a s a v a g e w h o i s th e re v er s e o f i n d u s t r i ou s
,
ma y nev erth el ess be s u bj ect t o a mu s c u l a r wa s t e not v ery

di fferent i n a m ou n t f rom th a t u nderg one by th e i n d u s
t ri ou s
. Wh en it i s a dd ed th a t a l a rger phys i ol o
g i ca l e xp en dit u re i s en t a il ed on th e u n civili zed th a n on th e
.
51 6 L A WS OF MU L T I P L I CA T ION .
c ivili zed by the a b s en ce of g oo d a ppli a nces for s h elt er a nd

p rotecti on— th a t i n s om e ca s es th ey h a v e t o m a k e g ood a
g rea t er l os s of h ea t a nd i n oth er c a s es s u ffer mu ch wea r f rom
,
i rri t a ti ng s wa rm s o f i n sects ; we s h a ll see th a t the t ot a l c os t

o f sel f m a i nt ena n ce a m on g th em i s p r ob a bly i n m a ny c a s es
-
li ttl e l es s a nd i n s om e c a s es m ore th a n it is a m on g ou rs elv es

, , .
S o th a t t h ou gh on t h e a v er a ge the c ivili zed a re p r ob a bl y

, ,
l a rger th a n th e s a v a g e ; a nd th ou gh th ey a re i n thei r ,
n er v ou s s y s t em s a t l ea s t s om ewh a t m ore c om pl ex ; a n d
,
th ou gh oth er thi ng s equ a l th ey ou ght t o be the l es s

, ,
p rolific ; yet oth er thin g s a re s o u n equ a l a s t o m a ke it

qu it e c onf orm a bl e t o th e g en era l l a w th a t th ey s h ou ld be
m ore p rolific I n § 3 6 5 we ob s erv ed h ow a m ong i n feri or
.
,
a ni m a l s high er ev ol u ti on s om etim es m a k es s elf p res er v a t i on

,
-
fa r ea s i er by op en in g th e w a y t o res ou r c es p revi ou s ly u n
,
a v a il a bl e : so in v olvi ng a n u ndi m in i s h e d or ev en a n i n ,
c r ea s ed r a t e o f g enes i s
,
A nd si m il a rly w e m a y exp ect th a t
.
a m on g r a c es of m en th os e wh os e s light fu rth er d ev el op
,
m ent s h a v e b een f oll owed by h a bit s a nd a rt s which imm ensely
f a ci l it a t e life w ill not exhibit a l ower d eg ree of f ertility a nd

, ,
m a y ev en exhi bit a high er .
— a ki nd red oh
§ 3 69 . m One
or e obj e cti on h a s t o be m et
jecti on t o whi ch th ere i s a kin d red reply C a s e s m a y be .
n a m ed of m en c on s pi cu ou s for a cti vity b o dily a n d m en t a l

-
, ,
wh o w ere a ls o not ed not for l es s genera tiv e p ower th a n u s u a l

, ,
bu t for m ore A s t h ei r su p eri oriti e s i ndi ca t e high er d eg rees

.
o f ev ol u ti on it m a y be u r g ed th a t s u ch m en sh ou ld a cc or d
, ,
i ng t o the th eory h a v e l ower d eg rees of rep rod u c tiv e a c tivity

,
.
T he f a c t th a t h ere a l ong w ith i n c rea s ed p owers o f s elf pr e

,
-
s erv a ti on there g o i nc rea s ed p ower s of ra ce—p r op a ga ti on

, ,
s eem s i r rec on c il a bl e w i t h th e g enera l d oct ri n e Rec on cili a .
ti on i s not difficu lt h owev er .
T he c a s es a re a na l og ou s t o s om e b ef ore na m ed i n which ,
m ore a b un d a nt f ood s imu lt a neou s ly a ggra ndi zes the i ndi

MU L T IP L ICA TI ON OF T H E H U MA N RA CE .
51 7
vid u a l and a dd s t o the p r odu c ti on of new i ndi vi du al s f the

di fl e renc e b et ween th e c a ses b ei ng th a t i ns t ea d of a b ett er
,
e xt ern a l s u pply of m a t e ri a l s there i s a b ett er i n t erna l

u t ili za ti on o f m a t e ri a l s C rea t ures o f the s a m e sp eci es n o t o
.
ri ou s l y di ffer i n g oo d nes s o f con s ti t u ti on H er e th ere i s s om e

.
vi s cera l d efect s h owi ng it s el f i n f eebl enes s of a l l th e fu nc

,
ti ons ; whil e h ere s om e p ecu li a r ity of org a n i c b a l a n ce s om e ,
high qu a lity o f ti s s u e s om e a bu nd a n ce or p ot ency o f the

,
diges tiv e j u i ces giv es t o the s y s t e m a p erp et u a l high tid e of

, ,
r i ch bl oo d w hich s er v es a t on ce t o en h a n c e th e vit a l a c tiviti es

,
a n d t o ra i s e t h e p ower of p ro p a g a ti on S u ch v a ri a ti on s ,
.
h owev er a re i n d ep end ent of ch a nges i n the p r op or ti on be

,
t w een I ndivid u a ti on a n d Genes i s T h i s rem a i n s th e s a m e

.
,
w hil e b oth a r e i n crea se d or d ec rea s e d by th e i n crea se or

d ec rea se of the com m on s t oc k of m a t eri a l s .
A n ill u s t r a t i on w ill b es t c l ea r u p a ny p erpl exity L et u s .
s a y th a t th e fu el b u rn t i n t h e fu rn a c e o f a l ocom otiv e s t ea m
e n gi ne a ns w ers t o t h e f o o d w hi c h a m a n c ons u m es
,
L et u s .
s a y th a t th e p r od u ce d s t ea m exp en d e d i n w o r ki n g th e en gi ne ,
co rres p ond s t o th a t p or ti on o f a b s o r b ed n u t ri m en t wh i c h
’
ca rr i es o n t h e m a n s f u nc ti on s a n d a c tiviti es A nd l et u s .
s ay th a t the s t ea m bl owi n g off a t the s a fety v a lve -

,
a n s wer s t o th a t p orti on of th e a b s orb e d nu t rim en t whi c h

'
g oes t o th e p rop a ga ti on of th e ra ce S u ch b ei n g the c ondi .
ti ons o f the ca s e s ev era l ki n d s of v a ri a ti on s a re p os s i bl e

, .
A l l oth er c i rcu m s t a nces r em a i ni ng the s a m e th ere m a y be ,
c h a n ges o f p r op o rt i o n b et w een th e s t ea m u sed for w or ki n g
th e en gi n e a n d the s t ea m th a t es ca p es by th e s a fety v a lv e -
.
T h ere m a y be a s t r u c t u ra l or or ga ni c c h a ng e of p r op ort i o n .
By enl a rgi n g t he s a fety v a lve or wea k eni ng i ts s p r i n g w hil e

-
,
the cyli nd ers a re red u ced i n s i ze th ere m a y be es t a bli s h e d a

,
c on s tit u ti o na lly s m a ll p owe r of l oco m oti on a nd a c on s t i t u

-
t i ona l l y l a rg e a m oun t of esc a p e st ea m ; a nd i nv ers e v a ri a ti on s

- -
s o p r o d u ce d will a nsw er t o the i nv ers e v a ri a ti ons b et ween

,
I ndivid u a t i on a nd Genes is whi ch diff erent t y p es of orga nism s

51 8 L A WS OF “
MUL T I P L I CA TION .
sh ow u s A ga i n th ere m a y be a f uncti ona l ch a ng e of pro

.
,
'
p orti on I f the engi ne h a s t o dr a w a c ons id era bl e l oa d the

.
,
a b s t ra c ti on of s t ea m by the cyli nd ers grea tly red u c es the

di scha rg e by th e s a fety v a lv e a nd if a high v el ocity i s k ept
-
u t h e di s ch a r g e f rom the s a fe ty v a lv e en ti r ely cea s es C on

p
-
.
,
versel y if th e v el ocity is l ow t h e e sc a p e st ea m be a rs a l a rg e
, ,
-
ra ti o t o t h e s t ea m c on su m e d by t h e m ot or a pp a ra t u s ; a n d if
th e engi ne b ec om es s t a ti on a ry th e w h ol e o f the s t ea m es c a p es
by the s a f ety v a lve T hi s i nv ers e v a ri a ti on a nsw ers t o t h a t
-
.
w hich we h a v e t ra c ed b et ween E xp endit u r e a n d Genes i s a s ,
di spl a y ed i n the c on t ra s t s b et ween s p eci es of t he s a m e typ e

bu t u nlik e a ctiviti es a nd i n th e c ont ra s ts b et ween a ctiv e a nd
,
i na ctiv e i ndividu a l s of th e s a me sp ec i es B u t now b ey ond .
t h e s e i nv ers e v a ri a ti on s b et w een t h e qu a nti t i es o f con s u m ed

s t ea m a n d es c a p e st ea m w hich a re s t ru c t u ra lly a n d f u nc ti on
-
,
a lly c a u s ed th ere a re c oi nc id en t v a ri a ti on s p r od u cibl e i n b o th

, ,
by ch anges i n th e qu a nti ty of s t ea m su ppli ed— ch a nges w hi ch

m a y be c a u s ed i n s ev era l w a ys I n th e fi rs t pl a ce t he fu e l
.
,
th rown i nt o the f urn a ce m a y be i nc rea se d or m a d e b ett er .
Oth er thi ng s equ a l t h ere will r es u l t a m ore a c tiv e l ocom o

,
ti on a s w ell a s a g rea t er es ca p e ; a n d this will a n s w er t o th a t

s imu lt a n eou s a dditi on t o i t s i ndivid u a l vig ou r a nd i t s r ep ro
du cti ve a ctivity ca u sed i n a n a nim a l by a l a rg er q u a ntity or

, ,
a s u p eri or q u a li ty of f oo d I n the s ec on d pl a c e th e s t ea m
, .
,
gen era t e d m a y be economiz ed L os s by r a di a ti on f rom th e .
b oil er m a y be l es s ened by a c ov eri ng of non c ond u cti ng s u b -
st a nc es ; a nd p a rt of the s t ea m th u s p revent ed f r om con

den si ng will g o t o i ncrea s e th e w orki n g p ow er o f the engi n e
, ,
whil e p a rt will be a dd ed t o the q u a n tity bl ow i ng off T hi s .
v a ri a ti on c orres p onds t o th a t sim u lt a neou s a d diti on to b odily

vig ou r a nd p rop a g a tiv e p ower whi c h resu lts i n a ni m a ls th a t,
h a v e t o exp end l es s i n k eepi ng u p th ei r t em p era t u res I n .
t h e thi rd pl a ce by i mp rov em en t o f the s t ea m —g enera ti ng

,
a pp a ra t u s m or e s t ea m m a
, y be obt a i ned f ro m a giv en w eight
o f fu el . A b ett er f orm ed ev a p or a ti ng s u r fa c e or b oil er t u b es
-
,
whi ch c ond u ct m ore ra pid l y or a n i nc rea s ed nu mb er o f th em

,
MUL T I P L I CA T I ON OF T H E H U MA N RA CE .
51 9
m ay c au e a s l a rger a bs orpti on of hea t f rom the bu rni ng m a s s

or th e h ot g a ses it giv es off ; a n d t h e e xt r a s t ea m g ener a t e d
by thi s ext ra h ea t will a s b ef ore a u gment b oth the m otiv e

, ,
f orc e a nd the e mi s s i on th r ou gh th e s a fety v a lv e A nd thi s -

.
l a s t c a s e of c oi ncid ent v a ri a ti on i s p a ra ll el t o the ca s e w i th

,
whi ch we a re h ere c oncerne d— the a u gm enta ti on of i n dividu a l

exp en dit u r e a n d o f rep r o d u c tiv e e n ergy th a t m a y be ca u s e d ,
by a s u p eri ority of s om e orga n on whi ch the u tili zi ng or

econ om i z i ng o f m a t er i a ls d ep en d s .
Ma nifes tly theref ore a n in c rea s ed exp endit u re for Gene

, ,
s i s or a n i ncrea se d exp en dit u re for I ndivid u a ti on m ay a r i s e

, ,
i n one of tw o qu it e di fferen t w a ys— e ith er by dimi nu ti on of

t h e a n t a g oni s tic exp endit ure or by a dditi on t o th e st ore w hi c h
,
s u ppli e s b oth exp en di tu res ; a nd c on fu s i on resu lt s f rom not

di s ti ngu i s hi ng b et ween th es e Giv en th e ra ti o 4 t o 2 0 a s
.
,
e xp re s s iv e o f th e r el a tiv e cos ts of Genes i s a n d I n divid u a ti on
th en the exp endi tu re for Gen es i s m a y be ra i s ed to 5 whil e the

exp enditu re fo r I n divid u a ti on is ra i s ed t o 2 5 w ith ou t a ny ,
a lt era ti on of typ e ,
m er ely by f a v ou ra bl e ci rcu m st a nces or
s u p er i ority of c on s tit u ti on On the oth er h a nd ci rc u m s t a nc es
.
,
rem a i ni n g th e s a m e t he exp endit u re for Genes i s m a y be

,
ra i s ed f rom 4 t o 5 ,
by l oweri n g the exp en dit u re for I ndi
vidu a ti on f rom 2 0 to 1 9 : whi ch ch a nge of ra ti o m a y be
e ith er fu n cti on a l a nd t emp ora ry or st ru ct u r a l a n d p er
,
m anent A nd only wh en it is the l a st d oes it ill u s t ra t e th a t

.
i nv ers e v a r i a ti on b et ween d eg ree of ev ol u ti on a nd d eg ree of

p r oc rea tiv e di s s ol u ti on whi ch we h a v e ev erywh ere s een
, .
3 7 0 T h ere i s n o
. pp ose th en th a t the l a ws
r ea s on to su , ,
o f m u ltipli ca ti on w hich h old of o th er b ei ng s d o not h old o f ,
th e hu m a n b ei ng . On th e c ont ra ry th ere a re s p eci a l f a ct s ,
whi ch u nit e with gener a l implic a ti on s t o sh ow th a t th ese

l a w s do h old of th e h u m a n b ein g Th e a b sence o f di rect .
evid en c e i n s om e c a s es wh er e it might be l ook ed for w e fi n d ,
fu lly expl a i ned wh en a l l th e f a ct or s a re t a k en i nt o a cc ou nt .
A nd c ert a i n s eem i ngl y a dv ers e f a c ts p r ov e on ex a m i n a ti on

-
, ,
5 20 L A WS OF MUL TIP L I CA T I ON .
t o be f a ct s b el on gi ng t o a di fferen t c a t eg ory fr om th a t i n
w hi ch th ey a re pl a c ed a n d h a r m oni ze with t he res t w h en
,
rightly i nt er p r et ed .
The c on form ity of h um a n fertility to t he l a ws of m u ltipli

c a ti on i n genera l b ei ng gr an t ed it rem a i n s t o i nq u i re wh a t
, ,
ef fec t s mu s t be c a u s ed by p erm a nen t ch a ng es i n m en s na t u res ’
a n d c i r cu m s t a n ce s T hu s fa r we h a v e ob s erv e d h ow by the i r
.
,
e x cepti ona lly high e v o l u ti on a n d ex cepti on a lly l ow fertility

- -
,
m a nki nd di s pl a y the i n v er s e v a ri a ti on betw een I ndividu a ti on

a n d Genesis i n one of i t s e xt r em es
, A n d we h a v e a l s o ob .
s erv e d h ow m a n ki n d lik e o th er ki n d s a r e fu n cti ona lly ch ange d

, ,
i n th ei r ra t es of m u ltipli ca ti on by c h a ng es of c onditi ons B ut .
we h a v e n ot ob serv ed h ow a lt era ti on of s t ru c t u re i n Ma n
ent a il s a lt er a ti on of f ertility The i n fl u ence of thi s f a ct or is
.
s o en t a n gl ed w ith t h e i n fl u en c es o f o th er f a c t or s w hi c h a re
for th e p r es ent m ore p ot ent th a t we ca nn ot rec og ni ze i t , .
H ere if we p rocee d a t a l l we m u st p roc eed d ed u ctiv ely

, , .
[ NOTE —
F rom a m ong the pu blic a ti ons of th e A m eric a n
A c a d emy of P olitic a l a nd S oci a l S c i en c e th ere w a s s ent t o
‘
m e s om e y ea r s a g o a n es sa y entitl ed T he S igni fi ca nc e of a
D ecrea s i ng B i rt h Ra t e by ( Mi s s ) J L B rownell F ell ow i n . .
,
P olitic a l S ci en c e B ry n Ma wr Coll ege

,
T hi s es s a y cont a i ns .
a nu mb er of el a b ora t e c omp a ris ons d ra wn f rom the vit a l
s t a ti s ti c s of th e t enth U ni t e d S t a t es C ens u s T he res u lt s .
of th es e c omp a ri s on s a re th u s s u mm ed u p
1 . Wh eth er or n ot i t be t ru e th a t t h e m ea ns s p ok en of by D r .
B i l l i ng s , M D um
. ont , M L e v a s s eur,
. a nd D r E d son h as b e c om
. e an
i m p ort a nt fa ctor i n th e d i m i ni sh i ng bi rth -

ra t e of c i v i l i z ed c ou ntri e s ,
i t i s e vi d ent t h a t i t i s not t h e onl y fa c tor, a nd t h at , q ui te a p a rt from

v ol u ntary pre v enti on , t h ere i s d i s ti nc t probl em t o b e i n v est i g a t e d
a .
T hi s i s sh ow n by t h e fa c t t h at t h e w h i t e a nd th e c ol ored bi r th ra te -
v ary t og et h er .
2 . Mr S p enc er s g eneral i zat i on th a t th e bi rt h

.
’ -
ra t e d i m i ni sh e s as
th e ra te of i n d i vi d u a l e v ol uti on i ncreases i s c onfi rm ed bv a c om p ari s on

of t h e bi rth -rat e s w i t h t he d e a th -rat e s from ner v ou s d i s e a s e s , a nd a l s o
MU L TI P L ICA TI ON OF T H E H UMA N RA CE .
5 21
w i th t h e d e ns i ty of p op u l ati on, th e v al u es o f a gri c ul tu ral a nd m a nu

”
fac t u rc d prod u ct s , an d t h e m ortg a g e i nd eb te d ne ss .
Of s mu ltit u di nou s differenc es of ra ce cl a s s m ode

cou r e ,
o f li vi ng o c c u p a ti on l oc a lity m a k e it diffic u lt t o d ra w p os i
, , ,
tiv e in ferences from the d a t a ; bu t th e i nferences a b ov e d ra wn

a re h e ld t o rem a i n ou t s t a n di ng a ft er a ll ow i n g for a l l th e
q u a lifyi ng c onditi ons ]

C H A P T E R XI I I .
H UM A N P O P U L A T IO N IN TH E F U T URE .
§3 71 . ANY fu rth er ev ol u ti on i n the m ost highly ev olv ed -
of t er rest ri a l b ei ngs Ma n mu s t be o f th e s a m e na t u re a s
, ,
e v ol u ti on i n g enera l S t ru c t u r a lly c ons id ered it m a y c ons ist

.
,
i n gr ea t er i nt egra ti on or g rea t er diff eren ti a ti on or b oth

, ,
a u gm ent e d b u lk or i n cre a s e d h et ero g eneity a n d defi ni t enes s

, ,
o r a c ombi n a ti on of th e t w o F u ncti ona lly c onsid ered i t

.
,
m a y c onsist i n a l a r g er s u m o f a cti ons or m ore mu ltipli e d ,
v a r i eti es of a cti ons or b oth— a l a rger a m ou nt of s ens ibl e a n d

,
i nsens ibl e m oti on genera t ed or m oti ons m ore nu m erou s i n

,
th ei r ki nds a nd m ore i nt ric a t e a nd ex a ct i n th ei r co ordi na -
ti ons or m oti ons th a t a re gr ea t er a lik e i n qu antity c om

, ,
p l e xi t
y , a n d p re ci s i on .
E xp res s i n g t he c h a nge i n t erms of th a t m ore sp eci a l

ev ol u ti on d i s pl a y ed by o rg a n i s m s ; we m a y s a y th a t it m u s t
be one whi ch f u rth er a d a pts th e m ovi ng equ ilib ri u m o f

org a n ic a c ti on s .A s wa s p oi nt ed ou t i n F i rs t P ri nci p l es ,
“
§ 1 7 3
,
th e m a i n t e n a n c e o f s u c h a m o vi n g eq u ilib r i u m re ,
q u i res the h a bit u a l genes i s o i i nt ern a l forces corres p on di ng

i n nu m b er di recti ons a nd a m ou nt s to the ext ern a l i ncid en t
, ,
f orc es— a s m a ny i nner f u ncti ons s i ngl e or c om bi ned a s th ere

, ,
”
a re s i ngl e or comb i n ed ou t er a ct i on s t o be m et A nd it .
“
wa s a l s o p oi nt ed ou t th a t th e s t ru c t u ra l com pl exity a ec om
p a nyi n g f u ncti ona l equ ilib ra ti on i s d efi na bl e a s o ne i n whi ch
,
th ere a re a s m a ny sp eci a li zed p a rt s a s a re c a p a bl e s ep a ra t ely ,
a n d j oi n tly o f c ou nt era c ti ng th e s ep a ra t e a nd j oi nt f o rce s

,
5 22
H U MA N P OP U L A TI ON I N T HE F U T URE .
5 23
a mid which the orga ni s m exi sts Cl ea rly th en si nc e a l l

.
, ,
i nc om pl et enes s es i n Ma n a s n ow c ons tit u t ed a re f a il u res t o ,
m eet c ert a i n of the ou ter a cti ons ( m os tly i n v olv ed rem ot e , ,
i rreg ul a r ) t o whi ch he i s exp osed ; every a dv a n ce i m pli es

,
a dditi ona l c o or di na ti ons of a c ti on s a n d a c c omp a n yi ng com

-
p l ex i ti es o f org a n iz a ti on .
Or t o s p eci a li ze s till fu rth er thi s c onc e pti on of fu t u r e pro

,
g ress we m a y c ons id er it a s a n a dv a nce t ow a rd s c ompl eti on

,
o f t h a t c on ti nu ou s a dj u s tm en t o f i n t ern a l t o ext erna l rel a
ti on s which L ife s h ows u s I n P a rt I of thi s wor k where

, . .
,
it w a s s h own tha t the c orres p ond ence be t ween i nn er a nd

ou t er a ct i on s w hi ch u nd er i t s ph en om en a l a s p ec t we c a ll ,
L ife i s a p a rticu l a r ki n d of wh a t in t erms of E v olu ti on w e

, , ,
c a ll e d a m ovi n g eq u ilib ri u m ; it w a s s h own th a t the d eg ree

o f li fe v a ri es a s t h e d eg ree o f c or res p ond enc e Grea t er evo .
l u ti on or high er li f e i m pli es th en s u ch m o difi ca ti on s o f

, ,
h u m a n n a t u re a s s h a ll m a k e m ore ex a c t the exi s ti ng c orre

i
s p on dences or s h a ll es t a bli s h a dditi on a l c or res p on d ences

, or ,
b oth C onnexi ons of ph enom ena o f a ra re di st a nt un obtru

.
, ,
s iv e or i n t ric a t e ki n d w hich w e e ith er s u f

, ,
fer f ro m or do not
t a k e a dv a n t a g e of h a v e t o be res p ond ed to by new c onnexi ons
,
o f id ea s a n d a c t s p r op er ly co mbi n ed a nd p r o p orti one d : th ere

,
m u s t be i ncrea s e of k now l edg e or s kill or p ow er or of a l l , , ,
th es e A nd t o effec t this m ore ext ens iv e m ore v a ri ed a nd

.
, ,
m ore a cc u r a t e c c o r di na ti on of a c ti ons t h ere m u s t be org a n i

,
-
,
za t i on o f still g rea t er h et ero g en eity a n d defi ni tenes s .
§ 3 72 . L et b efore p roc eedi ng c on s id er i n wh a t pa r

u s, ,
t i cu l a r w a y s thi s fu r th er e v o l u ti on thi s high er li fe this , ,
grea t er co or di n a ti on o f a cti ons

-
m a y be e xp ec t ed t o s h ow
,
it s elf .
W ill it be i n s t ren gth ? P rob a bly n ot t o a ny c ons id er a bl e

d eg ree Mech a ni ca l a ppli a nc es a re f a s t s u ppl a nti n g b ru te
.
f orc e a n d d ou btl es s will c onti nu e d oi ng this T h ou gh a t

,
.
p r esent civili zed n a ti ons l a rg ely d ep end for self p res erv a ti on -
on vig ou r o f li m b a n d a r e lik e ly t o d o s o w hil e w a r s con

,
5 24 L A WS OF MU L T IP L I CA T ION .
t i nu e ; yet th a t p rogr ess iv e a d a p t a ti on t o the s oci a l s t a t e

Whi ch mu s t a t l a s t b ri ng w a rs t o a n end will l ea v e t h e ,
a m ou n t of m u s cu l a r p ow er t o a dj u s t it s el f t o th e r eq u i rem ent s
of a p e a c e f u l r egi m e T h ou gh t a ki n g a l l thi ngs in t o a ccou n t

.
, ,
t h e mu scu l a r p ower th en requ i red m ay n ot be l es s th a n n ow ,
th ere seem s n o rea s on why m ore s h ou ld be requ i red .
Will it be s wi ftnes s or a gility ? P rob a bly not I n s a v a ges .
t h ese a r e imp ort a n t el em en t s of th e a bility t o m a i nt a i n

li fe ; bu t i n c ivili zed m en th ey a i d s el f p r es erv a ti on i n qii i te
-
mi nor d eg rees a n d th ere s eem s n o ci rcum s t a n ce lik ely t o

,
nec es s it a t e a n i n c r ea s e o f th em By ga m es a n d gymn a s ti c
.
c omp etiti on s s u ch a tt rib u t es m a y i n d ee d be a rtifi c i a lly i n

,
c r ea s ed ; bu t n o a rtifi c i a l i n c rea se w hi ch d o es n ot b ri ng a
p rop orti ona te a dv a nt a ge c a n be p erm a nen t ; s i nce other ,
thi ngs equ a l i ndivid u a l s a nd s oci eti es th a t d ev ot e the s a m e

,
a m ou n t s of en er gy i n w a y s whi c h s u b s erv e li fe m or e effectu
a lly m u s t by a n d by p r ed om i na t e
, .
Will it be i n m ech a ni ca l s kill t h a t i s i n the b ett er , ,
c c or di na ti on o f co mpl e x m o v em en t s ?
-
Mos t lik ely i n s ome
d egree A wk wa rd n es s i s c on ti nu a lly ent a ili ng i nj u ri es a nd
.
d ea th s M oreov er the c om pli ca t ed t ool s whi ch c ivili za ti on

.
b ri ngs in t o u se a re c on s t a ntly req u i ri ng g rea t er d eli ca cy of

,
m a nip u l a ti on A l l the a rt s i nd u s t ri a l a nd m s th eti c a s th ey

.
, ,
d ev el op i m ply a c orresp ondi ng d e v el opm ent of p erceptiv e a n d

,
e x ecu t iv e fa cu lti es i n m en : th e t wo a ct a nd re a c t .
Will it be i n i nt el l igence ? L a rgely no d ou bt T h ere i s ,

.
a m pl e r oom for a dv a nce i n thi s di rec ti on a n d a m pl e d em a n d ,
for it . Ou r liv es a re u niv ers a lly sh ort ened by ou r ignora nce .
I n a tt a i ni ng c ompl et e k n owl edge o f ou r own na t u res a nd o f

th e n a t u res o f s u rrou n di ng thi ngs— i n a s c ert a i n i ng t he con
d i ti ons o f exi s t ence to whi ch we m u s t c onf or m a nd i n di s ,
c ov eri n g m ea ns o f c on fo rmi ng t o t h em u nd er a l l v a ri a ti ons
o f s ea s ons a nd c i rcu m s t a nce s ; we h a v e a b u n d a nt S c op e for
i nt ell ect u a l p rogres s .
Will it be i n m ora lity th a t is i n gre a t er p ow er of self

, ,
H U MA N P OP U L A T I ON I N T H E F U T U RE .
5 25
re gul a ti on ? l p erh a p s m os t l a rgely Right con

L a rgely a so : .
d u ct i s u s u a lly s om e s h ort of m ore f rom d efect of w ill th a n

d efect of kn owledge For t h e right c o or di na t i on of th os e
.
-
c ompl ex a cti on s w hi c h con s tit u t e h u m a n li fe i n i t s c ivili z ed
for m th ere g oes n ot only th e p re requ i s it e — r ec og niti on of

,
-
th e p r op er c ou rs e ; bu t the fu rth er p retr equ i s i te —a du e

i m p u l s e t o p u rs u e th a t c ou rs e On c a lli n g t o mi nd ou r .
d a ily f a ilu res t o fu lfil oft en rep ea t ed res ol u t i on s we sh a ll

-
,
p erceiv e th a t l a ck of the need fu l d es i re r a th er t h a n l a c k of ,
t he need fu l i n s ight i s th e chi ef c a u se of f a u lty a c t i on

, A .
fu rt h er en d owm en t of th os e feeli ng s which c ivili za ti on i s

d ev el opi ng i n u s— s enti m ent s res p on di ng t o the requ i rem ent s
o f th e s oci a l st a t e — e m ot iv e fa c u lti es th a t fi nd th ei r g ra t i fi
ca ti on s i n th e d u ti es d e v ol vi n g on u s— m u s t be a cqu i re d
be fore the cri m es e xc es s es di sea s es imp rovid ences di sh ones

, , , ,
ti e s a n d cru el ti es th a t now so grea tly dimi ni s h th e d u ra ti on

, ,
of lif e c a n c ea se
,
.
T hu s l ooki ng a t th e sev era l p os s ibiliti es a nd a ski ng

, ,
wh a t di recti on thi s fu rth er ev olu ti on thi s m ore c ompl et e ,
m ovi ng eq uili b ri u m thi s b ett er a dj u st m ent of i nn er to ou t er

,
r el a ti ons this m ore p er fe ct co or din a ti on of a cti on s i s lik ely

,
-
,
t o t a k e ; we c on cl u d e th a t it mu s t t a k e m a i nly t h e di recti on
of a high er i n t ell ect u a l a nd em o ti ona l d ev el op m ent .
§ 3 73 T his
c oncl u s i on we sh a ll fi nd equ a lly f orc ed on u s
.
if we i nqu i re for t he c a u ses whi c h a r e t o b ri n g a b ou t s u ch

res u lt s N o m or e i n the c a s e of Ma n th a n i n th e c a s e of
.
a ny ot h er b ei ng c a n w e p resu m e th a t ev ol u ti on h a s t a k en
,
pl a ce or w ill h er ea ft er t a k e pl a c e s p ont a n eou sly

, I n th e , .
p a s t a t p resen t a nd i n the fu t u re a l l m odific a ti on s fu nc

, , , ,
t i ona l a nd or g a nic h a v e b een a re a n d m u s t be imm edi a t ely

, , , ,
or r em ot ely c on s eq u en t on s u rrou n di n g c ondi ti on s Wh a t .

,
th en a re th os e ch a nges i n the envi r onm ent t o w hich by

, ,
di rec t o r i n di rect equ ilib r a ti on t h e h u m an or ga n i s m h a s be en

,
a dj u s ti n g its elf i s a dj u sti ng it s elf n ow a nd will con ti nu e t o

, ,
a dj u s t it s elf ? A nd h ow d o th ey neces s it a t e a high er evol u

ti on of the org a ni s m ?
C ivili z a ti on ev er ywh ere h a vi ng for its a nt ec ed en t th e i n
,
c rea s e of p op u l a t i on a nd everyw h ere h a vin g for on e of i ts

,
c on sequ ences a d ec r ea s e o f c ert a i n r a c e d est royi ng f or c es h a s -

,
for a fu rth er c on s eq u enc e a n i n crea se o f c er t a i n oth er r a ce

d es t royi ng f orces D a nger of d ea th f rom p red a t ory a ni m a l s
.
l es sens a s men g row m ore nu m er ou s T h ou gh a s th ey s p rea d .

,
o v er t h e E a rth a n d divid e i nt o t rib es m en b ec om e w ild ,
b ea s t s t o one a n oth er yet th e d a nger of d ea th from thi s

,
c a u s e a l s o di m i n i s h es a s t rib es c oa l esce i nt o n a ti on s B u t th e .
d a ng er o f d ea th w hi ch d oes not di m i ni s h i s th a t p rodu ced by ,
a u g m en t a ti o n o f nu mb ers it s elf— th e d a ng er f rom d efici ency
o f f oo d S u pp os i n g h u m a n n a t u re t o rem a i n u nch a nged th e

.
,
m ort a lity h en ce res u ltin g w ou ld on th e a v era g e r i s e a s , ,
h u m an b ei ng s m u ltipli ed I f m ort a lity u n der s u ch c on di

.
,
ti ons d oes not r i se it m u st be b eca u se the s u pply o f f ood

, ,
a l s o a u g m ent s ; a n d thi s i m pli es s om e ch a ng e i n h u m a n
h a bits w rou ght by s t ress of h u m a n need s H ere th en is .

, ,
the p er m a n en t c a u s e of m o difi c a ti on t o which civili zed m en

a re e xp osed T h ou gh the i n t en s i ty of its a cti on i s ev er b ei ng
.
mitig a t ed i n on e di r ecti on by grea t er p r od u cti on of food it ,
i s i n th e oth er di rec ti on ev er b ei ng a dd ed t o by t h e g rea t er

, ,
p rodu cti on of i n divi d u a l s Ma nifestly th e wa nts of th ei r .

,
re d u nd a n t nu mb ers c on s tit u t e th e on ly s timu l u s m a n ki nd
h a v e t o obt a i n m ore nec es s a ri es of life Were n ot th e d em a nd .
b ey on d th e su pply there w ou ld be no m otiv e t o i nc rea se th e

,
s u pply A nd m a n i fes tly thi s ex c es s of d em a n d ov er s u pply

.
,
i s p erenni a l : t hi s p res s u re o f p op u l a ti on of which it i s th e ,
i nd ex c a nn ot be el u d ed
,
T h ou gh by th e em igr a ti on th a t
.
t a k e s pl a c e w h en th e p ressu re a rriv es a t a c ert a i n i nt en s i t y ,
t emp ora ry reli ef i s from tim e t o tim e obt a i ned ; yet a s by ,
thi s p roces s a l l h a bit a bl e c ou n t ri es mu s t b ecom e peo pl ed it

, ,
f oll ows th a t i n th e end the p res su re wh a t ev er it m ay th en ,
be m u s t be hom e i n f u ll
,
.
T hi s c on st a nt i nc rea s e of p eopl e b ey ond the m ea ns of s ub

H U MA N P OP U L A TI ON I N TH E F U T U RE .
527
s i s t en ce c a u s es t h en a nev er c ea si ng r equ i rem ent for skill

, ,
-
,
i nt elligence a nd s el f c ont rol— inv olv es th eref ore a c ons t a nt

,
-
, ,
e x erc i se o f th e s e a n d g r a d u a l g r owth o f t h em E v ery i nd u s .
t ri a l i m p rovem en t i s a t once th e p rodu c t of a high er f or m

o f h u m a n i ty a nd d e m a n d s th a t high er f or m of h u m a n ity t o
,
ca rr y it i n t o p r a c t i c e T he a ppli ca ti on of s ci enc e t o th e a r ts
.
,
i s th e b r in ging t o b ea r g rea t er i nt elligenc e for s a ti s fyi n g ou r

w a n t s a n d i m pli es c ont i nu ed p rogres s of th a t i nt ellig ence
,
.
T o g et m ore p r od u ce from th e a cr e th e f a rm er m u s t s t u dy,
ch em i s t ry m u s t a d opt new m ech a n i ca l a ppli a n ces a n d m u s t

, , ,
by the m u ltiplic a ti on of p roc esses cu ltiv a t e b oth his own ,
p ow er s a n d th e p ower s of h i s l a b ou rers T o m eet th e .
req u i r em en t s o f th e m a r k et the m a nu f a ct u rer i s per

,
p etu a l l y i m p r ovi n g h i s ol d m a c hi n es a n d i n v en ti n g n ew
o ne s ; a nd by th e p r em i u m o f high wa g es i n cit e s a rt i z a n s t o
a cqu i re gr ea t er s kill T he d a ily wid eni ng ra m ifi c a ti on s of

.
-
c o m m er ce ent a il on t h e m er c h a n t a n ee d for m or e k no w
l edge a n d m ore c om pl ex ca l cu l a t i ons ; w hil e the l es s eni n g

p r ofit s of the s hip own er f or ce him t o b u ild m ore s ci enti fi
-
ca lly t o g et ca pt a i n s of high er i n t ellig en ce a n d b ett er c r ew s

, .
I n a l l c a s es p r es su r e of p op u l a ti on i s th e origi n a l c au s e .
Were it n ot for th e c omp etiti on this ent a ils m ore th ou ght ,
a n d ener gy w ou ld n ot d a ily be s p en t on t h e b u s i nes s o f li fe

;
a nd gr owth of m ent a l p ow er wou l d n ot t a k e pl a ce
‘
D i ffi cu lty i n g ettin g a l ivi ng i s a lik e th e i n c entiv e t o a

high er edu ca ti on of c hild ren a n d t o a m ore i n t en s e a n d
,
o n —
l g con t i nu ed a ppli ca ti on i n a d u lts I n th e m oth er it
.
p rom p ts f ores ight econ om y a nd s kil fu l h ou s e k eepi ng ; i n th e

, ,
-
f a th er l a b ori ou s d a y s a n d c ons t a nt self d eni a l N othi n g bu t

,
-
.
n ec es s ity c ou ld m a k e m en s u b m it t o thi s di s cipli n e

; a nd
n othi n g bu t thi s di s c ipli ne c ou ld p ro d u ce a conti nu ed ro
p
g ress 1 on .
I n thi s ca se, a s in
m a n y oth ers N a t u r e s ecu res ea ch s t ep
,
i n a dv a n ce by a s u c c es s i on of t ri a l s ; whi ch a r e p er p et u a lly
rep e a t ed a n d c a nn ot fa il t o be rep ea t ed
, u n til su cc ess is ,
a chi ev ed .A l l m a n ki nd i n t u rn s u bj ec t th em s elv es m ore or

5 28 L A WS OF M UL T I P L ICAT I ON .
l es s t o th e di s c ipli ne d es c ribed ; th ey eith er m a y or m ay not

a dv a n c e u n d er it ; bu t i n t h e n a t u r e of t hi n gs only th os e
, ,
wh o d o a dv a n ce un d er it ev entu a lly s u rviv e F or nec es .

,
s a ri l y f a m ili es a n d r a ce s wh o m
,
thi s inc rea s ing diffi cu lty o f
g ett ing a l ivi ng whi ch ex c es s of fert ili ty ent a il s d oes n ot ,
s t i m u l a t e t o imp ro v e m ent s i n p ro d u cti on— th a t i s t o g r ea t er ,
m ent a l a ctivity— a re on th e high r oa d t o exti nct i on ; a n d

m u s t u l tim a tely be s u ppl a nted by t h os e wh om the p ress u re
d oes s o s timu l a t e T hi s t ru th we h a v e rec ently s een exem
.
p l i fi e d i n I r el a n d A n d h ere i n d ee d w i t h ou t f u rth er ill u s

.
, ,
t ra t i on it w ill be s een th a t p r em a t u re d ea th u nd er a l l its

, ,
f orm s a nd from a l l i ts ca u s es c a nnot fa il t o work i n th e ,
s a m e di r ec ti on F or a s th ose p r em a tu rely c a rri ed off m u s t

.
,
i n t h e a v era g e of c a s es be th os e in w h om th e p ower o f s el f
,
p res erv a ti on is th e l ea s t it u n a v oid a bly f oll ows th a t th os e ,
l eft b ehi n d t o c onti nu e th e ra ce m u s t be th os e i n wh om th e ,
p ower o f s el f p res erv a t i on i s the g rea t es t mu s t be the s el ec t

- -
o f th ei r g en er a ti on S o th a t w h eth er the d a n gers t o exi s t

.
,
enc e be o f th e ki nd p ro d u ced by e xc ess o f f ertility or o f a ny ,
o th er ki n d it i s c l ea r th a t by th e cea s el ess ex ercis e of th e

,
f a cu lti es need ed t o cont end w ith th em a n d by the d ea th of ,
a l l m en wh o fa il t o con t en d wi t h th e m s u cce s s f u lly th ere i s ,
e ns u r e d a c on st a n t p r ogr ess t ow a r d s a high er d e gr ee of s kill ;
i n t ellig e n c e a n d se lf re g u l a ti o n— a bett er co ordi n a ti on of

- -
,
a c ti on s— a m ore c om pl et e lif e
*
.
§ 3 74 . Th e
p rop os iti on a t which we h a v e th u s a rr iv ed is .
th en th a t ex ces s o f fertility th r ou gh the ch a n ges it i s ev er

, ,
A g ood dea l o f th i s ch a pt er ret a ins i ts ori gi n l form ; and th e a bo v e a
p r a g ra ph i s re p ri nte d v e rb a t i m from th e U es tm i ns tcr Revi ew for A p i l

’
a r ,
1 8 5 2 i n whi ch th e v i e w s de v e l op ed i n th e foreg oi ng h u nd e d p age s w e r e

,
r
fi rs t s k et ch e d out Thi s p a ra gra ph sh ows h o w ne a r one m a y be t o

. a grea t
genera l i za ti on wi th out s e e i ng i t Thou g h th e s t r ugg l e for l ife i s the
. a l l eged
m oti v e force ; t hou gh th e p roc e s s of n a t ur a l s el ec ti on i s reco gni ze d ; a nd
though to i t i s a s c ri b ed a sh a re i n th e e v ol u ti on of a h igh er t y pe ; ye t the

conce p ti on i s not th a t wh i ch Mr . D a r w i n h as w or ked out w i th s u ch w onder
ful s kil l a nd I n th e fi rs t p l a c e, na tu ra l s e l ect i on i s h e re de
knowl ed ge .
sc ri b ed onl y a s furt h e ri ng d i rect a da p t a ti on— o nl y a s a i d ing p rog res s by t he

.
p re se rv a t i on o f i ndi v i dual s i n w hom functi onal l y-produce d m odi fica t i ons h a v e

H U MA N P OP U L A T ION I N T H E F U T UR E .
5 29
worki ng i n Ma n s envi ron m ent i s it s elf the c a u s e of Ma n s

’
,
’
fu rth er ev olu ti on ; a n d t he obvi ou s c oroll a ry h ere t o be d r a wn

i s th a t Ma n s f urth er e v ol u ti on so b rou ght a b ou t it se lf
’
, ,
n ec e ss it a t es a d ecli ne i n hi s f ertility .
A l l fu t u re p rog ress i n civiliz a ti on whi ch th e nev er

c ea s i ng p res s u re of p opu l a ti on mu s t p rod u ce will be a ecom ,
a n i e d by an e n h a n c e d c os t o f I n divid u a ti on b oth i n
p ,
s t ru c t u re a n d f unc ti on ; a nd m ore es p ec i a lly i n ner v ou s
s t r u c tu re a n d f un ct i on The p ea cefu l s t ru ggl e for exi s t en ce

.
i n s oc i eti es ev er gr owi ng m ore crow d ed a nd m ore c ompli

c a te d m u s t h a v e for its c on comit a n t a n i nc rea s e of th e gr ea t
,
ner v ou s c ent res i n m a s s i n com pl exity i n a ctivity ,

T ha t , .
l a rg er b ody of e m oti on needed a s a f ou nt a i n of energy for

m en w h o h a v e t o h old th ei r pl a ces a n d r ea r th ei r fa mili es
u n d er th e i nt en s i fyi n g comp et iti on o f s oci a l li fe i s oth er , ,
thi ngs equ a l th e c orrel a tive oi l a r ger b ra i n T h os e high er

'
, .
f eelin g s p res u pp os ed by th e b ett er sel f regu l a ti on whi ch i n -

,
a b ett er s oc i ety c a n a l one en a bl e th e i n divid u a l t o l ea v e a

,
p ers i s t ent p os t er ity a re other thi n g s equ a l the c or rel a tiv es

, , ,
o f a m ore c ompl e x b ra i n ; a s a re a l s o th os e m or e nu m erou s ,
m ore v a ri ed m ore genera l a nd m or e a b s t ra ct id ea s wh i ch

, , ,
mu s t a l so b ecom e i nc rea s i n gly requ i s it e for s u cc ess fu l life a s

s oci ety a dv a n ces A nd th e g enes i s of t hi s l a r ger qu a ntity o f
.
.
go ne on m os t fa v ou ra bl v I n th e s econd p l ace, th e re i s no t ra ce o f the i dea

.
th a t n a tu ra l s el ec t ion m a y by c o-op e ra ti on wi th th e ca us e a ss i gne d, or w i th
p roduce di verg ences of s tru ct ure ; a nd of cou rs e, i n th e a b s ence

ot h e r c au s e s .
of thi s i de a , th e re i s n o i m p l i c a t i on t h a t n a t u ra l s el ec ti on h as a nythi ng to do
w i th th e or i gi n o f s p ec i e s A nd i n th e th i r d pl a ce, t h e a l l i m port a nt fa ctor

.
-
of va i
r a t on i s p on ta n eo u s ,
”
or i nc i dent a l as we m ay oth e rwi s e cal l i t— i s
wh ol l y i gnor e d . Th ou gh I th i nk , m u c h m ore pot e nt
u se a nd di s us e a re,
ca u s e s of o rga ni c m o dific a ti on t h an Mr D a r wi n s uppos e s— th ou gh , wh il e p ur

.
su i ng t h e i n q ui ry i n de t a il I h av e b e en l ed to b e l i e v e th a t di rect e q uil i b ra ti on
,
h as p l a y e d a m ore a cti v e p a rt e v en th an I h a d m y s el f a t one t i m e t hought

ve t Mr D a rwi n to h a v e
I h ol d . s h own b evond q ues ti on , th a t a grea t p a rt of
the fa cts— perh a ps th e g re a te r p a rt
— a re e xp l i cabl e onl y as re s u l t i ng fro m
th e s ur v i v al of i ndi v i du al s wh i ch h a v e de v i a ted i n
i ndi rectl v-c au s ed wa y s om e
from th e a n c e s tral type Th u s , the a b o e p a ra g ra p h cont a i ns m erel y a p a s s

. v
v
i ng rec ogni ti on of t h e s e l e ct i e p roc e s s ; a n d i ndi ca tes no s u spi ci on of th e
e norm o u s r a nge of i t s e ffec ts , o r of th e cond i ti on s u nder whi ch a l a r e p a rt
g
of i ts e ffe ct s a re p rodu r ed .
80
5 30 L A WS OF MU L T IP L ICA TION .
feeli ng th ou ght i n a b ra in t h u s a u gm ent ed i n s i ze a nd

a nd ,
d ev el op ed i n st ru ct u re i s oth er thi ngs equ a l the c orrel a tive

, , ,
of a g r ea t er w ea r of n e rv ou s ti s su e a n d g rea t er c on s u m pti on
of m a t eri a l s t o r ep a i r it S o th a t b ot h i n or igi na l cos t of c on

.
stru c ti on a n d i n s u bs equ ent c ost of w or ki n g th e n erv ou s ,
s y s t em m u s t b ec o m e a h ea vi er t a x o n th e org a n ism A l rea dy .
the b r a i n of the civili zed m a n i s l a rger by nea rly thi rt y per

cent th a n t h e b ra i n o f th e s a v a g e
. A l rea dy t oo it p res ents .
, ,
a n i n c rea s ed h et er og eneity— es p ec i a lly i n th e di s t rib u ti on o f
i ts con v ol u ti ons A n d fu rth er ch an g es lik e th es e which h a v e

.
t a k en pl a c e u n d er t h e dis c ipline of civiliz ed lif e we i n fer ,
will c onti nu e t o t a k e pl a c e Bu t ev erywh ere a n d

.
a lwa y s ev ol u ti on i s a nt a g oni s tic t o p r oc rea tiv e di s s ol u ti on

,
.
Wh eth er it be i n gr ea t er growth of the orga n s which su b

s erv e s elf m a i nt ena nc e
-
w h eth er it be i n th ei r a dd ed c om
,
pl exi ty of st ru ct u re or wh ether it be i n th ei r hi gh er a ctivity

, ,
t he a b s t ra c t i on of t h e requ i red m a t eri a ls impli es a dimi

n i s h ed r es er v e of m a t eri a ls for ra c e m a i nt en a n c e A nd we -
.
h a v e s een rea s on t o b eli ev e th a t this a nt a gonism b et ween

I ndivid u a ti on a nd Genes is b ec om es u nu su a lly m a r k ed where
‘
t he ner v ou s s y s t em i s c onc erned b ec a u s e of th e c os tli ness of ,
n erv ou s s t ru ct u r e a n d f u n cti on I n § 3 4 6 wa s p oi nt ed ou t
.
t he a pp a ren t connexi on b et ween high c ereb r a l d ev el opm ent

a nd p r ol on g ed d e l a y of s ex u a l m a t u r ity ; a n d i n 3 66 3 67 ,
,
t he evid en c e w ent t o s h ow th a t wh ere exc epti on a l fertility

e xi s t s th ere i s s l u ggish n es s o f mi n d a nd th a t wh ere th e re ,
h a s b een d u ri ng edu c a ti on exc es s iv e exp en di tu re i n m ent a l

a c ti on t h ere frequ en tly f oll ows a c ompl e t e o r p a rti a l i n fer
,
t il i ty . H en c e the p a r ti cu l a r ki n d of fu rth er ev ol u ti on which

Ma n i s h erea ft er t o u n d erg o i s one which m ore th a n a ny , ,
ot h er ,
m a y be exp ect ed t o c a u s e a d ecli ne i n h i s p ower o f
rep ro d u cti on .
T he high er n erv ou s d ev el opm ent a nd gr ea t er exp endit u re

i n n erv ou s a cti on h ere d esc rib ed a s i n di rectly b rou ght a b ou t
,
by i n c rea s e of nu mb ers a nd a s th erea ft er b ec om i ng a ch eck

,
o n t he i nc r ea s e of nu mb ers m u s t n ot be t a k en t o imp l y
,
H U MA N P OP U L A TI ON I N T H E FU T URE .
53 1
an i nt enser s t ra i n— a m ent a lly l a b ori ou s life T he grea t er

-
.
em o ti on a l a nd i nt ell ec t u a l p ower a n d a c tivity a b ov e c on
tem pl ated m u s t be u n d ers t oo d a s be c omi ng by s m a ll i nc re

, ,
m en t s or g a ni c s p ont a neou s a nd pl ea s u ra bl e
, , ,
A s ev en w h en .
,
reli ev ed f r o m the p res s u re of n eces s ity l a rg e b ra i ned E u ro,

-
p ea ns v ol u nt a r ily ent er on ent erp ri s es a nd a cti viti es w hi ch

th e s a v a g e c ou ld n ot k eep u p e v en t o s a ti s fy u rgent w a n t s ;
s o th ei r s till l a r g er b r a i ned d es ce n d a n t s will i n a s till high er
,
-
,
d eg ree fi nd thei r gra ti fi ca ti ons i n c a reers ent a ili ng s till

,
g rea t er m ent a l exp en dit u res T hi s enh a nc ed d em a n d for

.
m a t eri a l s t o es t a bli s h a nd c a rry on the p s y chi ca l fun cti ons ,
will be a c on s tit u ti ona l d em a nd We mu s t c onceiv e the .
typ e gr a d u a lly s o m odifi ed th a t the m ore d ev el op ed nerv ou s

,
-
s y s t e m i rres i s tibly d r a w s off for i t s n orm a l a nd u nforc e d

,
a c tivi ti e s a l a r g er p r op orti on of th e c o mm on s t oc k of nu
. tr i
m ent ; a n d whi l e s o i ncr ea s i ng the i nt en s ity c ompl et enes s , ,
a n d l engt h o f th e i ndivid u a l life n ec ess a r ily di m i n i s hi ng the

,
reserv e a pplic a bl e t o th e s etti n g u o f ne w liv e s— n o l onger

p
requ i red t o be so nu m erou s .
T h ou gh th e w orki n g o f this p rocess will d ou btl ess be

i nt er fere d with a nd m odifi ed i n the fu t u re a s it h a s b een ,
i n th e p a s t by th e f a cilit a ti ons of livi ng whi c h c ivili za ti on

,
b ri n gs ; yet nothi ng bey on d t em p ora ry i nt err upti ons ca n s o

be c a u sed H owev er m u c h th e i nd u s t ri a l a rt s m a y be i m
.
p r ov ed th ere m u s t be a limit t o the i m p rov em ent ; w hil e

, ,
w ith a ra t e of m u ltiplic a ti on i n exc es s of th e ra t e of m or

t a lity p op ul a ti on m u st con ti nu a lly t rea d on th e h eel s of
,
p rodu cti on S o th a t th ou gh d u ri ng the ea rli er s t a ges of

.
,
c ivili z a ti on a n i ncrea sed a m ou nt o f f oo d m a y a c c ru e f rom a

,
g iv e n a m ou n t of l a b ou r th e r e m u s t c om e a ti m e w h en this
,
r el a ti on will be r ev ers ed a n d w h en e v ery a dditi on a l i nc r e

,
m ent of f ood will be obt a i ned by a m ore th a n p rop orti on a t e

l a b ou r : the di s p ro p orti on gr owi ng ev er high er a nd the dimi ,
nu ti on of th e re p rod u ctiv e p ower bec omi ng g rea t er .
§ 3 75 . T h ere n ow rem a n i s bu t to i nqu i re t owa r ds wh a t

5 32 L A WS OF M UL TI P L ICA TI ON .
li m it thi s p rogress t end s S o l ong a s the f ertility of th e

.
ra c e i s m ore th a n s u ffici en t t o b a l a n c e t h e di m i nu ti on by
'
d ea th s p op u l a ti on mu st c onti nu e t o i ncrea s e S o l ong a s

,
.
p opu l a ti on c onti nu es t o i ncrea s e th ere m u s t be p res su re on

,
the m ea n s o f s u b s i s t en c e A n d s o l ong a s th e re is p r ess u r e

.
on th e m ea n s o f s u b s i s t enc e fu rth er m ent a l d ev el opm ent

,
m u s t g o on a nd further dimi nu ti on of f ertility m u s t res u lt ;

,
p rovid ed th a t the a cti ons a nd rea cti ons whi ch h a v e b een ,
d es crib ed a re not a rt i fi ci a lly i nt erfered w ith I a pp en d this .
qu a li fyi n g c l a u s e a dvi sedly a nd es p ec i a lly emph a s i ze it be

, ,
c a u s e th es e a cti ons a n d rea cti ons h a ve b een hithert o a nd ,
a r e n ow ,
g rea tly i n t erfered with by g ov ernm ent s a nd the ,
c on ti nu a nc e of t h e i n t er feren ces m a y r et a rd if not s t op , ,
th a t fu rth er ev ol u ti on which w ou ld el s e g o on .
I refer t o th ose hi nd ra nces t o the s u rviv a l of th e fitt es t

whi ch i n ea rli er tim es resu lt ed f rom th e u ndis crimi na ti ng
ch a riti es of m ona s t eri es a n d i n l a t er times f rom the op era
ti on of P oor L a w s .Of c ou rse if th e c ompe ti ti on w hich
i nc rea s i ng p res su re o f p op u l a ti on ent a il s i s p revent ed f rom
,
acti n g on a c on s id er a bl e p a rt of th e c om mu n ity s u c h p a rt , ,
s a v ed f rom the neede d i n t ell ect u al a nd m ora l s t res s will not ,

'
u n d erg o a ny fu r th er m ent a l d ev el o pm ent ; a nd m u st ev er
t end t o l ea v e a p os t erity a n d a n i nc rea s i n g p os t erity i n

, ,
whi ch n on e o f th a t high er i n divid u a ti on which ch eck s

genes i s t a k es pl a c e S u ch S t a t e m eddlin g s with th e n a t u ra l
.
-
pl a y of a cti ons a nd rea cti on s p rod u ce a fu rth er evil eq u a l l y

grea t or g rea t er For th os e wh o a re fi ot s elf m a i nt a i ned or
.
-
,
bu t p a rti a lly s elf m a i n t a i n ed a re s u ppli ed with th e m ea n s

-
,
th ey l a ck bv th e b ett er m emb er s of the c ommu ni ty ; a nd

th ese b ett er m emb er s h a v e thu s n ot only t o s u pp or t th em
s elv es a n d th ei r off s p r i ng bu t a l s o t o s u pp ort or a id th e
,
i nferi or m emb ers a nd th ei r offs p ri ng T he u nd er worki ng of

.
-
o ne p a rt is a c c om p a ni ed by t h e ov er w orki n g o f the oth er

-
p a rt by a w orki ng whi ch a t ea ch s t a ge of p rog res s ex ceed s

-
th at whi ch th e n orm a l c onditi ons n ecessit a t e a nd resu lt s ,
s om etim es i n ill nes s p re m a t u re a g e or d ea th or i n l es s ened

, , ,
H U MA N P OP UL A T ION I N T H E F U T U RE .
533
nu mb er of c hild ren or i n i m p er fect rea ri ng of child ren : th e

, ,
ba d a r e f ost ere d a n d th e g ood a r e r ep r e s s e d .
I t d oes n ot foll ow t h a t th e s t ru ggl e for life a nd th e su r

vi va l of th e fi tt e st m u s t be l eft t o w or k ou t t h e i r e ffec t s
wi th ou t mitig a ti on I t i s cont end ed only th a t th ere s h a ll

.
n ot be a forc ibl e b u r d en i ng o f th e s u p eri o r for th e s u pp or t
o f th e in fer i or . S u c h a i d t o th e i n fer i or a s the s u p er i or

v ol u nt arily yi eld k ep t a s it will be wit h in m od era t e li m it s
, ,
m a y be giv en with b enefit t o b oth— re li ef t o t h e one m or a l ,
cu lt u re t o t he o th er A n d a i d willi n gly giv en ( li t t l e t o th e

.
l ea s t w orthy a nd m ore t o the m os t worthy ) will u s u a lly be

s o giv e n a s n ot t o f u rt h er t h e i nc r ea se o f th e u nw orthy .
F or i n p rop or ti on a s th e em oti ona l n a t u r e b ecom es m ore

ev olv ed ,
a n d th ere g ro w s u p a high er s en se o f p a ren t a l re
sp ons ibili t y t he b eg etti ng of child ren th a t c a nn ot be p rop erly

,
rea re d w ill be u niv ersa lly h eld i n t o l era bl e I f a s w e s ee

.
, ,
p u blic opi ni on in m a ny pl a ces a n d t im es b ecom es c oerciv e

en ou gh t o force m en t o fight d u e l s we ca n s c a r cely d ou bt th a t
,
a t a high er s t a g e of e v ol u ti on it m a y b ecom e s o coerc iv e a s t o
p rev en t m en from m a rryi n g imp rovid ently I f the f rowns .
o f t h e i r fe ll ow s ca n m a k e m en com m it imm ora l a ct s s u rely ,
th ey m a y m a k e m en ref ra i n fr om i m m ora l a cts—e s p eci a lly

wh en the a ct ors th em s e lv es f ee l th a t th e th rea t ened frown s
w ou ld be j u s tifi ed . H enc e with a high er m ora l n a t u re will
c om e a restri cti on on the mu ltipli ca ti on of the i n fer i or .
I n b r i ef th e s ol e r eq u i rem en t i s th a t th ere s h a ll be n o
,
ext en s iv e s u s p en s i on of th a t n a t u r a l r el a ti on bet w een m erit
a n d b enefit w hi c h con s tit u t es j u s ti ce H oldi n g th en th a t

.
, ,
th i s a l l ess enti a l c on diti on will it s el f c om e t o be rec ogni zed

-
a n d enfor ced by a m or e ev o lv e d hu m a n i t y l et u s con s id er

,
wh a t i s th e g oa l t owa rds whi ch the r es t ra i nt on g enes i s bv

i n dividu a ti on p rog resses .
S u pp os i n g th e S u n ’
3 75a . light a nd h ea t on whi ch a l l
s ,
t erres t ri a l li fe d ep en d s t o , con ti nu e a b u n d a n t for a p eri o d
l ong enou gh t o a ll ow the en ti re ev o l u ti on w e a re c ont em

5 34 L A WS OF MU L T I P L I CA TION .
pl a ting ; th ere a re s till c ert a i n ch a nges which mu st p rev ent

s u c h c o mpl et e a dj u s tm en t of h u m a n n a t u re t o s u rrou n di ng
c onditi on s a s w ou ld p er mit th e ra t e o f m u ltiplic a ti on t o

,
be com e equ a l t o the ra t e of m ort a li ty A s be f ore p oi nt e d ou t

.
1 4 8 ) d u ri ng a n ep och o f yea rs ea ch h emi s ph ere g oes

t hr ou gh a cycl e of t emp era t e s ea s ons a nd s ea s ons ext rem e i n
thei r hea t a nd c old— v a ri a ti ons whi ch a re th em s elves a lter
n a tel y ex a gge ra t e d a nd mitig a t ed i n t h e c ou r s e of fa r l ong er
c ycl es ; a nd we s a w th a t th es e c a u s e p erp et u a l ebbi ng s a nd
fl owin gs of sp ec i es ov er di fferent p a r ts o f th e E a rth s s u rf a ce

’
.
Fu rther by s l ow bu t i nevit a bl e g eol o gic ch a nges es peci a lly

, ,
th os e of el ev a ti on a nd s u bsid enc e the clim a t e a nd phys i ca l

,
ch a ra ct ers of ev ery h a bit a t a re m odifie d ; whil e ol d h a bit a t s

a r e d es t roy e d a n d new a re f or m ed T hi s t oo we n ot ed a s
.
, ,
a c ons ta n t c a u s e of migra ti on s a n d of re s u lti ng a lt era ti on s
o f en vi ronm ent N ow th ou gh th e hu m a n ra c e differs f rom

.
o th er ra ce s in h a ving a p ower of a rtifici a lly c oun t era ct i ng
e xt ern a l ch a n ge s yet th ere a re l imi ts t o t hi s p ower ; a nd

, ,
e v en w ere th ere no limits t he ch a n g es c ou ld n ot f a il t o w ork

,
'
th ei r eff ec t s i ndi rectly if not di rectly

, I f a s is th ou ght .
,
p rob a bl e th es e a s t ron omic cycl es ent a il rec u rrent gl a c i a l

,
p eri ods in ea ch h em i s ph ere then p a rt s of the E a rth which are

,
a t one tim e thickly p eopl ed will a t a noth er time be a lm os t

,
d es ert ed a nd vi ce vers é The g eol ogi ca lly c a u s ed a lt er a ti ons

,
.
-
o f clim a t e a n d s u r f a c e m u st p rod u c e f u rth er s l ow re di s tri

,
-
b u ti on s of p opul a ti on ; a nd oth er cu rrents of p eopl e t o a nd ,
f rom different regi ons will be neces s it a t ed by the r is e of su c

,
c es si ve c ent res o f high er civi li z a ti on C ons equ ently m a n .

,
ki n d c ann ot bu t c onti nu e t o u nd erg o c h a nges of envi ron

m en t phys ic a l a n d m ora l a na l ogou s t o th os e which they
, ,
h a ve thu s far b een u nd erg oi ng S u ch ch anges m a y eventu

.
a lly be c om e s l ow er a nd l es s m a r k ed ; bu t th ey c a n nev er
c ea s e A nd if th ey ca n n ev er c ea s e th ere can n ev er a ri se a
.
p er fect a d a pt a ti on of hu m a n na tu re t o i ts c onditi on s of e xi s t
e n ce . T o e st a blish th a t c ompl et e c orr esp ond en c e be t ween
i nner a nd ou t er a cti ons which c onstit ut es the hi ghest life a nd
H UMA N P OP U L A T I ON I N T H E F UT U RE .
535
g rea t est p ow er of s elf p res erv a ti on th ere mu st be a p rol onge d

-
,
c on vers e b et ween the org a ni s m a nd ci r cu m st a nc es which re

m a i n the s a me I f the ext erna l rel a ti ons a re b ei ng a lt ered
.
whil e the i nt erna l rel a ti on s a re bei ng a dj u s t ed to th em the ,
a dj u st m en t ca n n ev er bec o m e e x a c t A n d in the a bs enc e of

.
ex a ct a dj u stm ent t h ere c a nn ot exi s t th a t th eoretic a lly high e s t

,
-
p ower of s elf p res erva ti on with w hich th ere w ou ld c c exi s t

- -
t he th eoretic a lly l owest p ower o f r a ce—p rod u cti on

-
.
H en c e th ou gh th e nu m ber of p re m a t u re d ea th s m a y u lti
m a t ely be com e v ery s m a ll it ca n nev er b ec om e s o s m a ll a s
,
t o a ll ow the a v era ge nu m be r of offsp ri ng f ro m ea ch p a i r t o

f a ll s o l ow a s two S om e a v era ge nu mber b et w een tw o a nd
.
th ree m a y be i nf erre d a s th e limit— a nu mb er h owe v er w hich , ,
i s n ot lik ely t o be qu it e c ons t a n t bu t m a y be e xp ect ed a t

,
on e ti m e t o i nc r ea s e s o m ewh a t a n d a ft er wa r d s t o d ec r ea s e
s o m ewh a t a cc or di n g a s v a r i a ti on s i n phy s i ca l a nd s oci a l con

,
di ti ons l ower or r a i s e th e c ost of s elf p res er v a ti on -

.
T o thi s qu a lific a ti on m u st be a dd ed a fu rth er qu a l i fi ca

ti on. T he for eg oi n g a rg u m en t t a citly a s s u m es th a t th e
c a u s es d es crib ed will c ontin u ou s ly ope ra t e on a l l m a nki nd ;
wh erea s a s u rv ey o f the f a ct s m a k es it cl ea r th a t s om e p a rt s
’
on ly o f t h e E a rth s s u r f a c e a r e ca p a bl e of b ea r i ng h igh
typ es of civili z a ti on a n d c on s eq u ently high typ es of Ma n

, .
T h ere m u s t rem a i n h erea ft er a s t h ere a re n ow con s id era bl e

, ,
p a rt s of i ts s u rfa ce whi ch can su pp ort only g rou p s of n om a d s ,
or oth er g r ou p s o blig ed by th ei r h a bit a ts t o l ea d s i m pl e
a nd i n f eri or ki nd s of l if e On ly by s u bj ecti on t o th e di s ci
.
pli ne we h a ve b een c ont empl a ti ng c a n th ere be p r odu ced the

f u lly d ev el op ed Ma n ; a nd evid ently i n m a ny p a rt s of the
-
w orld thi s disciplin e will c onti nu e t o be el u d ed N ot only .
mu st w e c oncl u d e th a t th e v ari e ti es of ou r ra c e n ow liv

i ng i n d eser t r egi on s a nd a rctic clim a t e s w ill c on ti nu e h er e
a ft er t o do so bu t we m a y c onc l u d e th a t a l wa y s a s now a
, , ,
c ert a i n p rop orti on of m en wh o a re b orn i n civili zed s oci eti es ,
i mp a ti en t of th e str es s whi ch p r essure of p op ul a ti on p u t s

on th em wi ll esc a p e i nt o u noccu pi e d or sp a rs ely p eopl ed -
,
530 L A WS OF M U L T I P L I CA TI ON .
re gi on s wh ere they c a n l ea d u n res tra i ned liv es th ou gh lives

,
o f h a rdshi p Recogn i z i ng a s we mu s t the p rob a bility th a t

.
i n c omm on with a l l oth er thi ng s h u m a n ity w ill c onti nu e t o

,
di fferen ti a t e a nd p rod u ce a m ore h et er ogeneou s a ss embl a ge

o f typ es w e m u s t i n f er th a t on ly i n s om e o f t h e high es t o f
,
t h es e w ill th e a nt a g on i s m of i n divid u a t i on a n d genes is h a v e

th e a ntici
B es t r i t o th es e th en we m a y c onc l u d e
, ,
th a t i n of p op u l a ti on a n d i t s a ccomp a ny
i ng evils will a lm os t dis a pp ea r ; a n d will l ea v e a st a t e of

things requ i ri ng fr om ea ch th a n a
n orm a l a nd d ecrea s e
o f f ertility i m pli es c es s a ti on i n the d ev el opment of the
n er v ou s s y s tem ; a n d thi s impli es a ner v ou s s yst em whi ch
h a s b ecom e equ a l t o a l l th a t is d em a n d ed of i t— h a s not t o do

m or e th a n i s n a t u ra l t o it Bu t th a t e x er cis e of fa cu lti es which
.
d oes n ot ex ceed wh a t i s na t u ra l c ons tit u t es gr a tifi ca ti on

, .
Th e n ec ess a ry a nt a g onis m of I n divid u a ti on a nd Genes i s .
n ot only th en f u lfil s the a p ri ori l a w of m a i nt ena nce o f

, ,
ra c e f ro m th e m on a d u p t o Ma n bu t en su res fi n a l a tt a i nm en t
, ,
o f th e high es t for m of thi s m a i nt en a n ce —a f or m i n whi c h
t he a m ou nt of lif e s ibl e a n d th e
bi rth s a nd d ea th s the fe th e b egi nni ng

p ressu re of p opu l a ti on h c a u s e of p r o
g res s . I t p rod u ced th e origi n a l diff u s i on o f the r a ce It
.
c om p ell e d m en t o a b a n d on p r ed a t ory h a bit s a nd t a k e t o
a g r ic u lt u re I t l ed t o th e cl ea ri ng o f th e E a rth s su r fa ce
.
’
.
I t forced m en i n t o th e s oci a l s t a t e ; m a d e s oc i a l org a ni za

ti on i ne vit a bl e ; a nd h a s d ev el op ed the s oci a l s entim ents I t .
h a s sti m u l a t ed t o p rog res s iv e imp rov em ents i n p ro d u cti on ,
a n d t o i n c rea se d s kill a n d i nt ellig en c e I t i s d a ily th ru s ti n g

.
u s i n t o c l oser c ont a c t a n d m or e m u t u a lly d e p en d en t re l a ti on

-
s hip s . A n d a ft e r h a vi n g c au s ed a s it u ltim a t ely m u s t th e

, ,
du e p eopli n g o f the gl ob e a nd th e ra i s i n g of its h a bit a bl e

,
p a rt s i nt o the high es t st a t e of cu ltu re —a ft er h a vi ng p erfect ed

a l l p roces s es for t h e s a ti s f a cti on of h u m a n wa nt s— a ft er
H U MA N P OP U L A T I ON I N T H E F U T U RE .
53 7
h a vi n g ,
at th e d ev el op ed th e i nt ell ect i nt o com
sam e tim e,
e t en ce fo r i t s w or k a n d th e f eeli ng s in t o fit n es s for s oc i a l
p ,
li fe —a ft er h a vi n g d one a l l th i s th e p res su re o f p op u l a ti on
,
mu s t gra du a lly a pp r oa ch to a n en d— a n end h owev er whi ch

w
, ,
for the rea s on s giv en i t ca nn ot a b s ol u t ely rea ch j
§ 3 77 . In l i g the a r gu m ent l et u s n ot ov erl ook the

c os n
s elf s uffi ci n g n es s o f th os e u n iv er sa l p ro ce s s e s by w hic h th e

-
res ul t s r ea c h e d th u s fa r h a v e b een w r ou ght ou t a n d whi c h ,
m a y be exp ec t ed t o w or k ou t th es e fu t u r e r esu l t s .
E v o lu ti on u n d er a l l i t s a s p ec t s g ener a l a n d s p ec i a l i s a n
, ,
a dv a n ce t owa r ds e qu ilib r i u m We h a v e s een th a t the theo

.
reti ca l li m it t ow a r d s wh ich th e i n t e gr a ti on a n d di ffer enti a
ti on of ev ery a ggrega t e a dv a nce s i s a s t a t e of b a l a n ce be,
t ween a l l the forces t o w h i ch i t s p a rt s a re su bj ect a nd the ,
f or ces whi ch i ts p a r t s opp os e t o t h em ( F i rs t P r in .
A n d w e h a v e s een th a t org a ni c e v ol u ti on is a p r o gr es s t ow a r d s
a m ovi ng equ ilib r iu m c om pl et ely a dj u st ed t o envi roni ng
a cti on s .
I t h a s b een a l s o p oi nt ed ou t th a t i n civili zed Ma n th ere is , ,
g oi ng on a n ew cl a s s of equ ilib ra ti ons— th os e b et ween his a c

ti on s a n d the a cti ons of the s oci eti es h e f orm s ( F i rs t P ri n .
S oci a l r est r a i nts a n d r e q u i r e m ents ar e e v er a lt eri n g

hi s a c t iviti es a n d by c on s equ en c e hi s n a t u r e ; a nd a s fa s t a s h i s
na t u r e i s a lt ered s o ci a l r es t r a i nt s a n d r eq u i r em en t s u n d er g o
,
m ore o r l es s re a dj u s t m en t
-
H er e t he orga n i s m a nd t h e c on
.
d i ti on s a re b oth m o difi a bl e ; a n d by su c c es s iv e conc ili a ti ons

o f th e t w o th er e i s effect e d a p r ogr es s t ow a r d s equ ilib r iu m
,
.
M ore recent ly w e h a v e s een th a t i n ev er y s p eci es th ere ,
e s t a bli s h e s it s elf a n eq u ilib r i u m o f a n i n v olv e d ki n d be t ween
th e t ota l r a ce d e s troyi ng f orces a n d th e t ot a l ra c e p r eser vi n g

- -
f orc es— a n equ ilib ri u m whi c h i m pli es th a t wh ere the a bility

t o m a i n t a i n i n divid u a l li fe i s s m a ll t he a bility t o p r op a ga t e
,
mu s t be g rea t a n d p i ce vers ei Wh en ce i t foll ow s th a t the

,
.
ev o l u ti on of a ra c e m or e i n equ ilib r i u m w ith the envi r on

m ent is a ls o th e e v ol u ti on of a r a c e i n w hich th er e i s a cor
,
~
538 L A WS OF MU L TI P L I CA T I ON .
l tive
re a a pp roa c h t owa rds equ ili b riu m b et ween the nu mb er
o f new i n divid u a ls p ro d u ce d a nd t h e nu mb er whi ch s u r viv e
a n d p rop a g a t e
.
T he fi n a l res u lt t o be ob serv ed i s th a t i n Ma n a l l th ese

,
e qu ilib r a ti on s be t we en c on s tit u ti on a n d con di ti on s b et ween

,
th e s tru ct u re o f s oci ety a n d th e n a t u re of i t s m em b ers ,
bet ween f ertility a nd m ort a lity a dv a n ce si m u lt a neou s ly

,
t oward s a c omm on clim a x . I n a pp r oa c hi n g a n e qu ilib r i u m

bet ween hi s n a t u re a nd the ev er v aryi ng ci rcu m s t a nc es of hi s
-
i norg a nic envi ron m ent a nd in a pp roa ching a n equ ilib riu m
,
b et ween hi s na t u re a nd a l l the requ i rem ent s o f the s oci a l

s t a t e Ma n i s a t th e s a m e tim e a pp r oa chi n g th a t l owe s t li m it
,
of f ertility a t w hich t he equ ilib r i u m of p op u l a ti on i s m a i n
t a i ned by the a dditi on of a s m a ny i nfa nt s a s th ere a re su b

t ra cti ons by d ea th i n ol d a g e B u t i n a u niv ers e of which a l l
.
p a rt s a re i n m oti on a n d ev ery p a rt is c ons equ ently s u bj ect t o

ch a nge of c on diti on s neith er thi s equ ilib ri u m nor a ny oth er
,
equ ili b ri u m ca n b ec om e c o mpl et e .
TH E E ND .
A P P E ND IX A .
SUBSTITUTI ON OF A X IAL FOR FOL I AR ORGANS I N P L A NTS .
I h ere th e e v id ences r eferre d to i n

A P P E ND 1 90 T h e m os t .
nu m ero u s a nd s t riki ng I ha v e m e t w i th a m ong th e Um bel l ifer ce .
Mo ns tro s i ti es h a v i ng t h e a lleg e d i m pli c a ti on, a re fre qu e nt i n th e

c o m m o n C o w P a rs ne p— s o fre qu e nt t h a t t h e y m u s t be fa m ili a r t o
-
b ot a ni sts ; a nd w i l d A ng eli c a s uppli es m a ny o v er d e v el op m e nts of

-
lik e m e a ni ng Om i tti ng nu m erous c a s es of m ore or l ess s i gnifica nc e ,

.
I wi l l li m it m y s elf t o tw o .
One of t h e m i s th a t of a t erm i na l
b el i n w hi ch ni ne of th e
um ,
o u te r u m bellu l e s a re v a r i o u s ly t ra ns form e d — h e re a s i ngl e fl o wer
b ei ng m a de m o ns t ro us by th e d e velopm e nt of s om e of i ts m em b ers
i nt o bu d s ; t he re s e v era l su ch m alfo rm e d flo we rs b ei ng as soc i a te d
w i th ray s t hat b ea r i m p e rfe ct u m b ellules ; a nd els e wh e re flo we rs ,
b ei ng re pl a ced by u m b ellules s om e o f wh i ch are p e rfe ct and oth ers ,
i m p e rfe c t o nly i n t he s h ort ne ss o f th e fl ow er s talks Th e a nne x e d

-
.
Fig 6 9 re pres enti ng i n a s o m e wha t conv ent i onali ze d w a y a part o f

.
, ,
5 41
5 42 A P P E ND I X A .
th e d ri e d s p ec im en w i ll gi v e a n id ea of thi s A ng eli ca A t a i s
, .
s h o wn a s i ngl e flow e r p a rt i a lly c h a ng e d ; i n th e u m b ellul e m ark e d
6 o ne o f th e ra y s b ea rs a s e co nd ary u m b ellule ; a nd t he re m ay
,
be s e e n a t c a nd ( 1 s e v e ral s u ch o v er d e v el op m e nts
,
- .
B ut t h e m os t conclu s iv e i ns ta nc e i s th a t of a C o w P a rs ne p i n -
,
w h i ch a s i ngl e t erm i nal u m b el b es i d es th e t ra ns form a t i o ns a lrea dy

,
m e nti o ne d e x h ibi ts h igh er d egrees o f s u ch t ra ns fo rm a t i ons

*
, The .
c o m p on e nts of th i s c o m l e x g row th are —th re e c ent ra l um b e ll ul es

p ,
a b no rm a l o nly i n m i nor o i nts ; o ne u m b ell ul e e xtern a l t o th es e

p , ,
w h i ch i s p a rt i ally ch a ng e d i nt o a n u m b el ; one ra th e r m ore out of th e

c e nt re w h i c h i s s o fa r m et a m orp h o s e d as t o b e m o re a n u m b el th a n
,
a n u m b e ll ul e : ni ne p e riph era l c lu s t e rs form e d by th e d e v el opm e nt
o f u m b ellu l es i nto u m b el s s om e of wh i c h a re p a rt i ally c o m p o u nd e d

,
s til l fu rt h e r E x a m i ne d i n d eta i l th es e s truc t u re s p res e nt th e fol

.
,
l owi ng fa cts — 1 The i nnerm os t um b ellul e i s nor m al s ave i n h a v i ng

.
,
a p e riph era l fl o we r o f w hi ch o ne m em b er app a re ntly a p e ta l

( ) si
tra ns form e d i nt o a fl o we r bu d 2 Th e ne x t u m b ellu l e not qu i te s o
-
. .
,
ce ntra l h a s o ne of i ts p e riph era l fl o we rs m a d e m ons t ro us by t he

,
g rowth o f a bu d from t he b as e of the c a lyx 3 T he thi rd of th e . .
c ent ra l u m b ellu l es h a s t w o a b norm a l o u t e r flo w ers One o f th em .
c a rri es a fl o w er bu d o n i ts e dg e i n pl a c e o f a foli a r m em b er
-
, .
Th e oth e r i s h al f flow er a nd ha lf um b ellu l e : b e i ng c om p os e d o f

t h ree p e t al s th re e s t a m ens a nd fi ve fl ow er b u d s gr ow i ng w h e re
, ,
-
t he oth er p et a ls a nd s t a m e ns s h ou ld g row 4 Ou ts id e of th es e . .
u m b ell ul e s c o m es o ne of th e m i x e d c l u s te rs Its fi ve ce ntral ,
fl ow e rs a re norm al Su rrou ndi ng th es e are s ev e ral flow ers tra ns

.
form e d i n di ffe re nt d e grees : o ne h a vi ng a s t am e n p a rt i ally ch a ng e d

i nto a flow er bu d A nd th en a t t he p eriphe ry of th i s m i x e d
.
,
c l us t er c om e thre e com pl e te u m b ell ul es a nd a n i ncom pl e te o n e i n

,
whi ch s om e p e t a ls a nd s ta m ens o f t he origi na l flow e r re m a i n .
5 A m i x e d cl us t e r i n wh i c h th e um b e l s tru c tu re p re d om i na t es
.
,
-
,
s ta nd s ne x t Its th re e c ent ra l flow ers a re norm a l: S urr ou ndi ng

.
t he m a re fi ve fl ow ers o v er d e vel op e d i n v a ri ous w a y s lik e t hos e

-
,
a l rea d y d e s c rib ed A nd on i ts p eriph e ry a re s e v en c om pl e te u m

.
bel l u l e s i n pl ac e of fl ow ers ; b es id es a n i nc om pl ete u m b ellu le that

c o nt a i ns t ra ces of t he o rigi na l fl o w e r o ne o f t he m b ei ng a p eta l ,
i m perfe ctly twi ste d up i nto a bu d 6 Of th e ni ne e x terna l . .
cl u s ters i n w hi c h th e d e v el op m e nt o f s i m pl e i nt o c om p o u nd u m
,
b els i s m os t d e cid ed nea rly a l l pres ent a nom a li es Three of them

,
.
h a v e e a ch a c e ntral fl ow e r untra ns form e d a nd i n oth ers th e c entra l ,
For t he i nform at i on of th ose who m a y wi sh to ex a m i ne m et am orph os es

o f t h e s e k i nds I m a y h ere st at e th a t I h a v e fou nd ne a l y a l l t he e x a m pl es
,
r
de s c ri b ed, i n th e nei gh bou rt of t h e s ea — th e l a s t-na m e d, o n th e s h ore of
L oc h e i l, nea r Fort W i l l i a m Whether i t i s tha t . h a e s ought m ore di l i I v

n t l for ca se s wh en i n s uch l oc a l i t i e s , or w h eth er i t i s t h a t th e s e a -ai r
g e y
v x v
fa ours th a t e cessi e nut ri t ion wh ence the se t ra ns form a t ions re s ul t, I a m
un a bl e t o s ay .
S U BS TIT UT I ON OF A XIA L F OR FOL IA R ORGA NS .
543
um b ell ul e is c om p os ed of t wo , three , or four flow e rs . 7 B ut th e

.
m os t r em a r a k bl e fa ct i s d ry
,
t ha t i n sun of th es e p e riph era l cl u s te rs ,
res ul ti ng fro m th e m e ta m orp h os i s of s i m pl e u m b e l s i nt o com p ou nd
u m b el s th e lik e m et a m orph os i s i s c arr i e d a s ta g e h igh e r

,
S om e of .
t he com po ne nt ray s ar e th em s e lv es the b eare rs of c om p ound u m b els

,
i nste a d of s i m ple u m b els In Fig 7 0 a po rt i on of th e d ri ed s p e ci

. .
,
m e n i s re pres ente d T w o of th e cent ra l u m b ellul es are m a rk e d a

.
a nd b th ose m ark e d c a nd d a re m i x e d cl u s te rs ; a t e a nd f are

c om o u nd u m b el s repl ac i ng s i m l e o nes ; a n d 9 sh o w s o ne o f t he
p p
ra y s o n wh i ch th e o v er d e v el op m ent g oe s s ti l l furt h er
-
.
D oes not th i s e vid e nce en force d as i t i s by m u ch m ore of li k e

,
ki nd g o far t o p rove th a t foli a r org a ns m ay be d e veloped i nto a x i a l

,
o rg a ns E ve n w ere not the tra ns it i ona l form s tra ce a ble th ere w ould ,
s t ill I th i nk be no oth er l e i ti m a te i nt e rp reta ti o n of th e fa cts l as t

, , g
d e t a ile d T he only w ay of c lu di ng t he con clu s i on h ere dra w n i s by
.
,
as s um i ng th a t wh ere a c l u s te r o f fl ow ers r epla c es a s i ngl e flo w e r, i t
i s b ec au s e th e a xill a ry bu d s w hi ch hyp oth eti cally b elo ng t o th e

s e v e ra l foli ar org a ns of th e flo w er b e co m e d e vel o e d i nt o a x es a nd
p
,
as s um i ng t hi s i s b as i n a n h yp oth es i s o n a noth e r h yp oth es i s th a t i s

,g
di re ctly a t v a ri a nce w ith fa cts The foli a r orga ns of flow ers do no t
.
b ea r bu d s i n th ei r a xil s ; a nd i t w ould ne v e r h a v e b e en supp os e d

t h a t s u c h b u d s a re ty pi ca lly p re s e nt h a d i t not b een for t ha t
,
m i s t a k e n c oncepti on of typ e w h i ch h as l e d to m a ny oth er erro rs

i n B i olo gy G o ethe w ri tes :
. Now as w e c a nnot re ali z e th e i d ea
“
o f a l e a f a p a rt from th e no d e ou t o f w h i ch i t s p r i n s o r o f a no d e
g ,
”
w itho ut a bu d w e m a y v e ntu re to i nfe r
, &c S e e h e re a n e x a m pl e
, .
o f a m eth o d o f h il os op hi zi n
p g not unco m m o n a m o ng the Ge rm a ns .
5 44 A PP E N D I X A .
Th e m e th o d i s t h i s — S u rv e yp ort i on o f th e fa cts a nd d ra w from

a ,
th em a g enera l c onc e p t i on ; p roj e c t th i s g enera l c oncep ti o n b a c k

i nto th e o bj ecti ve w o rld as a m oul d i n w hi c h Natu re c a s ts h er
,
p ro du cts ; e xp e ct to fi nd i t e v ery w h ere fulfill ed ; a nd a ll eg e p ote n

t i a l fulfil m e nt w h ere no a c t u a l fulfil m e nt i s v i s ibl e .
I f i ns t ea d of i m p os i ng our id ea l fo rm s o n Na tu re w e a re c on ,
t e nt t o g ene ra li z e th e fa ct s a s Na tu re p res e nts t hem w e s h a ll fi nd ,
no w a rra nt for th e m orp h ol og i c a l d oc tri ne a b o v e e nu nc i a te d Th e .
o nly c o nc e p ti o n of typ e j us tifie d by t h e l ogi c o f s ci e nc e i s — t h a t ,
c o rre l a t i o n o f p a rt s w h i c h rem a i ns c o ns t a nt u n d er a l l m o difi ca ti ons
o f t h e s t ru c tu re t o be d efi ne d T o as c e rta i n th i s w e m u s t co m p a re
.
,
a l l t h es e m o dific a ti o ns a nd no t e w h a t t ra i ts a re c o m m o n to th em
, .
O n d oi ng s o w i th t h e s u cc e ss i v e s egm e nts o f a ph aenoga m i c a x i s ,
w e a re brou ght t o a c o ncl u s i on w id ely di ffere nt from t ha t of Go et he .
A x ill ary bu d s a re a l m os t u ni v ers a lly a b s ent from th e c otyl ed o ns ;

t hey a re h a bi tu a lly p r es ent i n t h e a x ils of fully d e v el op e d l ea v e s -
h igh er u p th e a x i s ; t he y a re o ft e n abs e nt from l ea ve s th a t a re cl os e

t o t h e flow e r ; t hey a re nearly a l w a y s a bs e nt from th e bra c t s ; a bs ent
fro m th e s ep a ls ; a b s e nt from t he p et als ; a b s e nt fro m t he s ta m e ns ;
a b s e nt from t he c a rp e l s Thu s o ut o f e ight le a di ng fo rm s w h i c h
.
,
foli a a s s u m e o n e h a s th e a x ill a ry bu d a nd s e v en a re w i thou t i t

, .
W i th th es e fa cts b efore u s i t s eem s to m e not di ffi cult to rea li ze

,
” ”
t he id e a of a n o d e w i th out a bu d If we a re not p os s e s s e d
.
by a foreg o ne c onclu si o n th e e v id e nc e w ill l ea d us t o i nfer t ha t

, ,
e ach n o d e b e a rs a fo li a r a pp e nd a g e a n d m a y b ea r a n a x ill a ry bu d .
E v e n ho we v e r w ere i t gr a nte d t ha t th e typi c a l s e gm ent of a

, ,
P h aenog a m i nclu d es a n a x ill a ry bu d wh i ch m us t b e rega rd e d a s ,
a l w a y s p o te nt i a ll y p re s e nt no l e gi t i m a t e c ou nt er i nt erp ret at i o n of
,
-
th e m ont ros i ti e s a b o ve d es c rib e d c ould th e nc e b e d ra w n I f w hen .
a n u m b e llul e i s d e v el op e d i n pl a c e o f a flow er t he e x pl a na ti on i s , ,
th a t i ts c om p one nt ray s a re a x illa ry t o th e foli a r o rg a ns of th e

flow er s up ers e d ed w e m a y fai rly re qui re that thes e foli ar orga ns to
w h i c h th e y a re a x ill ary s h a ll b e s h o w n
,
B ut th e re a re none I n
. .
th e l as t s p e c i m en figu re d t h e i nne r ra y s of e a c h s u ch u m b ellul e a re

,
w i th ou t th e m ; m os t o f t h e ou t e r ra y s a re a ls o w i th ou t t he m ; a nd
i n o ne clu s te r o nly a s i ng l e ray h a s a b ra ct a t i ts p o i nt of o rig i n
,
.
T here i s a rej oi nd er rea dy h o we v e r : th e foli a r orga ns a re s aid t o

,
”
be s u pp re s s e d . Th ou g h Goeth e c ould not re a liz e th e id ea of
“ “
”
a no de w i th ou t a bu d t h os e w ho a c ce p t h i s typi c a l form a pp ear to
,
fi nd no d i ffi culty i n r eali z i ng th e id e a of a n a xill a ry bud w i th out

a ny th i n t o wh i c h i t i s a x ill a ry B u t l etti ng th i s p ass s u pp os e w e
g .
,
ask w h a t i s th e w a rra nt for th i s a s s u m e d s u pp res s i o n A xill a ry .
b ud s no rm ally o c cur w he re th e nu tri ti on i s high e nough to p ro du ce

fully d e v el op ed l e a v es ; a nd wh e n a xill a ry b u d s a re d em ons tra bly
-
re s e nt i n fl o w ers t h e y a c com a ny foli ar org a ns th a t are m o re l ea f

p , p
lik e t ha n u s ua l— a l way s g re e ne r i f not a l way s l a rg er That i s t o .
S UBST IT U T I ON OF A XI AL . FOR F OL IA R ORGA NS .
5 45
s a y, th e norm a l b norm al a x illary bu ds a re a lik e the c on

an d th e a ,
c o m i t a nts o f foli a r o rg a ns c ol ou re d by th a t chl orop h yll w h i c h
h a bi t u a lly fa v ou rs foli a r d e v el op m e nt H o w th e n c a n i t b e s u p
.
, ,
o s e d t h a t w h e n o u t o f a fl o w e r t h ere i s d e vel o e d a c lu s te r o f
p , p
fl ow e r be a ri ng ra ys th e i m pli e d e x cess of nut ri ti on c a us es th e foli ar
-
,
o rg a ns t o a bo rt I t i s t ru e t h at v e ry g e nera lly i n a bra nc h e d i n

fl ores ce nc e t he b ra cts of th e s e v era l fl o we r b ra nch e s a re v e ry s m all
,
-
e i r s m al l nes s b e i ng p ro b a bly du e t o th a t d efe ct i v e s u pply o f

( t h
c e rt a i n c hl o ro p h yll form i ng m a tte rs w h i ch i s th e p ro xi m a t e c a u s e
-
,
o f fl o w eri ng a nd i t i s t ru e th a t u n d er th es e con di t i ons a fl o w e r

) , ,
i ng a x i s of c o ns id era bl e s i ze for th e d e vel op m ent o f w h i c h chl oro

,
ph yll i s less ne ed ful gro ws from th e a xil of a d warfe d l e af B ut

,
.
t h e i nferen ce t h at t h e foli ar o rg a n m a y th e refore be e nti rely s u p

p res se d s eem s to m e i rr e co nc i l a ble with the fa ct th a t th e foli ar
, ,
o rg a n i s a lw a y s d e v e l op e d to s om e e x tent befor e th e a xill ary bu d
a pp ears U nt il i t h as be en s h o w n tha t i n s om e cas es a l at era l bu d

.
fi rs t a pp e ars a nd a foli a r org a n after wa rds g ro ws out be nea th i t t o

, ,
form i ts a x il t h e c onc ept i o n o f a n a x illa ry bud of w h i ch th e foli a r

,
o rg a n i s s u pp res s e d w ill rem a i n a t v a ri a nce w i th th e es ta bli s he d

,
t ru t hs of d e v el opm e nt .
Th e a b o v e origi nally form e d a p o rt i on of 1 9 0 I h a ve tra ns .
fe rre d i t t o th e A pp endix , p a rt ly b eca u s e i t c o nta i ns t oo m u ch

d eta il to re nd er i t fit for t he g ene ra l a rg um ent, a nd p a rtly b e ca u s e
t h e i nt erp r et a t i o ns b ei ng o p e n t o s om e qu est i on , i t s e em e d u nd e
s i ra bl e t o ri s k c o m p rom i s i ng th a t a r u m e nt by i nc lu d i ng t h em
g .
T h e c ri t i ci s m s p as s e d u p on t hese i nt e rp re t a ti ons h a v e not , h ow

e v er, s u fli c e d to c on v i nc e m e of t h e i r i n c o rre c t ness U nfort u .
na t el y, I h a v e s i nc e h a d no o pp o rt u ni ty o f v e ri fyi n th e abov e
g
s t a t e m e nts b
y m i cros co pi c e x a m i na ti o ns , as I h a d i ntend e d .
T h ou g h u n a bl e to e nforc e th e i nfe renc e d ra wn by fu rth er fa cts

m or e m i n u t ely l ook e d i nt o , I m a
y a dd s om e a rgum ents b a s e d o n
fa cts th a t a re w ell k now n One of th es e i s th e fa ct th a t th e s o
.
ca ll e d a x ill a ry bu d i s not u ni v ers a lly a x ill a ry — i s n ot u niv ers a l l y
s e a t e d i n t h e a n l e m a d e by th e a x i s a n d a n a p e nd e d fo li a r o rg a n
g p .
I n c e rta i n pl ants t h e a x illa ry bud i s pl a c e d far a b o v e th e n o d e,

h a lf wa y be t we en i t a nd th e s u c c ee di ng n o d e S o th a t n ot o nly m a
-
y.
a s e gm e nt o f a p h aenog a m i c a x i s be w i th o ut th e a x ill a ry bu d bu t
,
th e a x il l a ry bu d, wh en p res e nt , m a y b e rem o v e d from th a t pl a c e i n

w hi ch , a c cordi ng to Go eth e, i t ne c ess a ril y e x i s ts A nother fa ct no t .
c o ng ruo u s w i th th e c u rre nt d oc tri ne , i s th e c o m m on o ccu rre n c e of
a d v e nt i t i o u s b u ds — th e buds t ha t a re put out from roots a nd fr om

o l d s t em s o r b r a nch es b are of l ea v es Th e na m e u nd er w h i ch th ey are
.
th u s cl a s s ed i s m eant t o i m ply tha t th ey m ay be left out of c ons id e

,
ra ti o n . Those , h o we v e r, w ho h a v e not g ot a theor y t o s a ve by

81
546 A P P E ND I X A .
putti ng a nom a li es out of s ight i ncline d to th i nk th at the

, m ay be
o c c u rre nc e o f bu d s w h e re th e y a re a v o w e dl y u nc onn e c t e d w i th
no d es a nd a re a x ill a ry t o noth i ng te l l s v ery m u c h a g a i ns t t h e a s

, ,
s u m p t i o n t h a t e v e ry bu d i m pli e s a no d e a nd a c or re s p o ndi ng fol i a r
o rg a n .A nd t h ey m ay a ls o s ee th at th e d e v el op m e nt of t hes e a d
ve nt i t i ou s b ud s a t pl a c es w h ere th e re i s e x c es s o f nu tri ti v e m a t e
ri al s fa v o u rs t h e v i e w a bo v e s e t fo rt h
,
For i f a bu d thu s a ri s es a t
.
a pl a ce w h e re i t i s not m orp h ol o gi c a ll y a cc ou nte d for s i m ply b e c a u s e

,
t he re h a pp e ns t o be a t t ha t pl ac e a n a bu nd a n ce of u no rg a niz e d p ro
to p l as m ; th e n c le a rly i t i s lik ely t ha t i f t h e m a ss o f p rot opl a s m
, ,
from wh i ch a l eaf w ou ld us ua lly ar is e i s grea tly i ncreas e d i n m as s

,
by e x c es s of nut ri ti on i t m ay d ev elop i nto a n a x is i ns t ea d of a le af

, .
Ma ny y ea rs after th i s p u bli sh e d I di s co v ered a m ong

w ork w as ,
m y p a p ers a m em o ra nd u m w h i c h u nfort u n a t e ly I ha d o v e rl ook e d ,
c o nt a i ni ng fu rt h er e vid e nce i n s upp ort of th e foreg o i ng c o n cl us i o n .
W i th th e om i ss i on of a n e rror c oncerni ng th e s peci es of pla nt I ,
rep ro d u c e t hi s m e m ora nd u m j us t as i t s t o o d
I found at D i epp e July 1 1 8 6 0 i n a g arden ne a r th e s ea a

, , ,
s a m pl e o f c ult i vat e d w ild fl o w e r ( I th ou ght i t was g ro wn a s a n

o rna m ent a l fl o we r
) i n wh i ch s om e o f th e s i ngl e fl o w e rs o f the
u m bel w e re d ev e l op e d i nto g roup s of flow ers thus
I n the case where th e tra nsform ati on w as fully effe cte d th e

u m b ell ul e h a d si x fl ow ers , a ns w eri ng t o th e si x p eta l s o f th e
o ri gi na l flo we rs I n oth e r c a s es th e tra ns form ati on w a s i nc om

.
l t Th ere w ere i ns t a nc es wh ere bu t two of th e p et a l s w e re

p e e .
d e vel op ed i nto flow ers ; a nd the oth er p etals rem a i ned uncha ng ed .
O thers i n whi ch three w ere d ev el op e d ; a nd o th ers wh ere fou r

w e re d e vel op ed In s om e c a ses to o, th e d ev el opm ent o f a p etal
.
,
i nto a fl o wer w as i m p erfe ct, i n th e ab s e nce of the fl o wer s ta l k

th e fl o w ers w ere s ess il e i n th e pl ac e wh ere th e p et al s w ould
h a ve b een I n one cas e th ere was a n i m perfect flo we r s es s il e ;
.
a n o th e r i m p erfect fl owe r o n a s h ort s talk ; a nd th re e p erfect
flowers on l ong lks

sta .
S U BST I T U T ION OF A XI A L F OR F OL IA R ORGA NS .
547
I w a s i n s om e d ou b t wh e th er th e p eta ls or t he s ta m ens w ere

d ev el op e d I n c as es of i m p erfe ct tra ns form a ti on th e p et al s a t
.
t he b as e of th e u m b ellu l e s eem e d t o s tan d i n the p os i t i o n of

c al yx o r i nv ol u cr um , g i vi ng th e id ea th a t th e s ta m e ns w e re d e
v e l o e d i nt o fl ow e rs B u t i n t h e c as e w h e re t her e w e re s i x flo w ers
p .
d e v elop e d th ere were no p eta ls a t th e bas e .
T ha t i t was a m a tte r o f e xtra nu tri t i o n w as s h ow n by this

1 T ha t th ey w e re c ult i va t e d as g ard en flo wers
. .
2 T ha t w h ere the re was o ne p e rfe ctly d e vel op e d u m b e llul e,

.
i t w as t h e o nly o ne i n t h e u m b el .
3 T h a t w h ere th ere w ere t hre e u m b ell u les th ey w ere al l i m

.
p erfec t .
4 Tha t i n th i s i m p erfe ct um b ell u l e th e p e rfe ct flowers w ere

.
o n l on s tal ks a nd th e i m p erfect o nes s es s il e

g .
5 Th at t he um b ellu l es w ere on s ta lk s b oth long er a nd

.
”
th i ck er th a n th os e of s i ngl e fl owe rs .
[ Conce rni ng th e foregoi ng

g at
a r u m e nt a n e x p e rt
‘
w rit es T he a b norm ali ti es you d es c rib e c e sh ow t ha t

a n a x is m a
y a ri s e a b norm ally i n t he pl a c e o f a norm al l e af
s tru ctu r e , a nd e v e ry m o d ern b ota ni s t w oul d be i n agr e em ent
w i th y o u i n y ou r c ri ti ci s m of th e old e r form of th e d oc tri ne of
a xill ary b u d s I thi nk w e a re l arg ely em a ncip ate d from th e
.
d e x tro u s j uggli ng wit h th e a rra ngem e nts a nd rel ati ons of orga ns
w h i ch u s e d t o p a s s c u rr e nt as m orp h olo y
g .
You ha ve q uot e d s u fli ci e nt e v id ence i n th e t e x t 1 9 0) t o

e s t a bli sh th e c o nc lus i on th a t n o s h a rp l i ne c a n b e d ra wn b etw een
a x e s a nd l eaf s t ru ctu re
; a nd a v ery gr ea t d e al m ore c ould be
-
a dd e d i n the s a m e s ens e P e ti ol es for i nsta nce e x i s t w hi ch t he

.
,
m os t h ighly t ra ine d h i s t ol ogi c al o b s erver c ou ld not di s ti ngu i s h

from s te m s .
B ut I d em u r to th e s ugg esti on th at th e repl a cem ent of

m us t
o ne by the o th er i s p ri m a rily a q u es ti o n o f nu t ri ti o n W e a re .
as i g no ra nt as e v er o f th e p ro x i m a te c a u s e o f th e p ro d u c t i o n
of a l eaf or a s h oo t a t a c e rt a i n s p ot i n m eri s t em a ti c t i ss u e
”
.
T o th i s l as t r em a rk I h a d a t fi rs t m a d e only th e reply th a t th e
pl a nts e xhibiti ng t he a b norm a liti es w ere i n all c a s es e x c ess i v ely
l uxuri a nt i n th ei r growt hs ; but t o thi s I a m now a bl e to a d d a
T h e e x p ert from w h om I h a v e ju s t qu ot e d
'
m ore d efini t e reply .

,
h a d re a d th i s a pp e ndix be fore th ere h a d b e en m a d e t o i t th e

a b o ve a ddi ti on d es cribi ng th e fl ow er fro m D i epp e
; a nd I w as not
m y s e lf a w a re u ntil I ca m e t o r ea d o v er thi s a ddi ti on wh a t cl ea r
, ,
e v id e n ce i t c ont a i ns t ha t e xtra nu t ri t i o n w as th e c aus e o f th ese
t ra ns form ati ons o f fo li a r s t ru c tu res i nt o a xi al s tru ctu re s ; but

t h e a bo v e p a ra gra p hs 1 2 3 4 5 cont ai n di fferent evid ences
, , , , ,
c o ns pi ri ng t o p r o v e t h i s
]
A P P E ND IX B
A CRI TI CI SM ON P ROF . OW E N S THE ORY OF THE

’
VE RTE BRA TE S KE L E TON .
[ F rom the BRITI SH FOREI GN ME D I CO CH IRU RGI CA L RE VI E W Ocr 1 858 ]

-
FOR .
,
I . On the A rchetyp e a nd H om ol og i es of the Ver tebr a te S kel eton .
B y RI C H A R D O W E N , F R S — L ond on, 1 8 4 8
. . .
pp 1 7 2 . . .
II . P r i ncip es d Os téol oy i e Com p a r es , ou R echerches

’
su r
’
l A rché
typ e et l es H om ol og i es da S qu el ette Ver tébr é P . ar RI C H A RD
O W E N — P a ri s
. .
P ri ncip l es of Com p a ra ti ve Os teol og y or , R esea r ches on the A rche
typ e a nd the H om o l og i es of the Ver tebra te S kel eton B y RI C H A R D .
OW E N .
III . On the N a tu r e f L i m bs
o A D i scou rse del i vered on F rida y ,
.
F ebr u a ry 9 a t a n E veni ng M eeti ng of the R oya l I ns ti tu ti on of

,
Grea t B r i ta i n B y RI C H A RD OW E N , R E S — L ondon, 1 8 4 9
. .
pp . 119 .
J U D G I N G w h e th er a noth er p ro ve s hi s p os i ti on i s a W id ely di fferent

th i ng fro m p ro v i ng y ou r o w n T o es ta bli s h a g eneral l a w re qu i res
.
a n e x t ens i v e k no w l e dg e of t h e ph e no m ena t o b e g enera li z e d ; but t o
d e c id e Wh eth e r a n all eg ed g ene ra l l a w i s es tabli sh ed by the e v id e nce

a ss ig ne d , r eq u i re s m erely a n a d e q u a te rea s o ni ng fa c u l ty E s p eci ally .
i s s u c h a d e c i s i o n e as y wh ere the p rem i s e s do no t w a rra nt the c on ‘
e lu s i on It m a y be d a ng erou s for one wh o h a s bu t littl e p r evi ou s

.
a c u a i nt a nc e w i th t h e fa ct s , t o s a y t h a t a g ene rali z a ti o n i s d em o n
q
s tra t ed ; s e e i ng th at the a rgum e nt m a y be o ne s id e d : th ere m ay be -
m a ny fa c ts u nk no w n t o hi m Wh i ch d i s p rov e i t B ut i t i s not
.
d ang erous t o gi v e a nega tiv e v erdi ct wh en t he alleged d em ons t ra

5 48
CRIT ICISM ON P R OF OWE N S T HE OR Y
’
A . .
54 9
ti o n i s m a nifes t ly i ns ufli ci e nt
. I f th e d ata p ut b efore h i m d o not
i nfe rence, i t i s com p etent for e very l ogi c al rea d er t o
'
b ea r ou t th e
s ay s o .
From th i s s t a nd -p oi nt, th e n, w e v e nture to c rit i ci z e s om e o f

P rofes s or O we n s o s t e ologi cal t heori es Fo r hi s k no wl e dg e o f
’
.
c om p ara ti v e o s t eol ogy w e h a v e th e h igh es t re s p e c t W e beli e v e

.
t h a t n o li v i ng m a n has s o w id e a nd d e t a il e d a n a c q u a i nt a nc e w i th
th e b ony s t ru c t ure of th e Ver tebra ta I nd ee d there p roba bly has
. ,
ne v e r b een a ny on e w h os e i nform a ti o n o n th e s u bj ec t w as s o n ea rly

e x h a u s t iv e Moreo v er, we confes s th a t ne arl y a l l w e k now of th is
.
d ep artm ent of bi ology has b een l earnt from h i s l e ctures a nd w ri t

i ngs W e p re t end to no i nd ep end e nt i nv estig atio ns , but m e rely to
.
s u ch k n o w l e dg e o f th e p h enom en a a s h e h as fu rni s h e d u s w i th .
Ou r p o s i ti on, t he n i s s u ch th at, ha d P rofess or O w e n s i m ply e nu n

c i s t e d hi s g e nera l i z a ti ons , w e s h ou ld h a v e a c c e p t e d th e m o n h i s
a u th ori ty .B u t h e h as b rought for wa rd e v id enc e t o p ro ve th em .
B y s o d oi ng h e h a s t a ci tly a pp eal e d to th e j u dgm ent s of hi s rea d ers

a nd h ea rers — h a s p ra c t i c a l l y s a id , H ere a re th e fa c ts ; d o the y
”
not w a rra nt th es e c o nc lu s i ons ? A n d a l l w e p ro p os e t o d o, i s to
c o n s id er w h eth e r th e c o nc l us i ons a re w a rra nt e d b t h e fac ts brou gh t
y
for wa rd .
L et u s fi rs t li m i t th e s cop e of ou r c ri ti ci s m s O n th at di v i s i o n
.
o f c om p a ra t i ve os teol o g y w h i c h d ea l s w i th w h a t P r o fess or O we n
”
dis ti ngui sh e s a s s p eci a l hom ologi e s , we d o not p rop os e to ent er .
Tha t th e W i ng of a bi rd i s fram e d u p on b ones ess e nti a lly p a ra ll el t o

th os e of a m a m m al s fore li m b ; th a t th e c a nn on b o ne of a h ors e s
’ ’
- -
l eg a ns w e rs to t he m iddl e m et a c arp al of th e hum a n h and ; th a t

v a ri ous b o n e s i n th e s k ull of a fish a re h om ologous w i th b one s i n
th e s kull o f a m a n— th es e a nd c ou ntl es s s i m il ar fa cts , w e t a k e t o be
w ell es ta bli s h e d I t m ay be , i nd ee d , th a t th e d oc t ri ne of s peci a l
.
h o m ol og i es is a t p re s e nt c a rri e d too far I t m a y be th a t , j u s t a s

.
t h e s w eepi ng g e nerali za ti o n at one ti m e fa v ou r e d , th a t th e e m b ry oni c

p has es of t he higher a nim als rep res e nt th e a d ult fo rm s of l ow e r
o n es , h a s be e n fou nd u nt ru e i n a li t e r a l s e n se a nd i s a c c e p t a bl e
,
o nly i n a qu alifi e d s ens e ; s o th e s wee pi n g g e n era li z a t i o n th a t t h e
s k e l et o ns o f a ll v e rt ebra t e a ni m a l s c ons i s t o f h o m ol og ou s p a rt s
,
w ill h a v e t o u nd erg o s om e m o difi ca t i o n B ut th a t th i s g enera l i z a

.
t i o n i s s u bs ta nt i ally tru e , a ll c om p arati v e a na tom i s ts a r e e

g .
Th e d oct ri ne w hi ch w e a re h e re t o c o ns id e r, i s q ui te a s ep ara t e
o ne — th a t of g ene ral ho m ol ogi es
“
.
”
T h e tru th or fa ls i ty of th i s
m a
y be d ec id e d o n qu i te a p a rt from th at of th e oth e r W h e th e r
.
ce rt a i n b one s i n o ne v ert e b ra t e a ni m a l s s k el e t o n corr e s p ond w i th

’
c e rt a i n b o n es i n a no th e r s , or i n e v e ry o th er s i s o ne q u es ti on
’ ’
, ; and
w h e th er th e s k el et o n o f e v ery v ert ebra t e a ni m a l i s di v i s ibl e i nto a
s eri es o f s eg m e nt s , ea ch o f w h i ch i s m o d ell e d afte r the s a m e t yp e
,
i s a noth er qu e st i o n W hile th e fi rs t i s ans we re d i n t h e afii rm ati ve,

.
5 50 A P P E ND IX B .
th e l as t m aybe a ns w e re d i n th e neg ati ve ; a nd we pr op os e to gi v e

reas on s why i t sh ou ld be a ns were d i n th e ne g a ti v e .
In so far hi s t h eo ry o f th e s k e l et on i s c onc erne d , P rofess or

as
O wen i s a n a vow ed dis cipl e o f P l a t o . A t t he c o n c l u s i o n o f h i s

A r chetyp e a nd H om ol og i es of the Ver tebr a te S kel eton, h e qu otes ap
p ro v i ngly t he P l atoni c hyp oth esi s of 286m , a s ort of m od els , or
m ou l d s i n wh i ch m a t t er i s c a s t , a n d w h i ch re gul arly p ro d uc e t he
”
s a m e nu m b er a nd di vers i t y of s p e c i e s . Th e vert e b ra te form i n
g enera l ( s e e di a g ra m of the A r chetyp u s) , or els e th e form o f e a ch
ki nd of v ert eb rate a nim al (s ee p 1 7 2 , wh ere thi s s eem s i m pli e d) ,
.
P rofess or O we n c onc e iv es t o e x i s t a s a n id ea —a n a rche

t yp al e x em pl ar on w hi ch i t h a s pl e as e d th e C rea tor t o fra m e
c e rta i n of h i s li v i ng c re a tu re s
”
. W heth er P rofes s or Ow e n h old s
t h a t the typi c al v ert e b ra a l s o e x i s ts a s a n id ea , i s not s o
“
c ert a i F th e t i tl e giv e n t o hi s fig u re of th e id e a l typi ca l

“
n . ro m
”
v ert e b ra , i t w ou ld s eem th a t h e d o es ; a nd a t p 4 0 o f h i s .
N a tur e of L i m bs , a nd i nd ee d th rou gh ou t h i s ge nera l a rgu m e nt , thi s

s upp os i t i o n i s i m pli e d . B u t o n th e l a s t tw o p a g es of th e A rchetyp e
a nd H om o l og i es , i t i s di s ti n ctly a ll eg e d th a t th e rep eti t i on of s i m i
l ar s egm ents i n a v ertebral colum n, a nd of s i m il a r el e m ents i n a
v e rt e b ra l s eg m ent, i s a nalog ous t o th e rep etiti on o f s i m ila r c ry s t al s
as th e res u lt of p o l a ri z i ng fo rc e i n t he gro w th of a n i n org a ni c
b o dy ; i t i s p oi nted out tha t, a s w e d es c e nd th e s cale of a ni m a l

“
life, th e form s of th e rep e at e d p a rts of th e sk eleton app roa ch m ore

”
a nd m ore t o g e om e t ri c a l fig ure s ; a nd i t i s i nferre d th a t th e
“
P la toni c iSea o r s p e c ific org a ni zi ng p ri nc ipl e or forc e , w ou ld s e e m

’
t o be i n a ntag oni s m w i th th e g ene ral p ol a ri zi ng force , a nd t o s u b

‘
d ue a nd m ou ld i t i n s u b s e r v i e nc y to th e e xig enc i es of t he res ult i ng

s p e cifi c form I f P rofess or Owen s d o ctri ne i s t o be u nd ers t oo d
’
.
as e x p ress ed i n th es e cl o s i ng p a ra g ra ph s of hi s A r chetyp e a nd H o m o
w h i c h p rod u c es th e di v e r
”
l og i es i f h e c ons id ers th a t th e i Sea
’
—
s i ty of form b e l o ngi ng t o li v i ng b o di es o f t he s a m e m a t e ri a l s ,
”
is
m e t by th e cou nt er o pe ra t i on
- of th e p ol a ri zi ng forc e p erv a di ng
”
a l l s p a c e, w hi ch p rod u ces t h e s i m il a ri ty o f form s , the re p et i ti on
o f p a rt s , t h e s ig ns o f u ni ty of o rg a ni za ti o n, a nd w h i c h i s su b
“
d u ed a s w e as ce nd i n th e s c al e of b ei ng
“
t h e n w e m a y p as s o n
w i th th e rem a rk th at th e h yp o th e s i s i s t oo c um brou s a nd i nv ol v e d
t o h a v e m u ch vr a i s em bl a nce .If, o n th e o ther h a nd , P rofess o r Ow e n
h old s , a s e v e ry re a d er w o u ld s up p o s e fro m th e g e ne ra l t e nor o f hi s
re a s oni ng s , th at no t o nly d oes t h e re e xi s t a n a rc h e typ a l or id e a l
v ert e b rat e s k ele ton, bu t th a t th e re a ls o e x i s ts a n a rch ety pa l or
i d eal v e rt e b ra ; th e n h e ca rri es th e P l a t oni c hyp o th es i s m u c h
fu rth er th a n P l a to d o es P l a to s a rg um e nt, th a t b efo re a ny s p eci es
’
.
o f o bje ct w as c rea t e d i t m u s t h a v e e xi s t e d as a n i d e a o f th e C r e a ti v e
l
I nt ellig e nce, a nd tha t h enc e a ll o bj e cts of s uch s p e ci es m us t be

A C RITI CIS M ON P R OF OWE N S T H E OR Y 5 51
'
. .
co pi es of th i s origi na l id e a i s t ena bl e enough from th e a nthrop o

,
m o rp h i c p o i nt o f v i e w B u t w h i l e t h os e w h o wi t h P l at o th i nk fi t
.
, ,
t o ba s e t h e i r t h eory o f c re at i o n u p o n t h e a na l ogy o f a c a rp e nt e r
’
d es igni ng a nd m a ki ng a ta bl e m u s t yi eld as s ent to P l a t o s i nfere nce
, ,
th ey a re by no m ea n s c om m i t t e d t o P rofess or O we n s e x pa n s i o n o f
’
it. T o s a y th a t be fo re c re a t i ng a v e rt e bra t e a ni m a l G o d m u s t ,
h a v e h a d th e c o nc e p ti o n o f o ne d oes not i n v ol v e s a yi ng t h a t Go d
,
g ratu it o u sly bou n d h i m s el f t o m a k e a ve rt e b rate a ni m al ou t of s eg

m e nts a l l m ou ld e d aft e r o ne p a t t ern A s t her e i s n o c o n cei va ble
.
d v g i l g d dh i f d l p —
a a nt a e i n t h s a l e e a es o n t o a u n a m e n t a a t t e r n as ,
fo r t h e fu lfil m e nt o f t h e i nt e nd e d e nd s i t i s not o nly n ee dl es s b ut

, ,
o ft e n a s P ro fes s o r O w e n a r gu es l es s a pp ro p ri a t e th a n s om e ot he r
, ,
s e e N a tu r e of L i m bs pp 3 9
c o ns t ru c ti o n w o uld be
( to su p , .
,
p o s e th e c rea t i v e p rocess es thu s regul at e d i s not a li ttl e s t a rt li ng ,

.
E v e n t h os e w h os e c o nc ep t i o ns a re s o a nth ro pom orp h i c as t o th i n k

”
th e y h ono u r th e C re a t or by c alli ng h i m t h e Gr e a t A rti fic e r w ill ,
s c a rce ly as c ri be t o h i m a p r oc e e di ng w h i ch i n a h u m a n a r ti fi c er , ,
t hey w ou ld co ns id er a n ot v e ry w o rt hy e x e rc i s e o f i ng e nu i ty .
B ut w h i c h e v e r o f t h es e a l te rna t i v es P ro fess o r O w e n c ont e nd s

for— w h et h e r t h e typi ca l v e rt e bra i s th a t m o re or l es s c ry s t all i ne
figu re w hi ch oss e o u s m a tte r e v er te nd s to as s um e i n s pit e of th e
or w h e t h e r t he typi c a l v e rt e b ra i s
”
iSea o r org a niz i ng p ri nc i pl e
’
i ts elf a n i Sea o r o rg a niz i ng p ri ncipl e — th ere i s alik e i m pli e d

“ ’
t h e b eli e f th a t t he ty pi c a l ve rt e b ra h as a n a bs t rac t e x i s t e nc e a p a rt
from a ct u a l v e rt ebrae It i s a form wh i ch i n e v e ry e nd os k el eto n
.
, ,
s t ri v es t o e m b o dy i ts e lf i n m a tt e r — a form w h i c h i s p ot e nti a lly
p re s e nt i n e a c h v e rte bra ; whi ch i s m a ni fe s te d i n e a ch v ert e bra

w i th m o re o r l es s cl e a rn es s ; bu t w hi c h i n c o ns e q u e nc e o f a nta g o
,
n i xi ng fo r ces i s n o wh e re c o m pl e t e ly re a liz e d
,
A p art from t he .
p hil os oph y of thi s hyp othe s i s l et u s h ere e x am ine th e e vid e nc e

,
wh i ch i s t ho u gh t t o j us tify i t .
An d fi rs t as to th e i l
es s ent a itu ents of th e id eal ty pi ca l
c ons t
“
v e rte bra . E x clu s i ve o f di verg i ng app end a g es w hi ch i t 2 ‘

m ay
” “
a l so s upp o rt , i t c o ns i s t s i n i t s typi c a l c om pl e t e ness of th e foll o w
”
i ng e l e m e nt s a nd p a rt s A cen tr u m r ou n d w h i ch t he re s t a r e
-
a r ra ng e d i n a s om ew h a t ra di a t e m a nn er ; a b o v e i t t w o neu r a o h s es
p p y
— c o n v erg i ng a s th ey a s c e n d a nd form i ng wi th th e c e ntru m a t ri a n
,
u l o i d s p a c e c onta i ni ng th e n e ura l a x i s a neu r a l s i ne , s u rm o u n ti n

g p g
t h e t w o ne u ra p o phy s es , a nd w ith t he m c om pl e t in g th e ne u ral a r ch
b e l o w th e c e nt ru m t w o hazm ap op hy ses a nd a haem a l sp i ne, fo rm i ng a
h aem al a rc h s i m il a r t o t he ne u ra l arch a bo ve , a nd e ncl o s i ng th e
b es m e l a x i s ; t w o p l eu rap op hy ses ra di a ti ng h o ri z o nt a lly from t h e
_
s id e s o f th e c e ntru m ; a nd t w o p a r ap op hy s es di v e rgi ng fr om th e
”
c e n tru m b el o w t h e pl eu r a p op hy s es Th es e, s ay s P rofe ss or
.
O wen, b ei ng us ually d e v elop e d from dis ti nct a nd i nd ep ende nt

552 A P P E ND I X B
'
.
c e ntre s , I h a v e te rm e d au tog enous e l e m ents .

’
Th e rem ai ni ng
”
e l e m e nts , w h i c h h e c l a s s es as e x o g e no u s , b e ca us e th e y s h oot
”
o u t a s c o nt i nu a t i o ns fro m s o m e o f t h e p rec e di ng e l e m e nt s , a re
t h e di ap op hy s es di v e rgi ng fro m t he upp e r p a rt o f th e c e ntru m as

t h e p a ra p op hy s es d o bel o w , a nd th e zyg ap op hys es w hi c h g row out
o f th e di s ta l e nd s of t h e n eu ra p op hy s es a nd haem a p o ph y s e s .
If, o ,
n w t h es e a r e t h e c o ns i
t t e
u n t s o f h
t e e e v rt b ra g
t e s e m ent i n
“
i ts ty pi ca l c o m pl e tene s s ; a nd if th e v e rte b ra t e s k el eton c o ns i s t s

o f a s u cc es s i o n of s u c h s e gm e nts ; w e o u gh t t o h a v e i n th e s e c o n
s t i t ue nts , rep res e nt a t i v e s of a l l th e e l e m e nt s of th e v e rt e b ra t e
s k e l e to n— a t a ny ra te , a l l i ts e ss e nti a l el e m ents . A re w e t h e n t o
”
c o n c l u d e th a t th e divergi ng a p pend a g es , whi c h P rofe ss or Ow en
rega rd s as ru di m e nta l li m b s , a nd fro m c e rt a i n of wh i ch h e c ons id e rs
a c tu al li m b s t o be d e ve l o p e d , a re typi ca l ly l es s i m p o rt a n t th a n s o m e
o f th e a b o v e s p e c i fied e x o g e n ou s p a rts — s a
y th e zyga p ophy s es ?
-
T h a t th e m e a ni ng of thi s qu es ti o n m a y be u nd e rs to od , i t w ill be
ne e d ful b ri efly t o s ta te P rofes s or O we n s theo ry o f The N a tu r e of
’
L i m bs a nd s u ch c ri ti c i s m s as w e h a v e to m ak e on i t m u s t be i n
c l u d e d i n th e p a re nth es i s . I n th e fi rs t pl a c e , h e a im s to s how th at
th e s c ap u l ar a nd p el vi c a rch es , g ivi ng i ns ert i o n to th e fore a nd h i nd
li m b s res pect i ve ly , a re di spla ce d a nd m odifie d haem a l arch es ,
o rigi na lly b e lo ngi ng i n th e o ne c a s e t o th e oc cipi t al v ert e b ra , a nd i n
th e oth e r cas e t o s om e t ru nk - ve rt e b ra not s p ecifie d . I n s upp ort of

th i s ass u m p t i o n o f di s p l a ce m e nt, carri e d i n s om e c as e s t o th e e xte nt
o f twenty s even v e rt e b rae , P rofes s or O w e n c i t e s cert a i n a ck no w
-
l edge d dis pl a cem ents wh i ch occu r i n th e h um a n s k elet on t o the e x

t e nt o f ha lf a ve rte b ra — a s om e wh a t s l end er j u s t ifi ca ti o n . B ut for
p roof tha t s u ch a di spl a cem ent ha s tak e n plac e i n th e s c ap ula r a rch ,
h e chi efly re li es o n th e fact tha t i n fish e s , th e p ect ora l fi ns , w hi ch
a re t h e h o m o logues of th e fore li m b s , a re di re c tly a rt i c ul a te d t o
-
c erta i n b o n es a t t he b a c k o f th e h e a d , w h i ch h e a ll eg es a re p a rt s
o f th e o c c ipi ta l v ert eb ra . Thi s a pp e al to th e c la s s of fish es i s

a vo w e d l y m a d e o n th e p ri nc ipl e t h at th es e l o w es t o f th e Ver tebr a ta
a pp ro a c h cl os es t t o a rch ety pa l regu l a ri ty , a nd m a y th e refore be
e x e ct e d t o s h o w t h e o ri gi nal re l a ti o ns o f th e b o n e s m or e nea rly

p .
S i m ply not i ng th e fa cts th a t P rofe s s or O w e n d oes not giv e u s a ny

t ra ns i ti ona l form s b et w e e n th e all eg e d norm a l p os i ti o n of t h e
s c a p u l a r a rch i n fi s h es , a nd i ts e x t ra o r di na ry di s pl a ce m e nt i n t h e
h igh er Ver tebr a ta a nd th a t h e m a k es no refe re n ce to th e em b ry o ni c

p hases of t he high e r Ver tebr a ta , w hi ch m ight be e x p ect ed t o e x
b ibi t t he p rogres s i v e di s pl a c e m e nt ; w e g o o n to r em a rk tha t i n ,
t h e c as e of th e p el v i c a rc h , h e a ba nd ons h i s p ri nc ipl e o f a pp ea l i ng
t o t he l o wes t v ert e b ra t e fo rm s for p roof o f th e typi ca l s t ruc tu re .
I n fis h es , th e ru dim e nta ry p el v i s , w id el y rem o ve d from t he s pi na l

c ol um n , s h o w s no s igns o f h a v i ng bel o nge d t o a ny v e rt e b ra ; a nd
h ere P rofess or O we n i ns ta nc es th e p e re nnib ra nc hi a te B a tra chi a as

CRIT I CIS M ON P R OF OWE N S T H E OR Y
’
A . .
5 53
e x h ibiti ng th e typi c al s tru c tu re : re m a r l bi rds ki ng th a t m am m a s , ,
a nd r e p t il e s s h o w th e rul e o f c o nne x i o n a n d fis h es th e e x c e p t i on
”
, .
T h u s i n th e c as e o f t he sc a p ul ar a r ch t he e v id enc e a ffo rd e d by ,
fis he s i s held o f gre a t w e ight beca u s e o f thei r a rch etyp a l regul a ri ty ;

,
w hi l e i n t he c a s e of th e p e l v i c a r ch th e i r e v id enc e i s r ej ecte d as ,
e x cep t i o na l B ut no w h a v i ng as h e c o ns id e rs s h o wn th at th e s e
.
, , ,
b ony fra m es t o w hi ch th e lim bs are art i c ul ate d are m o difie d h aem a l

a rc h es P rofess or Ow e n p o i nts o u t th a t th e h aem a l ar ch es h a bi tu a lly
,
be a r c ert a i n di v ergi ng a pp end age s ;

“
a nd h e a i m s t o s h o w t h a t ~
the di v ergi ng a pp end a g es of th e s c ap u la r a nd p el v i c a rch es re

s e c t i ve l y a re d e v e lo p e d i nt o th e fore a nd h i nd li m b s T h e re a re
p , .
s e v e ra l i ndi rec t w a y s i n w hi ch w e m a y t e s t th e p roba bil i ty o f thi s
c o nclu s i o n I f th es e div e rgi ng a pp e nd a e s a re ru di m ent a l li m b s

.
g
”
futu re p oss i ble o r p ote nt i a l a rm s l egs w i ngs or fe et w e m ay
, , , ,
fa i rly e x p ec t th e m al w a y s t o b ear t o th e h aem a l ar ch es a r el a ti on

s u c h as t h e li m bs d o B u t th e y by no m ea ns d o t hi s
. A s the .
v ert e b rae a pp ro a ch th e t a i l th es e app en d a g es a re ofte n t ra ns ferre d

,
g ra d ua lly from th e pl eura p oph ys i s to t he p ara p ophysi s or e ven to ,

”
t he c e ntrum a nd n e u ral a r c h ( A rch a nd H om p . A g ain
. .
, .
,
i t m i ght n a tu rally be ass u m e d th a t i n th e l ow es t v ert e b r a t e form s ,
w h ere t h e li m bs a re bu t li ttl e d e v el op e d th e y w ou ld m o s t c l ea rly ,
di s pla y the i r alli an ce w i th th e a pp end a g es or ru di m ent al li m b s , ,
by th e s i m ilarity of th ei r a tta chm ents I ns te a d of thi s h ow ev er .

, ,
P rofess or Owe n s dra w i n s s h ow t h a t w h e re as th e a pp enda g e s a re

’
g
h a bi tu a lly a tt a ch e d t o th e pl eu ra p op hy s e s th e li m bs i n th ei r e a rli es t , ,
a nd l o w e s t p h a s e a lik e i n fis h e s a nd i n th e L e i d osi r en a r e a rt i cu
, p ,
l at e d t o the h aem apophys e s Mos t a no m al ou s of al l h ow e v er i s.

, .
th e p roc e s s o f d e v elop m e nt Wh en w e s p e ak of one th i ng as be i ng

.
d e v e lop e d ou t of a noth er w e i m pl y th a t th e p arts ne x t t o th e g e rm

,
a re th e firs t t o a pp e a r a nd th e m os t c o ns ta nt
, I n th e e v olu t i o n of .
a t re e o u t of a s e e d t h er e c om e a t th e o u t s et t h e s t e m a nd th e
,
ra di c l e ; afte rw a r d s th e bra nc h e s a n d di v e rg e nt roots

; a nd s till
la t er th e b ra nchl ets a nd rootlets ; th e re m o tes t p a rts b e i ng th e l a tes t
a nd m os t i n c o ns t a nt I f t hen a lim b i s d e v e lop e d ou t of a
.
, , di -
v e rgi ng a p pend ag e of t he h aem a l a rc h th e ea rli es t a nd m o st c on ,
s t a nt b o n es s h o u ld b e th e h u m e ru s a n d fe m u r ; n e x t i n o rd e r o f
t i m e a nd c ons t a n c y s hou ld c o m e th e c ou pl e d b o ne s bas e d o n th e s e ;

w hil e th e t e rm i na l g rou ps o f b o ne s s h ou ld b e th e l as t t o m ak e th e i r
a pp e a ra nc e a nd th e m o s t li a bl e t o b e a b s e nt
, Ye t as P ro fes s o r .
,
O wen hi m s el f s ho w s th e a ct u a l m o d e o f d e v el opm e nt i s t h e v ery re

,
v ers e of thi s A t p 1 6 of th e A r che typ e a nd H orno l ogies he s ay s

. .
,
The ea l i e s ta ge s i n th e de v el op m ent of a l l l oc o m oti v e e xt re m i t i es a re

r r
p e rm a nent l y re t a i ne d or r e p r e s ente d
i n t h e p a i re d fi n s o f fi s h e s Fi rs t t he .
e s s e nti a l p a rt o f t h e m e m be r , t h e h a nd or foot a pp e a r s : t h e n t h e for e a r m or

,
-
fl
l eg bo th m u c h s h or t en e d, a t t ene d, a nd e x p a nded, a s i n a l l fi ns a nd a l l e m
,
b ryoni e r u di m e nt s o f l i m b s : fi n a l l y co m e th e h um e ra l a nd fem oral s egm e nts

b t t i s s ta e I
u h g h a ve t fo nd a tta i ned i n ny fi s h
no u a
”
.
5 54 A P P E ND I X B .
T hat i s to s ay, a lik e i n as ce ndi ng th ro ugh th e Ver tebr a ta g ene

ra lly , a nd i n t ra c i ng u p t h e s u cc ess iv e p h as e s o f a m a m m a li a n e m
bryo, th e l as t d e v e l op e d a nd l eas t c ons t a nt di v i s i on of the li m b , i s

-
t ha t b a s i c one by w hi ch i t a rt i c u l a t es w i t h th e h aem al a r c h It .
s ee m s t o u s th a t , s o fa r fro m p ro v i ng hi s hyp o th es i s , P rofess er

O w en s o wn fa cts t en d t o s h o w th at l im bs do not bel ong t o th e
’
v e rt e brae a t a l l ; tha t t hey m a k e th ei r firs t a pp ea ran c e p e ri p h erally ;

t h a t th e i r d e v e l op m ent i s c e ntrip e t a l ; a nd t h a t th ey be com e fi x e d
t o su ch p a rts of the v ert e b ra t e a x i s as th e r e qu i re m e nt s o f th e c a s e
d et erm i ne .
Bu t n ow , e n di ng h ere thi s digr es s i v e e x p os iti on and c ri ti ci s m ,
a n d gra nt i ng t h e p o s i t i o n th a t li m b s a re d e v el op m e nt s o f c os ta l
a pp e nd a g es
”
l et u s re turn t o th e qu es ti on a b o v e pu t— W hy a re not
,
t hes e a pp e nd a g es i nclu d e d a s el em e nts of t h e id ea l typi c al ver

“
t e b ra It c a nnot be b ec a u s e of th ei r c om p a ra ti v e i nco ns t a n c y ;
for j u dgi ng from th e i l lu s tra t iv e figu res th e y s e em t o be a s c o n
,
s t a nt a s the h aem a l s pi ne w h i ch i s o ne o f th e s o c a ll e d a u t og e no u s
,
-
e l em e nts : i n th e di agr a m o f th e A r che typ u s th e a pp e nd a e i s re

, g
p res ente d as a tta ch e d t o e v ery v ert eb ra te s egm ent of th e h e a d a nd
t runk w h i ch th e h aem a l s pi ne i s no t
, I t c a nnot be from the i r c om
.
p a ra t i ve u nim p ort a nc e ; s e ei ng th a t a s p ot enti al li m bs th ey a re

e s s e nt i a l p a rt s o f ne a rly a l l th e Ver tebr a ta — m u ch m ore o b v i ou s ly
s o th a n t h e di a p o ph y s es a re . If a s P rofes s or Ow en a rgu es
,
the ,
divi ne m i nd “ h i c h pl a nned th e a rchetyp e a ls o forekne w a ll i ts

m o difi ca ti on s a n d if a m o ng th es e m o di fi c a t i o ns th e d e v el op m e nt
, ,
o f li m b s o u t of di v e rgi ng app e nd a g es w as o ne i nt en d e d t o c h a ra c
t e ri z e a l l th e h i gh e r Ver tebr a ta th en s u rely th ese di v e rgi ng a p

, ,
p end ag es m us t h a v e b ee n p art s o f th e id eal typi c al v e rte b ra

“ ”
.
O r i f th e id eal typi c a l v e rt ebra i s to be u nd ers t oo d a s a c ry s t a l

,
“
li ne form i n a ntago ni s m wi t h th e orga ni zi ng p ri nciple ; th e n w hy

s h ou ld n ot th e a pp e nd a g es be i ncl ud e d a m o ng i t s v a ri ou s of fs h oo ts
W e d o no t ask th i s q u e s ti o n b ec a u s e of i ts i nt ri ns i c i m p o rt a nc e .
W e a sk i t for th e p u rp os e of a s c ert ai ni ng P rofes s or Owen s m eth o d ’
o f d e t e rm i ni ng w h a t a re t ru e v e rt e b ra l co ns t i t u e nts H e p res e nt s
.
u s w i th a di a g ra m o f th e typi c a l v e rte b ra i n w h i c h are i ncl ud e d

,
c e rt a i n b o ne s a n d fro m w h i c h a re e x c l u d e d c e rt a i n o th ers
,
I f re .
l a t i v e c ons ta ncy i s th e c rit e ri o n th e n there ari s es the qu es ti on

,
W h a t degree o f c ons ta ncy e nti tles a bone t o be i ncl u d e d ? If re

l a ti v e i m p ort a nc e i s the c ri teri on the re co m e s not o nly the qu es ti on
,
— VVh a t d eg ree o f i m p ort a nc e s u ffi c e s ? bu t t he fu rt h er q u es t i o n

— H o w i s i m p ort a nc e t o be m ea s u re d I f nei th er o f t h e s e i s t he
c ri t e ri o n th e n w h a t i s i t ?
,
A nd i f t he re i s no cri te ri o n do es i t ,
no t foll o w t h a t th e s el ecti o n i s a rbi tra ry
Th i s
qu es ti o n s erv es to i ntrod u ce a m u ch w id er one z— II a s th e
id ea l typi ca l v erte b ra a n es s e nti a l c o ns ti t ue nts a t a l l i I t m igh t
y
CRITI CIS M ON P R OF OWE N S T H E OR Y
’
A . 555
n a tu ra lly be s u pp os e d th a t thou gh s om e b ones are s o rarely

d e velop e d as not t o s e em w orth i nclu di ng a nd th ou gh s om e t h at ,
a re i nc lu d e d a re v e ry a t t o be a bs e nt v e t th a t c e rt a i n o th e rs a re
p ,
i nva ri a bl e : for m i ng as i t w ere t he bas rs of th e id e al typ e L et

, ,
.
u s s ee w h eth e r t h e fa c t s b e a r ou t th i s s u pp os i t i o n I n h i s s um m a ry .
”
o f m o di fi c a ti ons of c orp o ra l v e rt e brae p
( P r ofes so r O w e n .
s ay s Th e hce m a l sp i ne i s m u ch l es s c o ns t a nt as t o i ts e x i s t e nce ,
a nd i s s u bj ec t t o a m u ch g re a t er ra ng e of v a ri ety w h e n p res e nt ,
t ha n i ts v e rt i c a l h o m otyp e a b o v e wh i c h c om pl et es th e n e u ra l a rch
,
.
A g a i n h e s ay s Th e hce m ap op hy ses a s o s s eou s el em e nts of a ,
v e rt e b ra a re le s s c ons ta nt tha n th e pl eu ra p op hy s e s
,
”
A n d a g a in .
Th e p l eu ra p op hys es a re l es s c on s ta nt e l em e nt s th an th e neu ra po
”
ph y s e s . A nd a ga i n A m ong s t a i r b reath i ng v ert eb ra te s the -
l e ur ap op hy s es o f t h e tru nk s e m e nts a r e p res e nt o nly i n th os e s p e

'
p g
c i es i n wh i c h th e s ep tu m o f th e h e a rt s v e nt ri cl e i s c om pl et e a nd i m
’
p erforate a n d h e re th e y a re ex og e n ou s a nd c onfine d t o t he c er v i c al
,
”
a nd a nt e ri or th orac i c ve rt ebrae A nd o n c e m ore b o th th e neu r a
.
,
o
p p y h se s a n d t h e ne u r a l sp i ne a r e a bs e nt u n d e r b o th h i s t o l o gi c al
c o ndi ti o ns a t t h e e nd o f t h e t a il i n m o s t a i r b re a th i ng v e rt e b r a t es
,
-
,
”
w he re t h e s eg m e nts a re re du ce d to th e ir ce ntra l e l e m ent s Th at .
i s to s a y o f a l l t h e p e ri ph era l el em ents o f th e
,
id ea l ty pi c al v er “
”
t e bra ,t he re i s n ot o ne w hi c h i s a lw ay s p re s ent I t w ill b e e x .
ec t e d h o w e v e r t h a t a t a n ra t e th e cen tr u m i s c o ns t a nt : t he
p , , y
b one w hi c h form s t he a x i s o f t h e v ert e b ra l c olum n a nd c om
“
,
m ou ly t h e c e nt ra l bo n d o f u n i o n o f th e p e riph er a l e l em e nts o f th e
v erte bra t e ( p i s o f c ou rs e a n i nv a ri a ble el em e nt

. No : n ot .
e ve n th i s i s ess e nt i a l .
The c e nt u m s d o not p a s s beyond th e p ri m i t i v e s tag e of th e not och o d

r r
(undi v i d e d co l um n) i n t he e xi s t i ng l ep i dos i ren a nd t h ey r t a ine d t he l ik e ,

e
r di m e nt a l
u s ta t e i n e v ery fi s h wh o s e rem a i ns h a v e b een fo nd i n s t a t a u I
ea l ie
r th an t h e p e m i a n {
r B a i n Geol ogy th o ugh t he n m b er of v e t eb rae i s
r r ,
u r
f e q e ntl y i nd i ca te d i n D e voni a n a nd Si l uri a n i ch thyol i tes by th e fos s i l i ze d

r u
n eu r a nd h eem a p o phys e s a nd t h e i r ep i ne s (p .
I nd e e d ,
P ro fe ss or O w en h i m se lf
ks th a t th e n e u ra po
r em a r
“
p hy s es are m ore co ns ta nt as o ss eou s or ca rt il agi nou s el em e nts of th e

v e rte b rae th a n th e c ent ru m s (p Th u s th e n i t a pp e a rs th a t
.
, ,
t he s e v e r a l el e m ent s i nc l u d e d i n th e id e al typi c a l v ert e bra ha v e

“
v a ri o u s d egr ees o f c o ns ta ncy a nd th at no o ne of t he m i s e s s e nti a l

,
.
T he re i s no o ne p a rt of a v e rt e bra w h i ch i n v a ri a bly a ns w e rs t o i t s
e x e m pl ar i n t h e p a t t e rn g ro u p H o w d o e s th i s fa c t c o ns i s t w i th th e
-
.
h ypoth es i s ? I f th e C re a t or s a w fi t t o m ak e t he v e rtebra te s k el et on
o u t o f a s e r i e s o f s e g m e nt s a l l form e d o n e s s e nt i a lly t h e s a m e m o d e l
,
— ii fo r th e m a i nt e na nc e of th e t y e o ne o f th e s e b o ny s eg m e nt s
, p ,
i s i n m a ny c a s e s form e d o u t o f a c o al es c e d g rou p o f pi ec es w h e re , ,
a s P r o fe ss o r O w e n a rg u e s a s i ngl e p i ec e w o uld h a ve s er v e d as w ell

,
o r b ett er ; th en w e o u gh t to fi nd th i s typi c a l r ep eti t i o n o f p a rt s

556 A P P E ND I X B .
un i form ly m a ni fes te d W ithou t a ny ch ange of s h a p e i t wou ld oh

.
,
v i ou s l h a v e b ee n qu i t e p o s s i bl e fo r e v e ry a c tu a l v e rt e b ra t o h a v e
y
i d l l h p f h id l — ru di m e nt a ll
co t n a ne a t e a rt s o t e e a o n e y w h ere th e y
we re not w a nte d E ven o ne of t h e term i na l b ones of a m a m m a l s
’
.
t a il m ight h a v e b ee n form e d ou t o f th e ni ne a u t og enou s pi e c e s ,
u ni t e d by s u t ure bu t a dm i tti ng of id e nt ifi c a t i o n . A s h ow e v e r
, ,
t her e i s n o s u ch u niform typi c a l re p eti t i on of p a rt s i t s e em s t o u s

,
tha t t o a cc ou nt for th e typi c a l re pe ti t i on w h i ch d oes occu r by s u p ,
p os i ng th e C rea t or to h a ve fi xe d on a p a tte rn v e rt ebra i s t o as cri be

-
,
t o hi m t h e i ncons i s t e nc y of form i ng a pla n a nd t hen a b a nd o ni ng i t .
,
If o n t he oth er h a nd P rofess or Ow en m e a ns th a t th e
,
id eal “
ty pi c a l v ert e b ra i s a c ry s t alli ne form i n a nt ag o ni s m w i th th e

id ea or org a ni zi ng p ri nciple th e n w e m igh t fa i rly e x p e ct to fi nd
i t m os t c l e a rly di spl a yi ng i ts c ry s t al li ne ch ara ct e r a nd i ts full c o m
,
pl e m ent o f p a rt s i n thos e pla c es where t h e org a ni zi ng p ri ncipl e

,
m a y be p res um e d t o h a v e i t to t h e s m al les t e x t e nt
”
s u bd u e d
“
.
Yet i n t h e Ver tebra ta genera lly a nd ev e n i n P rofe s s or O wen s

’
,
A r chetyp u s t he v ert e b rae of th e t a il wh i ch m us t be c ons id ere d a s

, , ,
if a ny th i ng l e s s und er t he i nflue n ce of t he orga ni z i ng p ri ncipl e

,
tha n th os e o f th e tru nk d o not m a nifes t th e id e a l form m o re c om

,
l et e l O n t h e c o nt ra ry a s w e a pp ro a ch t h e e nd o f th e t a il t he
p y
.
, ,
s u ccess iv e s egm ents n ot o nly l os e th ei r re m a i ni ng typi c a l e l e m e nts ,
but b ec o m e a s u n cry s talli ne l ooki ng a s c a n be c once i v ed

-
.
S upp os i ng, h o w e v e r, th a t the as s u m p t i on o f s u ppr es s e d or u nd e

v e10 e d e l e m e nts b e g ra nt e d — s u pp o s i ng i t t o be cons i s t e nt w i th
p
th e hy poth e s i s o f a n id ea l ty pi c al v e rt e bra , tha t th e cons ti tuent
”
p a rts m a y s everally be a b s ent i n g reat er or l ess nu m b er, s om et i m es

l ea vi ng o nly a s i ngl e b one to rep res e nt t hem a ll m ay i t not be that
s u ch p a rts as a r e p res ent , s ho w th e i r re s pec t i ve typi ca l na t u res by
s om e co ns t a nt c h a ra c t er : s a y th e i r m o d e of o ss i fica t i o n ?
T o th i s qu es t i o n s om e p a rts o f th e A rchetyp e a nd H om ol og i es
s ee m t o reply , Yes ; w hil e oth ers cl ea rly a ns w e r, No C ri ti c i s
“
.
i ng th e opi ni ons of Ge ofl roy St H il a i re a nd C u v i e r, w h o a gre e d

.
t hi nki ng t ha t os s ifi ca ti on from a s ep a ra te c e nt re w a s t h e t es t of a
s ep a ra t e b o ne , a nd th a t thu s th e re we re as m a ny e l e m enta ry b o nes
i n th e s k e le ton a s th e re were c e ntres of oss ifi ca ti on, P ro fes s or O wen

p o i nts out th a t , acco rdi ng to thi s t est, th e h u m a n fem ur, w h i ch i s
o s s i fie d from fou r c e nt re s , m u s t be reg a rd e d a s fo u r bo nes ; w h il e
th e fem u r i n bi rd s a nd rep tiles , w h i ch i s o ss i fie d from a s i ngl e

c e ntre , m u s t be re g a rd e d a s a s i ngl e bo n e Y e t, o n t h e oth e r h a nd ,
.
h e a ttach e s w e igh t t o th e fa ct t h a t the s k ul l of th e h u m a n fte t us

p res e nts the s am e oss ific cent res a s d o th os e of th e e m bry o ka n
g a ro o a nd the y ou ng bi rd ( N a tu r e of L i m bs , p
. . A nd a t p .
1 04 o f th e H om ol og i es , after gi v i ng a num b e r of i ns t a nces , h e s a y s
Th e se and th e l ik e corre s pondences b e tween th e poi nts of ossi fica ti on of

CRI T I CIS M ON P ROF OWE N S T H E OR Y
’
A . .
5 57
th e h u m foe ta l s k el e ton, a nd th e s ep a ra te bone s of the a du l t s kel e to ns of

an
i nfe ri or a ni m a l s , a re p regnant w i t h i n te res t , a nd rank a m ong t h e m os t s tri k i ng
il l us tra ti ons o f u ni ty of pl a n i n t h e v e rte bra te orga ni za t i on
”
.
I t i s t ru e th a t on t h e foll owi ng p ag e he see ks to e xpl a in th is

s eem i ng c o ntra di cti o n by dis t i ngui s h i ng
b e twe en th ose ce ntres of os s i fi ca t ion t h a t h a v e hom ol ogi es ] rel a ti ons , a nd

t h os e th a t h a v e te l eol ogi cal ones b et wee n th e s e p a ra te poi nt s of os s i fi ca:
t i on of a hu m a n bone w hi ch typify v e rt eb ral e l e m ent s , oft e n p e rm a nent l y d i s
t i n ct bone s i n t h e l o we r a ni m a l s ; a nd t he s ep ara t e p oi nts wh i ch , wi th ou t s u c h
s i gni fi ca t i on, fac il i t a t e t h e p rogres s of os teogeny, a n d ha v e for t h ei r obv i ou s
fi nal ca use t he well -b ei ng of t he gro wi ng a ni m al ” .
B ut i f t h ere a re thu s ifi cati on wh i ch h a v e h om o

c e ntr es o f o s s
l og i cal m ea ni ngs a nd oth ers w hi ch h a v e n ot th ere a ri s es th e qu e s

, ,
t i o n— H o w a re th ey a l w ay s t o be di s t i ngu i s h e d ? E v id ently i n
d ep end ent oss ifica ti on c eas e s to be a h om ol ogi cal tes t i f there a re ,
i nd ep e nd e nt os s i fi c ati ons th at h a ve n othing t o d o w i th t h e h om o

l ogi es A nd th i s b e c om es t he m ore e v id ent wh en w e learn that
.
th e re a r e c a s es w h e re ne i th e r a h o m ol ogi c a l n or a t el e ol ogi ca l
m ea ni ng c a n b e gi v e n A m o ng v a ri ou s m o d es of os s ifica ti on of th e
.
c e nt ru m P rofess or Ow en p o i nts ou t th a t t h e b o dy of th e hu m a n
,
a t l a s i s s om e ti m es o s s ifie d fro m tw o ra rely fro m th re e di s ti nct

, ,
c e ntr es pl a c e d s i d e by s id e (p 8 9 ) w h il e a t p 8 7 h e s a ys
. In .
o ss eo u s fis h e s I fi nd t h a t th e c e nt ru m i s u s u a lly oss i fi e d from s i x

p oi nts ”
. I t i s cl ea r th at th i s m od e of oss ificat i on h as h ere no h o m o
l ogi ca l s ignificat i on ; a nd i t w ould be difli cu l t to gi ve a ny t el eo
l ogic al reas on why th e s m al l c e ntru m of a fish s h ould h a v e m ore
c e ntr es of o s s i fic a t i o n th a n th e l arg e c e ntru m o f a m a m m al Th e .
t ruth i s th a t as a c ri t e ri on o f the id ent i ty or indi vidu a li ty of a b one

, ,
m o d e of os s ifi ca t i on i s qu i t e u ntr u s tw o rth y T h ough i n hi s id e al “

.
,
”
typi c a l v ert ebra P rofess or O w en d eli nea t e s a nd c l as s ifi es as s ep a
,
ra t e a u t o g e n ou s e l e m e nt s th os e p a rts w hi ch a re
, u s u a ll y
d evelop e d from dis ti nct a nd i nd ep en dent c entres a nd th ou gh by
d oi ng s o h e e re cts thi s ch arac te ri sti c i nt o s om e s ort of crit e ri on ;

e t hi s o w n fa cts s h o w i t to be no cr i t eri on Th e p a ra p oph y s es
y .
a re c l as s e d a m o ng th e a u tog en ou s e l e m ents ; y et th ey are a u to
e n o u s i n fi s h es a l on e a n d i n th es e o nly i n th e trunk v e rt e b rae

g , ,
w h il e i n a l l a i r b rea th i ng v e rt e b rate s t h ey a re w h e n p r es ent a t al l

-
, ,
e x o e nou s Th e n eu ra p oph y s es a g a i n l o s e t h e i r p ri m i t i v e i n ;
“
g .
, ,
di v i du al i ty by v a ri ou s ki nd s a nd d egr ees o f co nflu e nc e i n th e

t a il s o f th e h igh e r Ver te br a ta th ey i n c om m o n w i th t he neu ra l
,
s pi ne b eco m e e x og e nou s
,
Na y e v en t he c ent ru m m a y l o s e i t s
.
,
a u t o g en ou s c h a ra c t er D esc ribi ng h ow i n s om e b at ra ch i a ns
.
, ,
“
th e o ss ifi ca t i on o f t h e c entru m i s c om pl et e d by a n e x te ns i on of
b one fro m th e b as es of th e neu ra p oph y s e s w hi ch e ffects als o th e ,
c oal es c e nc e o f th e s e w i th t h e c e nt rum P rofe s s o r O w e n a dd s

,
I n P el oba tes fu sca s a nd P el oba tes cu l trip es Mulle r found t he e u ,

558 A P P E ND I X B .
t i re ce ntru m os s i fi ed fro m -
thi s s o u rce, w i th out a ny i nd ep e nde nt
p oi nts ificati on (p
of o s s T h a t i s to s ay th e centrum i s i n
.
,
t hes e cas es a n e x og enou s p rocess of t he n eu ra p op hy s e s W e see .

,
th en t ha t t h e s e s o c a l le d typi ca l el e m ents of v erte b rae h a v e n o

,
-
c o ns ta nt d e v el o p m e nta l ch ara c t er by w hi c h th ey c a n b e id ent ifi e d .
N o t o nly a re they u ndi s ti ngui sh a ble by a ny sp ec ific t es t from o th e r

b ones no t i nclu d e d a s vert e bra l el e m e nts ; not o nly do they fa il t o
s h o w t h e i r t ypi c a l c h a ra c t e rs by th e i r cons ta nt p res e nc e ; bu t ,
w h en p res ent th ey e x hibi t no p ers i s t e nt m ark s o f i ndi v id u a lity

, .
T he cent ra l el e m ent m a y be os s i fie d from s i x fo ur th ree or t w o , , ,
p o i nts ; or i t m a y h a v e no s ep a ra te p oi nt of os s ific at i o n at al l
a nd s i m il a rly w i th v a ri ou s of t he p eriph e ra l e l em e nt s T h e w h ol e .
group o f b ones fo rm i ng the id ea l typi ca l v e rt eb ra m ay s evera lly

“
h a v e th e i r one o r m ore o ss i fic c e nt res ; or th ey m ay as i n a m a m ,
m al s t a il l o s e t h ei r i ndi v id u alit i es i n a s i ngl e b one oss i fie d from

’
,
o n e o r t w o p o i nts .
A noth er fa ct wh i ch s eem s v e ry di ffi cult to r econc il e w i th th e

hyp oth es i s of a n id ea l typi c a l v ert eb ra
“
i s t he no t i nfre q u e nt
p res ence of s om e of t h e typi ca l el e m ents I n d upli ca te N ot o nly , .
as w e h a v e s een, m a y t hey s e v era lly be a bs e nt, bu t th e y m ay s e v e
ra lly be p res e nt i n g rea t er nu m b er th a n th ey sh ou ld be W he n .
w e s e e , i n th e id ea l di ag ram , o ne c e nt rum , t wo neu ra p ophy s es ,

t w o pl eu ra p ophy s es , tw o h aem ap ophys es , o ne neu ral s pi ne , a nd o ne
h re m a l s pi n e , w e n atu rally e x pe ct to fi nd th em a l wa y s hea ri ng to
e a ch o th e r t h es e nu m eri ca l re l a ti ons T h ou g h w e m ay not be
.
g reatly s urp ris e d by th e a b sence of s om e of th em , w e a re h ardly

p rep a re d t o find others m ultipli e d Yet s u ch cas es a re com m on .
.
T hu s t he n eu ra l S pi ne i s d ou bl e i n th e a nt eri or v e rte b rae of s o m e

“
fishe s ( p A g a i n, i n th e a bd om i na l reg i on o f e x ti nct s au ri a ns ,

.
a nd i n c roco dile s , th e freely s u sp end e d h ae m a p ophys es a re com

“ -
p ou nd ed of t w o or m ore o v erla ppi ng b ony pi eces ( p Ye t .
a g a i n, a t p 9 9 , w e rea d
.
— “
I h a v e o b s e r v e d s om e o f t h e e xp a nd e d
pl eura p ophy s es i n th e g reat Tes tud o el ep ha ntop u s o s s i fie d from t wo
c e nt re s , a nd t he res ul t i ng di v i s i o ns c o nti nu i ng di s ti nct , bu t u ni t e d
”
by s ut ure O nce m ore t he neu ra ophys e s , w hi c h do n o t a d v a nc e
.
“
b eyond th e c a rtil agi nou s s ta g e i n t e s turg eo n, consi s t i n tha t fish

o f t wo di s t i nc t pi ec es o f c a rt il a g e a nd th e a nt eri or pl e u ra o phy s e s
p
a l s o c ons i s t o f t w o o r m o re c a rtil a g e s , s et e nd o n e nd
”
(p .
A nd els e wh ere referri ng t o th i s s t ructure , h e s a y s

Vegeta t i v e peri verteb ra l pa rt s not onl y m a ni fes ts i t s el f i n
rep eti t i o n o f
th e com p os i te ne u ra p op h ys e s a nd p l e u ra pophys es , but i n a s m a l l a c ce ssory

i ura l ) c arti l a e, a t th e fo re a nd ba ck
( nt erne g p a rt of th e bas e of th e neu ra
p ophys i s ; a nd b y a s i m i l a r (i nt e rhaem a l ) one a t the fore a nd b ac k p a rt of
m os t o f t h e p a ra pop hys e s (p .
Th us th e ne u ra l a nd h aem al s pi nes , th e neu ra p oph y s es , th e pl eu ~

A CRIT ICIS M ON P ROF OWE N S T HE OR Y
’
. .
5 59
ra p Oph ys es , th e h aem h s es , m a s e v e rally c o ns i s t of tw o or

a
p p y o
y
m ore pi e c es . Thi s i s not al l th e lik e i s t ru e e v en o f th e c entru m s .
“
I n Hep ta nc hus ( Squa l us ci ner ens ) th e v e rt eb ra l ce n tre s a re fe eb l y a nd
v egeta ti v el y m a rked out by n um erous s l ender ri ngs o f h a rd ca rti l a ge i n th e
notoc h ord a l ca p s ul e , th e n um be r of v e rt e brae b e i ng m ore defini t e l y i n di c a t e d
b y th e ne ura po phys es a nd pa ra pop hys e s In t h e p ik ed dog-fi s h
.
h d fi s h ( S c l l i u m ) th e v e rt eb r a l ce nt res co i n
( A c a nt i a s ) a nd t h e s p ot t e d og
-
y
c i de i n num be r wi th t h e ne ural a r ch es (p .
Is i t n ot s tra n ge th a t th e p attern v e rt eb ra s h ould be s o li ttle

a dh e re d t o, th a t e a ch of i ts s i ngl e t ypi ca l pi ec es m ay be t ra ns
form e d i nt o t w o or three
’
’
é
B u t th e re a re s til l m ore s ta rtli ng d ep a rt ure s from th e a ll eg e d
typ e . T he num eri c a l rel a t i ons of th e e l em e nts v a ry not o nly i n
t h i s w a y, bu t i n th e o pp os i t e w ay A giv e n p a rt m a y be p res ent
.
n o t o nly i n g rea ter nu m b er th a n i t sh ou ld be , bu t a l s o i n l es s In .
t he t a il s o f h om o c e rcal fi sh es , th e cent ru m s a r e re nd e red by cen
t ri p eta l s h orte ni ng a nd b o ny co nflu enc e fe w e r i n nu m b er th a n t h e

p ers i st ent neu ral , a nd h aem al a rch es o f th at p a rt —th at i s , the re
,
i s o nly a fra cti o n o f a centru m t o e a ch v ert e bra N ay , e v e n t h i s.
i s not th e m o s t h e tero cli t e s tru ct ure P ara d o x i c a l as i t m a y s eem ,

.
th ere a re cas es i n w hi ch th e s a m e v e rte b ral e l e m e nt i s , c ons id ere d

u nd er di f fere nt as p e cts , a t once i n e x ce ss a nd d efec t Sp e a ki ng .
o f th e h aem al s pi ne , P rofes s or O w en s a y s
The hori zonta l e xtens i on o f thi s v erte bra l el e m ent is s om e tim e s aecom
p a ni ed b y a m ed i a n d i v i s i on, or i n o th er w ords , i t i s os s i fie d fro m t wo
l a te ra l cent res ; t hi s i s s ee n i n t h e de v el opm e nt of p a rt s o f t he hum a n
s te rnu m ; t he s a m e v e geta t i v e ch a ra cte r i s cons t a nt i n th e b roa der th orac i c
h aem a l s p ine s of b i rd s ; th ough , som etim es , a s e g , in t he s truthi oni dae,
. .
oss ifica ti on extends fr om the sa m e l a ter a l c entr e l eng th wis e for wa r ds a nd

ba ck wa r ds , ca l cifyi ng the con na te ca r ti l ag i nous hom olog u es of ha l ves of fou r
o r fi ve harm a l s i nes , be
p fore thes e fi na lly coa les ce wi th thei r fel l ows a t the m edia n
l i ne ( p .
S o t hat th e s ternum of th e os tri ch , w hi ch a ccordi ng to th e h yp o

th es i s , s h ou ld , i n i ts ca rt il agi nous s ta g e , h a ve c ons i s te d of fou r or
ve tra nsver se pi e ces , a ns w e ri ng t o th e v ert e b ral s e gm e nts , a n d
fi
s h o uld h a v e b een os s ifi e d from fou r or fi ve c e nt re s , on e t o e a ch
c a rt il ag i no u s pi e ce, sh ow s n ot a t ra ce o f th i s s tru ct u re ; but i n
s t e a d , c o ns i s t s o f two l ong i tu di na l pi e ces of ca rtil ag e , ea ch oss ifi e d
fro m one c entr e , a nd fi nally c oa l esc i ng o n th e m e di a n li ne Th e s e .
fou r o r fi v e b es m e l s pi nes h a v e a t th e s a m e ti m e d ou bl e d th ei r i n
d i vidu a li ti es t ra ns v e rs ely , a nd ent i rely l o st th em l ongi tu di nally !
Th ere s till rem a i ns t o be c o ns id ere d th e te s t of rel a ti v e p os i ti on .
I t m i ght be h eld th a t s pi te of al l th e foregoi ng a nom a li es , i f th e

,
t ypi ca l p a rts of th e v ert eb rae al wa y s s t oo d to w ards e a ch ot h er i n

th e s a m e rel ati ons — al w ay s p re s erve d th e s am e c on ne x i ons , s om e
t hi ng li k e a c as e w ould be m a d e out D o u b tl es s , re l a ti v e p o s i t i on
.
i s a n i m p ort ant p oi nt ; a n d i t i s on e on w hi ch P rofess or Ow e n m a ni

fe s tl y pl a c e s gre at d ep e nd e nc e . I n h i s di s cu ss i on o f m o ot c as es
“
”
o f s p ec i a l h o m ol o gy , i t i s th e ge n eral t es t t o wh i ch h e a pp e al s .
Th e typi c a l na tu re s of th e ali sp h enoid , th e m as to id , the o rbi te

s p h e no id , t h e p refro nta l , th e m a l a r, th e s qu a m os a l , & e , h e d et e r
.
m i nes a l m os t w holly by refe renc e t o th e a dj ac e nt ne rv e p e rfora ti o ns

-
a nd th e a rt i cu l a t i ons w i th n eigh b ou ri ng b on es s e e pp 1 9 t o 7 2
( .
)
t h e g e neral form o f th e a rgu m e nt be i ng— T hi s b one i s to be cl as s e d
as s u ch o r s u c h , beca u se i t i s co nn ec te d t h u s a nd t hu s w i th th es e
o th e rs , w h i ch a re s o a nd s o . More ov er, by p u tti ng forth a n i d ea l “
”
typi ca l v e rt e b ra , c o ns i s ti ng of a nu m b er o f el e m ents s t a ndi ng
t o wa rd s each oth er i n c e rta i n d efinit e arra ng em e nt, thi s p ers i st ency
of r ela ti ve p os i ti o n i s m a ni fe s tly a ll e ge d . T he e s s enti al a tt ribu te
of thi s g ro u of b o n e s , co ns id e re d as a t pi c a l g rou p , i s th e co n
p y
s t a ncy i n t h e c o nne x i o ns of i ts p a rts : c ha ng e th e c onne xi o ns , a nd
th e typ e i s c ha ng e d . B ut th e c ons tancy of rel at i v e p os i t i o n th us

t a c i tly a s s e rt e d , a nd a pp ea l e d t o as a conclu s i v e t es t i n “
m o ot
”
c as e s o f s p ec i a l h o m ol o y , i s cl early n eg a tiv e d by P ro fes s or
g ”
O we n s ow n fa c ts For i ns ta nc e, i n th e id ea l typi c al vert e b ra ,
’ “
.
t he h aem al a rch i s rep res ent e d as for m e d by th e tw o h aem a pophys es

a nd th e h aem a l s pi ne ; bu t a t p 9 1 w e a re t old t ha t
.
Th e contra c ted h ae m al a rch i n t he ca uda l regi on of the b ody m ay be

form ed by di ffe rent el e m ent s o f th e t yp i ca l v ert ebra : e g , b y t he pa r a
. .
p Oph ys e s (fi she s gene ra l l y) ; by the p l eura p oph y s e s (l e pidos i re n) ; by bo th

p a ra pophyse s a nd pl eura pophys e s ( S ad ie, L ep i dosteus) , a nd by h aem a po
phys e s, s h ort ened a nd d i rec tl y ar ti cul ated wi th t he centrum s ( repti l e s and
A n d furth er, i n th e th ora x of rep til e s , bi rd s , and m am m al s , the

h aem a poph ys e s a re r em o v e d from th e c e ntru m , a nd a re a rt i cu lat e d t o
t h e di s t a l e nd s o f t h e pl e u ra p oph y s es ; th e b ony h oo b ei n c om
p g
pl et ed by t h e i nt erc ala ti o n of t he haem a l s pi ne ( p. S o th a t
t here are fi ve di ffere nt w a y s i n whi ch t h e haem a l a rch m a be form e d
y
— fou r m od es of a tta ch m ent of th e p a rts di ffe re nt from th a t s h own
i n th e typi c al di ag ra m ! No r i s th i s a ll The pl eu ra p op hy s es m ay
.
be qui te d eta ch e d from th ei r p rop er s egm ent, a nd s u s p end e d to the

h aem a l a rch o f a noth er ve rt ebra z as w e ha v e al rea dy s ee n, th e
”
e nti re h ae m al a rch m a
y be d eta ch e d a nd rem ov e d t o a di s t a nce ,
-s e v en v
s o m e ti m es reachi n
g t h e l e ngt h of t w e nty ert e b rae ; a n d, e v e n
m ore re m a rk a bl e, th e v ent ra l fi ns o f s om e fi s h es w hi ch th eore ti c a lly
,
be l ong to th e p el v i c a rch , a re s o m uch a dvanc e d forw ard as to be

a rti cul a t e d t o th e s ca p u l a r a rch th e i sch i u m el ong at i ng t o j o i n
t he c ora coid .
”
W i th thes e a dm i ss i ons i t s eem s t o u s th at rel ati v e
p os iti on and connexi ons ca nnot be app eal e d to as t es ts of h om ol ogy ,
no r as e vid e nce o f a n
y origi na l ty pe of v erte b ra .
I n no cl as s o f fac ts , th en d o w e fi nd a good fou nd ati on for the

,
hyp o thes i s of a n i d eal ty pi c al v e rt e b ra

“
. Th ere i s n o o ne c on
CRITI CIS M ON P ROF OWE N S T H E OR Y
’
A . .
5 61
cei v a bl e a ttri bute of th is a rchetyp a l form wh i ch i s h a bi tu ally reali s e d

by a ctua l v erte b rae Th e a ll eg e d group of tru e v e rt e b ral el em ents
.
i s not di s t i ngu i s h e d i n a ny s p e c ifie d w a y fro m b o ne s not i nclu d e d i n

it. Its m e m b e rs h a v e v a ri ou s d eg re e s o f i nc ons t a nc y ; are rarely
al l p res ent to g eth e r ; a nd n o o ne o f t h e m i s ess e nt i a l T h ey a re
.
s e ve ra l l y d e v e l op e d i n n o u nifo rm w a y : e a ch o f th e m m a a ri s e
y
e i t h e r o u t o f a s e p ara t e pi e ce o f c a rt il ag e or o ut o f a pi e c e c o n
,
t i nu o u s w i th th a t o f s om e o th er e l e m e nt ; a nd e a ch m a y be o ss i fie d
from m a ny i nd ep end ent p oi nt s from o ne or from n one
, , No t o nly
.
m a y t h e i r res p e c t i ve i ndi v i du a li t i es b e l os t by a b s e nc e or b c o n
y ,
fl u e nce w i th ot h ers but th e y m a y be d ou bl e d or tripl e d o r h al v e d , , ,
o r m a y be m ultipli e d i n one di re c ti o n a n d l o s t i n a n o th er T h e eu .
t i re grou p of typi ca l el em e nts m a y c oa l es ce i nto o ne s i m pl e b one

rep res e nt i ng th e w h o l e v e rt eb ra ; a nd e v e n as i n th e t erm i na l pi ece
,
o f a bi rd s ta il h a lf a d oze n v e rt e b ra w i th a l l t h e i r m a ny el e m e nts
’
- -
, , ,
m a y b ec o m e e nt i rely l os t i n a s i ngl e m as s L as tly th e res p ec ti v e

.
,
e l e m e nts w h e n p r es e nt h a v e n o fi xi t o f rel a t i v e p os i ti on : s u nd ry
, , y
o f t h em a r e fo und a rt i c ul a t e d t o v a ri o u s o th e rs th a n t ho s e w i t h
w hi ch th e y a re typi ca lly co nnec t e d ; th ey a re fre qu e ntly di s pl a c e d

a nd a tta c he d t o n e i h bo u ri ng v e rt e b rae ; a n d th e y a re e v e n r e m o v e d
g
t o qu i t e rem ot e p a rt s of th e s k el et o n I t s eem s to u s th a t if thi s
.
w a nt o f c o ngru i ty w i th th e fa c ts d o es not di sp ro v e th e hyp o t hes i s ,
no su c h hyp o th es is a d m i ts o f di s p r o of .
U ns ati s fa ctory as i s th e evid enc e i n th e eas e of th e tru nk and

t a il v e rt e bra , t o w h i ch w e h a v e h i t h e rt o c onfine d ou rs el v e s , i t i s fa r
w ors e i n th e c as e o f th e a ll eg e d c ra ni a l v e rtebra T h e m ere fac t
.
th a t th os e w h o h a v e c ont e nd e d fo r th e v e rt e b ra te s truct u re o f th e
s ku ll , h a v e di f fere d s o as t o ni s h i ng ly i n t h e i r s p ec i a l i nt e rp ret a ti o ns
o f i t , i s e no u g h t o w a rra nt g r ea t d o u b t as t o th e g e neral t ru th o f
th e i r th e o ry From P rofe s s or O w en s hi s tory of th e d octri ne of

’
.
g ene ral h om ol ogy w e g ath e r th at D um é ri l wrote up on l a t et e

,
“
cons id é r é e c om m e u ne v e rt eb re th a t Ki e h eyer, i ns t ea d of
“
c a lli ng th e s ku ll a v e rt e b ra , s a id e a ch v e rt e b ra m igh t be c a l l e d a
”
s k ul l ; th a t Ok e n re c og ni z e d i n th e sku ll thr ee v e rt e b ra a nd a
ru d i m e nt ; th a t P rofe s s o r O w e n h i m s el f m ak es o u t fo u r v e rt e b ra ;
th at G oeth e s id e a, a d opt e d a nd d e v el op e d by C a r u s , w as , th at th e
’
s k u ll i s c om p os e d o f si z e v erte b ra ; a n d th at G eo f fr e y S t H ila i re .
divid e d i t i nto seven D o es not th e fa c t th at di ffere nt com p a rat i v e

.
a n a to m i s t s h a v e a rr a ng e d th e s a m e rou p of b o nes i nt o o ne, thr ee,

g
i d v b l h o w th a t th e m o d e o f d e
f ,o u r s x , a n seven ert e ra s e
gm e nt s s .
t erm i na t i o n i s a rbi t ra ry , a nd th e c on clu s i ons a rri v e d a t fa nci fu l ?

May w e n ot p rop erly e nt erta i n grea t d ou b ts a s t o a ny on e s ch em e
b ei ng m ore valid th a n th e oth ers A nd i f ou t o f th es e c onfli cti ng
s c h e m e s w e a re as k ed t o a c c ep t on e, o u gh t w e n o t t o a c c e p t i t only
o n t h e p ro d u c t i o n o f s o m e th oro u ghly c o ncl u s i v e p ro of— s om e
82
r ig orou s tes t sh owi ng irrefra gably th a t th e oth e rs m u st be w rong

a nd t h i s a l o n e ri g h t E vid e ntly w h e re s u c h c ont ra d i c tory o pi ni ons
h a v e b e e n fo rm e d by s o m a ny c o m p e t e nt j u dg e s w e ou ght befo re , ,
d ecidi ng i n fa vou r of o ne o f th em t o h a v e a cl e a rnes s o f d em on ,
s tra t i o n m u ch e x c e e di ng t h a t re qu i re d i n a ny o rdi n ary c as e L et .
u s s e e w h e th e r P ro fe s s or Ow e n s u ppli es u s w i th a ny s u ch cl ea r
nes s o f d e m o ns t ra ti on .
T o b ri ng th e firs t or o c c ipi t a l s egm ent of th e s ku ll i nt o c orre

s o n d e nc e w i th th e id e a l t y pi ca l v e rt e bra P rofe s s or O we n a rgu es
p , ,
i n th e cas e o f th e fi s h th at t h e p a ra p oph y s es a r e di sp l a ced a nd

, ,
we dg e d b etw een th e ne u ra p o p h y s es a n d t h e n e u ra l s pi n e — r em o v e d
from th e h a m al a r ch a nd bu ilt i nt o t h e u pp er p a rt o f th e neu ra l

a rch .
Fu rt h e r he c ons id e rs th a t th e pleu ra p ophy s es a re tel eol og i
,
ca l l y com p ou nd . A nd th en i n a l l th e high er v e rt ebra ta h e a ll eg es

, ,
t h a t t h e h a m a l a rch i s s ep a ra ted from i ts c e nt rum t a k en t o a d i s ,
t a nce a nd t ra ns form e d i nt o th e s ca p ul a r a rc h
,
A dd t o wh i c h h e .
,
s a y s th a t i n m a m m a l s th e di s pl a c e d p a ra p o phy s es a re m ere p roc es s es
o f th e n eu ra p o p hy s es ( p 1 3 3 ) th e s e v e rt e b ra l e l em e nts ty pi c a lly
. . ,
b el ongi ng t o th e low e r p a rt o f th e c entrum a nd i n nea rly a ll c a s es ,
c o nfl u e nt w i th i t a re n o t o nly rem o v e d t o t h e fa r e nd s o f e l e m e nts

,
pla c e d a b o v e th e c e nt rum but h a v e b ec o m e e x o genous p art s of th em !

,
C onform i ty of th e s e cond or p a ri eta l s e gm e nt of th e c ra ni um wi th

th e p a tt e rn
- v e rt e b r a i s p,
ro d u ce d t h u s z— T h e p et ro s a ls a re excl u ded
as b e i ng p a rt i ally os s ifie d s e nse c a p s u l e s no t form i ng p a rts o f t h e
- -
,
t ru e v e rt e b ra l s y s t e m but b el o ng i ng t o th e
”
,
s pl a nc h no s k el eto n -
.
A c ent ru m i s a r tifici a l l y obt a i ne d by s a w i ng i n t w o t h e b one w hi ch

s erv e s i n c o m m o n as c e ntru m t o thi s a nd th e p r ec e di ng s e gm ent ; a nd
th i s th ou gh i t i s a d m i tt e d t h a t i n fis h es w h ere th e i r i ndi v id u ali ti e s ,
o u gh t t o be b e s t s e e n t h es e t w o h yp oth et i c a l c ent ru m s a re n ot
,
s i m ply c o al e s ce nt bu t c o nna t e
,
Ne xt a s i m il a r a r bi tra r y bi secti on
.
,
i s m a d e of cert a i n el e m e nts o f th e h a m a l a rch es A nd t he n t he .

,
“
p ri ncipl e of veg eta ti ve r ep eti ti on i s s till m ore m a ni fest i n th is arch

t h a n i n t h e o c c ipi ta l o ne e a ch pl e u ra p o ph y s i s i s d o ubl e ; e ac h
h a m a pophys i s i s d ou bl e ; a nd th e h a m a l s pi ne c ons i s ts of s i x pi e c e s !
Th e i nt erp ret ati o n of t he thi rd a nd fo u rth s egm ents b e i ng o f
th e sa m e g e ne ra l c h ara ct er nee d not be d eta il e d , Th e o nly p o i nt .
c a lli ng for rem a rk b ei ng th a t i n a ddi ti o n t o th e a b o ve v a ri ou s

,
m o d es o f g et ti ng o v e r a nom a li es w e find ce rta i n bone s referre d t o ,
t he der m o- s kel eton .
No w i t s e em s to u s, th a t
v en s upp os i ng no a ntag oni s t i nterpr e
e ~
t a ti ons h a d b e e n gi v e n a n hyp oth es i s reconc il a bl e w i th t he fa cts

,
o nly by th e a i d o f s o m a ny qu es ti ona bl e d e v i c es c o uld no t be c on ,
s id ere d s a ti s fac t ory ; a nd t h a t w hen a s i n th i s c a s e v a ri ou s co m , ,
ara ti ve a na t om i s ts h av e c o nt e nd e d for oth er i nte rp r e ta ti ons th e

p ,
c h aract er o f thi s o n e i s ce rt a i nly no t o f a ki nd t o w arra nt t h e re
e c ti o n of t h e o the rs i n i ts fa v ou r ; bu t ra the r o f a ki nd to m ak e
j
CRITICIS M ON P ROF OWE N S T H E OR Y
’
A . .
5 63
us d ou b t th e p o ss ibility of all s u ch i nte rp ret ati ons . Th e questi on

w h i ch n atural ly a r i ses i s w h eth e r by proce e di ng afte r thi s fas h i on
, ,
g rou p s of bone s m ight not be a rra ng e d i nto endles s typi cal fo rm s .
If w h e n a gi v e n e l e m e nt w as no t i n i ts pl a c e w e w ere a t lib erty t o

, ,
c o ns id er i t as supp r es sed o r conna te w i th s o m e n eigh b ouri ng e l em e nt

, ,
o r r em oved to s om e m ore o r le s s di s ta nt p os i t i o n — i f o n fi ndi ng a ,
b one i n e x ces s w e m igh t c ons id er i t now as p art o f th e der m o

, ,
s kel eto n now as p a rt of t h e sp l a nc hno s lcel eton n o w as tra nsp l a n ted

,
-
,
fro m i ts typi ca l p os it i on n ow as res u lti ng from veg eta ti ve r ep eti ti on

, ,
a nd n o w as a b o n e te l eol og ic a l l y com p ou nd
( fo r th e s e l as t t wo a re
i ntri ns i c ally different th ough ofte n u s e d by P rofess or O w e n as

,
e qu i v al ents ) — i f i n o th er c as e s ,
a b o n e m i ht b e re g a rd e d as
g ,
spu ri ou s
( p . or ag a i n as h a vi ng u su rp ed th e pl a ce of a n oth er ;
— if w e s a th e s e v a ri ou s lib e rt i e s w e re all ow e d u s w e s h ou ld
, y , ,
no t d es p a i r of reconc i l i ng th e fa cts w i th v ar i ou s d ia gram m at i c

typ e s b es id e s th at a d op te d by P rofess or Ow e n .
Wh en i n 1 8 5 1 w e a tt end e d a c ou rs e of P rofess or Owen s l ec

, ,
’
tu re s on C om p a ra t i v e Os te ol ogy b egi nni ng th ou gh w e did i n t h e a t ,
t i tu d e of di s c ipl es h ip o u r s c ep t i ci s m g r e w a s w e l i s tene d a nd rea c h e d

, ,
i t s c lim a x wh en w e c am e t o th e s ku l l ; th e re du ct i o n o f w hi ch to th e
v e rt e bra t e s t ru ctu re rem i nd ed u s very m u ch of t he i nte rp reta ti o n
,
of p roph e cy Th e d el i v e ry a t th e Roy a l S o c i ety of th e C rooni a n

.
, ,
L ecture for 1 8 5 8 i n wh i ch P rofess or H u x l ey c onfi rm i ng t he s t ate

, ,
m e nts o f s e v e ra l G erm a n a n a t o m i s ts h as s hown th a t th e fac ts of ,
e m b ry ol ogy d o n ot c ou nt ena nc e P rofes s or Ow en s v i ew s res p ec ti ng

’
th e fo rm a t i on of th e c ra ni u m h as i nd u ce d u s t o re cons id e r th e v ert e,
b ral th eory a s a w h ole C l os er e x am i nati on of P rofessor Ow en s

.
’
d octri nes as s et fort h i n hi s w orks h as c erta i nly not rem ove d th e

, ,
s c e p t i c i s m g e n e r a te d y e a rs a g o by hi s l ect u res On t h e c ontrary .

,
t ha t s c ep t i c i s m h as d eep en e d i nt o di s b eli e f A nd w e v e ntu re t o th i nk .
th at th e e v id e nc e a b o ve c i t e d s h ow s t hi s di s b eli ef t o be w a rra nte d .
Th e re re m a ns qu e s ti on— W h a t g eneral vi ew s a re w e to
i th e
ta k e res p ec ti ng th e v ert ebra t e s t ru ctu re I f th e h yp oth es i s o f a n
“
id eal t ypi c al v ert e b ra is not j u st ifie d by th e fa cts h o w a re w e ,
t o u nd ers t a nd th at d e gree of s im il a ri t y w h i c h v e rte b ra d i s pl a y ?

W e b eli e ve th e e x pla nati on i s not fa r t o s eek Al l th at ou r .
s p a c e w i l l h ere a ll o w i s a bri ef i ndi ca t i o n o f w h a t s e em s t o u s th e

,
na t u ra l vi e w o f t h e m att er .
P rofes s or Ow en i n c om m on w i th oth er com p ara t ive a na tom i s ts

, ,
re g ard s th e di v erg enc es o f i ndi vid u a l v ert e b ra from th e a v e ra ge

form a s du e t o a d ap t i v e m o difi cat i ons
,
I f h e re o ne v e rt e b ra l el e .
m ent i s l a rg ely d e v e l op e d w h il e el s e w h e r e i t i s s m a l l— if n ow th e
,
fo rm now th e p os i t i on no w th e d e gre e of c o a l es ce nce o f a g i v e n

, , ,
p a rt va ri e s ; i t i s th a t th e loca l re qu i rem e nts h a v e i nvolve d thi s

c ha n e T h e e nti re te a chi ng o f c o m p a ra tiv e os te ol o y i m pli es th a t
g .
g
5 64 A PP E N D I X B .
di ffere nces i n th e con di t i ons of th e res p e ct i v e v ert e b ra ne c e s s ita te

differences i n t h ei r s t ru c tures .
No w, i t s eem s t o u s th a t th e fi rs t s t e p t ow ards a rig h t co nc e p

t i on of th e ph e nom e na , i s t o r ec og ni ze th i s g enera l l a w i n i t s con v e rs e
a p li c a t i on If v e rt e bra a r e u nlik e i n p ro p o rt i on t o t h e u nlik e
p .
n ess o f th e i r c i rcu m s t a nce s , t he n, by i m pl i c a t i o n, th ey w ill be lik e i n

p ro port i o n to the lik e nes s of t hei r c i rcu m s ta nc es W hil e s u cces si v e
.
s e g m ents o f th e s a m e s k e l et o n, a nd o f di f fe rent s k el e tons are a ll i n

,
s om e res p e c ts m ore or l ess di f fere ntly a cte d o n by i nc id e nt fo rc e s ,

a nd a re t h ere fore re qu i re d t o be m ore or l es s di f fere nt ; th ey a re
a l l , i n oth e r res p ect s , s i m il a rly a c t e d o n by i n c id e nt fo rc es , a nd a r e
t here fore re qu i re d t o be m ore o r l es s s i m ila r . I t i s i m p o s s ibl e t o

d e ny th at i f di fferences i n th e m e ch a ni ca l functi ons of th e v ert e b ra
i nvolve di ffe rences i n th ei r form s then, c om m u ni ty i n th e ir m e ch a ni
c a l fu nct i o n s , m us t i n v ol ve com m uni ty i n th e i r form s . A nd as w e
k now th at th rough out the Ver tebr a ta g ene rally, a nd i n ea ch v ert ebra te
a ni m a l , th e v e rt e b ra , a m id a l l th ei r v a ryi ng c i rc u m s t a n ces , ha ve a
c ert a i n com m u ni ty o f funct i o n, i t foll o ws n e c es s arily t ha t th e y w ill
h a v e a c ert a i n ge ne ra l res e m bl a nce— th ere w ill rec u r th a t a v e ra g e

s hap e wh i ch h a s s u gg es te d th e no ti on o f a p a tt e r n v e rt e bra .
A gl a nce at th e fa c ts at o nc e s h ow s th e ir h arm ony w i t h t hi s

c on cl u s i o n. I n a n e el or a s na k e , w he re th e b o dily a cti o ns a re s u c h
as t o i nv o lv e grea t h om og en e i t y i n th e m ec h a ni c a l c on di ti o ns of th e
v e rt e b ra , th e s e ri es of th em i s c o m p a ra ti vely h om og e ne ou s On t he .
c o nt ra ry , i n a m a m m a l o r a bi rd w he re t h e re i s c o ns id era bl e h e t ero
,
e nei ty i n t he i r c i rcu m s t an c es , t h e i r s i m il a rity i s no l ong e r s o re a t

g g .
A nd i f, i ns t ea d of c om p a ri ng t h e ve rt e b ra l c ol u m ns of di ffere nt
a ni m a ls , w e c om p a r e th e s uc c es s i v e v e rt e b ra o f a n o n e a ni m a l , w e
y
r e cog ni ze th e s a m e l a w . I n th e s eg m e nt s of a n i ndi v id u al s pi ne ,
w h e re i s th ere th e g re a t es t di v e rg e nc e fro m t h e c om m on m e ch a ni c a l
c o n di ti ons ? a nd w h e re m a y w e t here fore e xp e c t t o fi nd th e wid es t
d ep a rt ure from th e a v erag e form O b vi ously a t th e t w o e x tre m i ti es .
A nd a c c ordingly i t i s a t th e tw o e x t re m i t i es t ha t t he ordi na ry
s t ru c tu re i s l o s t.
S t ill cl e a re r b e c om e s th e tru th of thi s v i e w , wh e n w e cons id e r th e

g enes i s of th e v e rte b ra l colu m n as dis pl ay e d th rough ou t t he as cend
i ng gra d es o f th e Ver tebr a ta . I n i ts fi rs t e m b ry oni c s ta g e, th e s pi ne
i s a n u ndi v id ed colum n o f fle xibl e s u b s t a nce . I n th e e arly fis hes ,
w h il e s om e o f th e p erip h era l e l em ents of t h e v e rt e bra w e re m a rk e d
o ut , t h e c e ntra l a x i s w as s till a c o nt i nu o u s u nos s ifi e d cord . A nd
thu s w e h a v e g oo d rea s on fo r th i nki ng th a t i n th e p ri m i ti v e v ert e
b rate a nim al , a s i n t he e xi s ti ng A m p hi oxu s, th e noto chord w as p e r
s i s t e nt
. T h e p rod u ct i on o f a h i g h er, m ore p ow erful, m ore a c ti v e
c re a tu re of t h e s a m e typ e by w h a te v er m et ho d i t i s c o nce i v e d t o
,
h a v e t a k en pl a ce, i n v ol v e d a ch a ng e i n th e not o ch ord al s tru ct ure .
Grea t e r m u s c ul a r e nd o wm e nts p res upp os e d a fi rm e r i nte r na l ful cru m

CRITI CIS M ON P ROF OW E N S T H E OR Y
’
A . .
5 65
— a l ess yi eldi ng c entral a xi s On th e o th er h a nd for th e centra l

.
,
a x i s to h a v e b e c om e fi rm e r w h i l e r em a i ni ng cont i nu ou s w oul d h a v e ,
e nt a il e d a s t if fness i ncom p at ibl e w i th th e cre a tu re s m o v em ent s

’
.
H e nce i nc reas i ng d e ns i ty o f th e c e ntral a xi s ne c e s s a rily we nt h a nd

,
i n h a nd w i th i ts s eg m e nt a ti on for s t reng th os s ifi c a t i o n was re ,
q ui re d for fle x ibility di vi s i on i nto p a rts T h e pro d uc ti on of v e r

, .
t e b ra res ult i ng th u s th ere o bvi o u s ly w o uld a ris e a m o ng t h e m a

,
g enera l lik e nes s due to the s i m ila rity i n th ei r m ech ani c a l c o ndi
,
t i o ns a nd m or e e s p ec i a lly th e m u s c ul a r for c e s b eari ng o n t h em

, .
A nd t h e n o bs e rve l as t ly tha t w h ere as i n t h e h e ad th e t erm i na l

, , , ,
p os i ti on a nd th e les s s p a ce for d e v elop m ent of m us cles e nta ile d ,
s m a ll er l a t e ra l be ndi ngs th e s e g m e nt a t i on w ou l d na tu ra lly be l e s s

,
d ecid ed less regula r a nd would be l ost as w e a pp roa ch e d th e

, ,
fro nt of th e h ea d .
B u t i t m a y b e repli e d th i s h yp oth es is d oe s not e x pl a i n all the

, ,
fact s . I t d oe s not t el l u s w hy a bone w h os e fu nct i on i n a gi v en

a ni m al re qu i res i t t o be s olid i s fo rm e d n ot o f a s i ngl e pi ece bu t by
, ,
th e coa l e s c e nc e of s e v e ra l pi ec es w hi c h i n o th er c re at u res a re s e a
,
p
ra t e ; i t d o e s n ot a ccou nt for th e fre qu e nt m a ni fes ta t i ons o f u ni ty
o f pl a n 1 n defi a nc e o f t el eol ogi c al r e q ui re m e nt s Th i s 1 8 qu i te tru e .
B ut i t i s not tru e as P ro fe s s or Ow e n a rgu es re s p ecti ng s u ch cas es

, ,
t hat if th e p ri nc ipl e of s pec i a l a d ap t a ti on fa il s t o e xpl a i n th e m a nd

“
,
w e rej ec t th e id ea th at the s e c o rr es p ond e nc es a re m a nifes ta ti ons of

s om e a rch e typ a l e x e m pl a r o n w h i c h i t h as pl eas e d t h e C r ea to r t o
,
fra m e c ert ai n o f h i s li v i ng crea tures th ere rem a i ns only th e a lt erna ,
t i ve t h a t th e org ani c a tom s h a v e c on cu r re d fo rt u i t ou s ly t o p ro du ce

s u ch h arm o n y
”
. T h i s i s not th e o nly a lt ern a ti v e : th e re i s a noth er ,
w h i ch P rofes s or Ow e n h as o v erl ook e d I t is a p e fe ctly te nabl e

r .
s u pp os i ti o n th a t a l l h igh er ve rt e bra t e fo rm s h a v e a ri s e n by the s u
p e rp osi ng of a dap ta ti on s up on a dap ta ti ons E i th e r of th e t w o a nta .
o ni s t c os m o g oni es c o ns i s ts w i th th i s s u pp o s i t i on I f o n th e o n e
g .
,
h a nd w e c o nce i v e s p ec i es t o h a v e res ult e d from a cts of s p ec i a l

,
c re a ti o n ; t h en i t i s qu i te a fa i r as s u m pti o n th a t to p ro d u ce a high er
v e rte b ra te a ni m al th e C rea t or did n ot b egi n afres h but t ook a
, ,
low er v ert ebra t e a ni m al and s o far m o difie d i ts pre exi s ting p a rt s

,
-
as t o fit th em for th e n e w re qu i rem e nts ; i n w h i ch cas e th e ori g i na l
s t ru ct ure w o uld s h o w i t self th rou h th e s u p e rp os e d m o difi c a ti o ns

g
.
I f on th e o th er h a nd w e conc e i v e s p e ci es t o h a v e res u lte d by

, ,
gra d u al di ffe re nti a ti ons u nd er the i nflu ence of ch a nge d c ondi ti ons
t h e n i t w ou ld m a nife s tly foll o w th a t th e high er h e te rog e ne ou s
, ,
form s w ou ld h ear t ra ces of t h e l ow er a n d m ore h o m og en eou s form s

,
from w hi ch th e y w ere e v ol v e d .
Thu s b e s id e s findi ng th at th e h yp oth es i s of an i d e al t ypi cal “
v ert eb ra i s i rre c oncila ble w i th th e fa cts w e fin d th at th e fa cts are ,
i nterpreta ble w ith ou t gra tui tou s as s u m pti ons Th e a vera g e com .
m uni ty o f fo rm wh i ch v ert e b ra di s p l a y i s e xpli ca bl e a s res ulti ng ,

5 66 A PP E N D I X B .
from na tu ral cau s es A nd th os e typi cal s im il ari ti es whi ch are

.
t racea bl e u nd e r a d a p t iv e m odifi ca ti ons m u s t ob vi ou sly e x i s t i f

, ,
t hrough ou t c rea ti on i n g enera l th ere h a s g one on t hat c onti nu ou s

,
s up e rp os i ng of m odifi c a t i ons u p on m odifica ti ons wh i ch g o es o n in
e ve ry u nfoldi ng org a nis m .
[I m ight with prop ri ety h a v e a dd e d to th e foregoi ng cri ti ci sm s ,
t h e re m a rk th a t P ro fes s or Ow e n h a s i ndi rec tly c o nferre d a g re a t

b enefit by th e ela b orate i nv es tig ati ons h e h as m a de w ith th e vi e w of
es ta bli s h i ng h i s hyp oth e s i s
. H e h as hi m s elf v e ry c onclu s i v ely
p ro ve d th a t t he t el eologi cal i nte rp reta ti o n i s qu i te i rre conc ila ble
w i th t he fa cts . I n ga theri ng t og e th e r e v id enc e i n s up port of hi s
o wn c o ncep t i o n of a rch etyp a l form s h e h as di s cl os e d a dve rs e
,
e v id en ce w hi ch I th i nk sh ow s hi s conc e p t i on t o be u nt ena bl e .
T he res ul t i s th at th e fi eld i s l eft cl ea r for th e hyp oth es i s of E ve

l uti on as th e o nly t e na bl e o ne ] .
A P P E ND IX C .
[ From the TRA NS A C TI ONS or r u n L I NNE A N Soc rs r x VOL . xxv .

]
XV . On Ci rcu l a ti on a nd the F or m a ti on of Wood i n P l a n ts B y .
H E R BE RT SP E N C E R, E s q Com m u ni ca ted by GE O R G E B U S K ,
.
E sq , . S ec L S . .
Rea d Ma rch 1 st , 1 8 6 6 .
O P I N IO N S p e cti ng th e fu ncti ons of th e v a scul ar ti s su es i n pla nts

res
a pp ea r t o m a k e bu t li tt l e p ro g res s t o w a rd s a reem ent T h e s u pp o

g .
s i t i o n th a t th es e v e s s el s a n d s t ri ngs of p a rti a lly u ni t e d c e ll s li ne d -

,
w i th s pi ra l a nnu l a r re ti cu l at e d o r ot h er fra m e work s a re ca rri ers

, , , ,
o f t h e pl a nt j u i c es i s o bj e ct e d t o o n th e g rou nd th a t th ey oft en
-
,
c onta i n a i r : a s th e pre s e nc e o f a i r arre s ts th e m o v em e nt o f bl oo d
t hrou gh a rt eri es a nd v ei ns i ts p res en c e i n t h e d u c t s of s te m s a nd

,
p eti ol es i s a s sum e d to u nfit them a s ch a nnels for s a p On the .
o th e r h a nd th a t th es e s tru c tu re s h a v e a res pi ra t o ry o fli ce a s s om e
, ,
h a v e th ou ght i s c e rt a i nly not m ore te na bl e s i n c e i f th e p res enc e

, , ,
of a i r i n th em neg a ti v e s th e b eli ef th a t t he i r fu n c ti o n i s t o di s
t ri bu te liqu id th e p res ence of li qu id i n th em e q u a lly neg a t i v es t h e
,
b eli ef th a t t h ei r fu ncti on i s t o di s tri but e a i r Nor c a n a ny bette r .
d e fence be m ad e for the hyp oth esi s whi ch I find p rop ou nd e d th at ,
to gi v e s tre ngth t o th e p a r e nchy m a

”
t h es e p a rts s erv e T u b es
“
.
w i th fen es tra t e d a n d ret i cu l a t e d i ntern al s k el et ons h a v e i nd ee d , ,
s om e p ow e r o f s u pp o rt i ng th e t i s s u e thr ou gh w h i ch th e y p a s s ; bu t
t ub es li ne d w i th s pi ra l th rea d s c a n yi eld e x trem ely li ttle s upp ort ,
w h il e tu b e s li ne d w i t h a nnuli or s pi r al s a l t erna t i ng w i th a nnu li ca n

, ,
yi eld no s u pp ort w h ate v er Th ough a l l th es e typ es of i ntern al

.
fra m e w ork a re m ore or l es s e ffi ci e nt for p r e v e nt i ng c l os u re by

l a teral p ress u re th e y a re s om e of th em qui t e u s eles s for h olding
,
up th e m as s th rou gh w h i ch th e v es s el s p as s ; a n d t h e b es t of th e m
a re for th i s p u rp o s e m ech a n i c a lly i nfe ri or t o th e s i m pl e cyli nd e r .
Th e s a m e qu a nti ty of m a tt er m a d e i nt o a c ont i nu ou s t ube w ou ld b e

m o r e e ffe ct i v e i n gi v i ng s ti f fness t o th e c ellu l ar t i s s u e a round i t .
In th e a bs e nce of a ny feas i bl e a l terna ti v e th e hyp oth e s i s th at ,
t h e s e v es s e ls a re di s t ri bu t ors o f s a p c l a i m s r e c ons id e rati on Th e .
o bj ec ti o ns a re n o t I th i nk s o s e ri o u s as the y s e e m
, , Th e h a bi tu a l .
5 67
5 68 A P P E ND I X 0 .
p res ence of ai r i n th e d ucts th at tra vers e w oo d c an s carcely be ,
h eld a no m al o u s if w he n t he w oo d i s form e d t h ei r fu ncti on c ea s es .
T h e c a n al s w h i ch ra m i fy throu gh a S ta g s h orn c onta i n a i r a ft e r

’
,
t h e S t a g s h orn i s fully d ev el op e d ; but i t i s not t h ere by re nd e r e d

’
d o ubtful w h eth e r i t i s th e func ti o n of a rt eri e s t o c on vey bloo d .
A ga i n th a t a i r s h ould fre qu entl y be fo u nd e v en i n th e v ess e ls o f

,
p eti o l es a nd l ea v es w i ll not a pp ear rem a rk a bl e wh en w e c a ll t o

,
m i nd th e c o ndi t i o ns t o wh i ch a l e af i s s u bj e ct E v a p o ra ti o n i s .
g oi ng on from i t T h e thi nner liqu id s p a ss by os m os e o u t of th e

.
v es s els i nto th e ti s s u e s c ont a i ni ng the liqu id s th i ck e ne d by e v a p o ra

t io n . A nd a s th e v es s el s a re thu s c onti nu a lly d ra i ne d a d ra ug ht i s ,
m a d e u p o n th e liqu id c o nt a i ne d i n th e s t em a nd roots S up pos e .
t ha t th i s d ra u gh t i s u nu s u a lly g rea t o r s u pp o s e t h a t a rou n d th e ,
ro o ts th e r e e x i s t s n o a d e qu a t e s u pply o f m o i s tu r e A s t a te of .
ca pill ar y t e ns i o n m u s t res u l t — a t end e nc y o f th e liqu id t o p a ss i nt o
t h e l e a v e s r es i s t e d b el ow by liq u id c oh es i on N o w h a d th e v ess el s .
,
i m p erm ea bl e c o ats only th ei r upp e r e x tre m i ti es w ou ld u nd er th es e

,
c on di ti ons be sl ow ly e m p t i ed B ut the i r c oa ts i n c om m on w i th a l l
.
,
th e s u rrou ndi ng t i ss u e s are p erm e a bl e by a i r

,
H e nce u nd er thi s .
,
s t a t e o f c apill ary t ens i o n a i r w ill e nt er ; a n d a s t h e upp e r e nd s o f

,
t h e t u b e s bei ng b oth s m all e r i n di a m et er a nd l ess p o ro us t h a n th e

,
lo we r w ill reta in th e liqu id s wi t h g reater t ena c ity the a i r w ill

, ,
e t er t e
n h w id e r a n d m or e p oro us t u b es b e l o w— th e d u c ts o f t he
s t em a nd b ra nch es Thu s th e e ntra nc e of a i r no m ore p ro ves th a t
.
th es e d u c t s a re not s a p c a rri ers t ha n d o e s th e e m p ti nes s of t ropi c a l

-
,
ri ver b e d s i n th e d ry s ea s on p ro v e th a t t he y a re not c h a nne l s for

-
w a t er Th ere i s h o w e v er a di ffi culty w h i ch s e em s m ore s e ri ou s

, , .
I t i s s aid th a t a i r wh e n p re s ent i n th es e m i nut e c a na ls m u s t be a

, ,
g rea t obs t a cl e t o t he m o v e m ent of s a p th rou gh th e m Th e i n v e s ti .
a ti o n s o f J a m i n h a ve s h o w n th a t bu bb l es i n a c a pill ary t u b e res i s t

g
t h e p a s sa g e o f liq u id a nd th a t th ei r re s i s t a nc e b ec om e s v ery gre a t
,
w h e n th e bu bbl es a re num e rou s — rea ch i ng i n s om e e x p eri m e nts as , ,
m u ch a s t h ree a t m os ph eres Ne ve rth e l ess th e i nference th at any

.
s u ch re s i s t a nc e i s o ffe r e d by t h e a i r bu b bl es i n th e v es s el s o f a -
pl a nt i s I thi nk a n e rroneous o ne W h at h app ens i n a c a pilla ry

, , , .
t u be h a v i ng i m p er v i o us s id e s w i th w h i c h th es e e xp e ri m e nts we r
, e
m a d e w ill by no m ea ns h a pp e n i n a c a pill ary tu b e h a v i ng p e r v i ous

,
s i d es . A ny p res s u re b rou ght t o bea r o n th e c ol u m n of li qu id c o n

t a i ne d i n th e p o r ou s d u c t o f a pl a nt m u s t qu i ckly c a u s e th e e xpu l
,
s i o n o f a c o nta i ne d a i r b u b bl e th ro u g h th e m i nut e o p e ni ngs i n th e

-
c o a t s of th e d uct Th e gr e at er m ol ecu l a r m o bili ty o f g as es th a n

.
liq uid s i m pl i es th at a i r will p as s out fa r m ore rea dily th a n s ap

,
.
W hils t th erefore a s ligh t t ens i o n on th e c o lu m n of s ap w ill cau s e

, ,
i t t o p a rt a nd th e a i r t o e nt e r a s light p re s s ure u p on i t w ill forc e

,
o ut t h e a i r a n d reu ni te th e di v id e d p a rts o f th e c ol um n .
T o o b ta i n d a ta for a n opi ni o n o n thi s v e x e d qu e s ti o n I h a v e ,

C IRCU L A T I ON A ND F O RMA T ION OF WOOD IN P L A NTS .
5 69
l a tely b een e xp eri m enti ng on th e a b s orp ti on of dy es by pl ants S o .
far as I c a n l e a rn e xp e ri m e nt s of t h i s ki nd h a v e m os t if not all of

, ,
t he m bee n m a d e o n s te m s a nd as i t w ou ld s eem from th e res u lts

, , , ,
o n s te m s s o fa r d e v e l op e d as to c o nt a i n a l l t h e i r c h a rac t eri s t i c
s tru ctu re s . Th e firs t e x p eri m e nts I m a d e m y s e lf w e re o n s u ch

p a rts a nd yi eld e d e vid e nce th a t s er v e d but littl e to elu c i d a te
,
m a t t ers . I t w as only afte r t ryi ng lik e e x p eri m ents w i th l ea v es of

di fferent a g es a nd diffe rent cha ra cters a nd w i th u nd e v elop e d a xes , ,
as w e ll as w i t h a x e s o f s p ec i al ki nd s t h a t c om p reh e ns ibl e res u l ts

,
w e re rea ch e d ; a nd i t th en b eca m e m a nifes t th a t t h e a pp ea r a nces

p res ent e d by o rdi nary s t em s w h en t hus t e ste d a re i n a great d egree ,
m is l e a d i ng L e t m e b ri efly i ndi c a te th e differe nces

. .
If a n a d u l t s h o ot of a t ree o r s h ru b b e c ut off a nd h a v e i ts l ow e r ,
e nd pl a c e d i n a n a l u m e d d e c o ct i on o f l ogw o o d o r a dilu t e s ol u t i o n
*
o f m ag e nt a th e dy e w ill i n t h e c ou rs e o f a fe w h ou rs as c end t o a
, , ,
d i s t a nc e v a ryi ng a cc o rdi ng t o th e ra t e of e v ap ora t i on from th e

le a v es On m a ki ng l ongi tu di na l s ecti ons of th e p art t ra ve rs ed by
.
i t t h e dye i s fo u nd t o ha v e p e ne tra t e d e x te ns i v e t ra cts o f th e

,
w o o dy t i s s u e a nd o n m a ki ng t ra ns v ers e s ect i ons th e o p eni ng s o f ,
t he d u c ts a pp e a r as em p ty s p a ces i n t h e m i ds t of a d eeply c ol ou re d -
p ros e nchy m a I t w ould thu s s eem t hat the liqu id i s c a rr i e d u p the

.
d ens er p a rt s o f t he v as cular bu ndles n egl ecti ng th e c a m bi um l ay e r ,
ne gl e c t i ng th e c e ntra l pi th a nd negl ec t i ng th e S pi r a l v e ss el s o f th e
,
m e d u ll a ry s h e a t h A pp a r ently th e s u b s ta nc e of t h e w oo d h as
.
af ford e d th e re a di es t c h anne l W he n h ow e v e r w e e x a m i ne th e s e
.
, ,
a pp ea ra n c e s c ri ti c a lly w e fi n d re as o ns for d ou bti ng t hi s c o ncl u s i o n

, .
If a t ra ns v erse s ect i on of th e l o w e r p a rt i nto w h i ch t h e dy e p ass e d,
fir s t a nd h as r em ai ned l o ngest be c om p a re d w i th a t ra ns v ers e s e c

,
ti o n of th e p a rt w hi ch t h e dy e h as bu t j u s t re a ch e d a m a rk ed ,
di fferenc e i s v i s i bl e I n th e one c as e th e w hole of th e d ens e ti ss u e

.
i s s t a i ne d ; i n th e oth er c as e i t i s n ot T hi s u ne v en di s tri buti on of

.
s t a i n i n th e p a rt w h i c h th e dy e ha s i ncom pl e tely p erm e a t e d i s n ot
a t r a nd o m ; i t a d m i ts o f d efi ni te d e s c rip t i on A t ol era bly r e gu l a r

.
c o nti nu o u s ri ng o f col ou r di s ti ngu i s h es th e ou t er p a rt of th e w oo d
fro m t h e i nne r m as s i m plyi ng a p as s ag e of l iq u id up th e e l ong a t e d

,
c ell s n e x t th e c a m bi u m l a y er A nd t h e i nner m ass i s colo ure d m o re

.
ro u nd t h e m o u th s o f th e pi tt e d d u c ts th a n el s ew h e re : t h e d ens e
t i s s u e i s d a rk es t c l os e t o th e e dg es of th es e d u c t s ; th e c ol o u r fa d es
a w a y gra d u a lly o n re cedi ng from th e i r e dg es th e re i s m os t c ol ou r
w h ere th ere a re s e v e ra l d u cts tog eth er ; a nd t h e d ens e t i ss u e w hi ch
Th e s e two dye s h a v e a ffini t i e s for di fferent com p onents of the t i s s ue s ,

a nd m a y b e a d v a nt a g eo u sl y u s e d i n di f ferent c as e s Ma g e nt a i s ra pi dl y
.
t a k en up b y wo od y m a tt e r a nd oth er s econda ry de po s i t s ; w h il e l og w oo d
col ou rs t he c e l l -m e m b ra ne s , a n d t a k e s bu t r el u cta nt l y t o t he s ub st an ce s s ei ze d
by m a g en ta By t r yi ng both o f th em on t h e sa m e s t ru c ture , we m a y gu a ni
.
o urs el v es a ga i ns t a ny e rror a ri s i ng from s el ec ti v e c om bi na ti on .

5 70 A P P E ND I X 0
.
i s ful ly dy e d for s om e s pace , i s th at wh i ch li e s b etw een t w o or m ore

d u cts T h es e a re i ndi ca ti ons that wh il e t he l a y e r of pi tt ed c ells
.
n e x t th e c a m bi u m h a s s e rv e d as a ch a nne l fo r p a rt o f t h e liq u id , t h e
res t h as a s c e n d e d t h e pi tt e d d u c ts , a nd o oz e d o u t o f t h es e i nto th e
p ros enchym a a round A nd th i s c onclu s i o n i s c onfirm e d by th e
.
c o nt ra s t b e t w e e n th e a pp e a ra n ces of th e l ow e st p art o f a s h oot
u n d er di f fe rent c ondi ti ons . For if, i ns t e a d o f a ll o wi ng t h e dye

t i m e for o oz i ng th rou gh th e p ros enc hy m a , th e e nd o f th e s h o ot b e
j us t di pp e d i nt o th e dye a nd t a k en o u t ag a i n, w e find on m aki ng ,
t ra ns ve rs e s e c ti ons of t h e p a rt i nto w h i ch th e dye h as b ee n ra pidly

t a k e n u p, th a t, th ou gh i t h as di ffu s e d to s om e di s ta nc e rou nd th e
d u cts , i t h as left tra ct s of w oo d b e tween th e du cts u ncol oure d— a
diffe renc e wh i ch w ou ld not e x is t h a d th e as cent b een throu gh t he
s u b s t a nc e o f t h e w o o d . E v en s till s t ronge r i s th e c onfirm a ti o n
o b ta i n e d by u s i ng o n e dye a fte r a n oth e r . If a sh oo t th at h as a h
s orb e d m a g e nta for a n h ou r h e pl a c e d fo r five m i nu t es i n t h e l og
w o o d d e c oc t i on , t r a ns v ers e s e ct i ons of i t t a k e n a t a s h o rt di s ta nc e
fro m i ts e nd s h o w th e m ou th s of th e d u ct s s urrou nd e d by d a rk
s t a i ns i n th e m id st o f t h e m u ch w id e r r e d s ta i ns
'
B as e d o n th es e c om p ari s o ns o nly , th e i nfe r e nce p o i nt e d ou t h as

li ttl e w eight ; but i t s w eig ht i s i ncre as e d by th e res u lts o f e x peri
m ents o n qu i t e y o u ng s h o o ts a nd s h o ots th a t d e v el o p e v ery li ttl e
,
w ood . T he b eh a vi ou r of th es e c orres p o nd s p e rfectly w i th th e e x

e ct a ti o n th a t a liq u id w ill a s ce nd c a pill ary t u b es i n p refe ren ce to
p
s i m pl e cell u l a r t i s s ue or t i ss u e n ot di f fe re nt i a te d i nto co nt i nu o us
c a na ls . Th e va scul a r b u ndl es o f th e m e d u l lary s h ea th a re h ere
t he o nly c h a nnels w h i ch th e c ol ou re d liqu id t a k es . I n s e cti o ns o f
t h e p arts u p to wh i c h t h e dy e h as bu t j u s t rea c h e d , th e s pi ra l , fen es
trat e d, s ca l a ri form , or oth e r v es s e l s c onta i ne d i n th es e b un dle s a re
a l o ne co l ou re d , a nd l o w e r d ow n i t i s o nly a fte r s o m e h ou rs th a t
s u ch a n e xu d a t i on o f dye t a k e s pl a ce a s s u f fi c e s p a rt i ally to colou r

t h e oth er s u bs t a nce s o f th e b u ndl e . Furth e r, i t i s to be note d th a t
a t t he t erm i na t i ons o f s h e e t s , wh e re th e v es s el s a re but i n co m pl etely
form e d ou t of i rre g ul a rly j o i ne d fib r ou s c ells w hi c h st ill re t a i n

-
t he i r origi na l s h a pes , t he dye r uns u p th e i nc i pi e nt v es s el s a nd

d o es not c olour i n th e s m a llest d e g ree th e s u rr o u ndi ng ti s su e .
E xp e ri m e nts w i th l e a v es b ri ng o u t p a ra ll e l fa c ts On pl a c i ng i n
.
a d ye a p et i ol e of a n a d ul t l e a f o f a t ree , a n d p u tt i n
g i t b e fore th e
fi re t o a cc el era te e v a p ora ti o n, t he dye w ill be fou nd t o as cend th e
m id rib a nd v ei ns a t v a ri o u s r a t es , u p e v e n t o a fo ot
p e r h ou r
. At
fi rs t i t i s c onfine d t o th e ves s els but by the ti m e i t ha s rea ch ed th e
p oi nt of th e l ea f, i t w ill c om m only b e s ee n tha t a t the l ow er p a rt i t
h a s d i ffu s e d i t s el f i nt o t h e s h ea th s o f th e v es s els
. I n a qu i te y ou ng
l ea f fro m th e s a m e s h o ot, w e find a m u ch m ore ri gorous r est ri ct i on
o f t h e d e t o t h e v es s e l s On m a ki ng o bliq ue s ec t i o ns o f i ts p e ti ol e ,
y .
m id ri b, a nd v e i ns , t h e v e s s el s h a ve t h e a pp ea ra nce o f g rou p s o f
CIR C UL A T I ON A ND FOR MA TI ON OF WOOD I N P L A NT S .
5 71
sh ar ply d efine d col oure d rods i m b e dd e d i n th e green p ros en chy m a ;

a n d t h i s m ark e d c o nt ra s t c o nti nu es w i th s c a r cely a n a pp r e c i a bl e
c h a ng e afte r pl e nty of ti m e h as b ee n a ll o w e d fo r e x u d a t i o n .
T h e fa cts thu s grou p e d a n d th u s c ont ras t e d s eem a t firs t s ight , ,
t o i m ply t h a t w h i l e th e y a re y ou ng t h e c oa ts of th es e r a m ifyi ng
c a na l s li ne d w i t h s pi ra l or a lli e d s t ru ct u res a re n o t re a dily p e rm s
a bl e bu t t ha t b ec om i ng p oro us as t h e y g ro w o ld t h ey a ll o w th e
, , ,
liq uid s th ey c arry t o es c a p e w i th increas i ng fa cili ty ; a nd h en ce a

p os s i bl e i nterp reta t i o n of the fa c t th a t i n t he old er p arts th e s ta i n
, ,
i ng of th e t i s s u e a rou n d th e vess el s i s s o ra pid as to s u gg es t th a t th e

dye h as as c en d e d di re ctly th rough th i s t i s s u e wh ereas I n th e y ou nger ,
p a rts the re v ers e a pp ea ra nce ne ces s ita t es the re v e rs e c onclus i on .
B u t now i s th is differe nc e d e t erm i n e d by di ffe re nc e of a g e o r i s i t

, ,
o th e r wi s e d e t erm i ne d ? T h e e vid e nc e as p r es e nte d I n o rdi na ry s te m s
a n d l ea v es s h o w s u s t h a t t h e p a rt s o f t h e v as cu l a r s y s t e m a t w h i c h
th ere i s a r a pid es c a p e o f dye a re n ot s i m ply old er p a rts bu t a re ,
p a rts wh ere a d ep osi t of wo o dy m a tt er i s ta ki ng pla ce I s i t th e n .

, ,
t h a t th e i nc r e a s i ng p e rm ea bi li ty of th e d u c t s i ns t ea d of b ei ng ,
di rec tly as s o ci a te d w i th th ei r i n c rea s i ng ag e i s di rectly a ss oc i at e d ,
w i th th e i nc reas i ng d ep os i t of d e ns e s u bs t a nce a rou n d th em

T o g e t p ro of th a t t hi s la s t c o nne x i on i s t h e t ru e o ne w e h a v e ,
bu t t o ta k e a cl as s of c as es i n w h i c h w oo d i s fo rm e d o nly t o a s m all
e x t e nt I n su ch c as es e x p e rim e nts s h o w u s a far m o re g ener a l a n d
.
c onti nu e d li m i ta t i o n of th e dy e t o t h e v ess el s O rdi na ry h er bs a nd

.
v e ge ta bl es wh en c ontr as te d w ith s h ru b s and tre es illu s tra te thi s ;

, ,
as i ns t a n c e th e p et i o l e s o f C e l ery or o f t h e com m on D oc k a n d th e
, ,
le a v es of C a b bag es or Tu rnip s A nd th en i n v ery s u c cu l ent pl a nts

.
,
s u c h as B r yop hy l l u m ca l y ci nu m K a l a nc ho e r o tu nd ifo l i a th e v a ri ou s
'
, ,
s p e ci e s o f Cra ssu l a Co ty l ed on K l ei n i a a nd o th ers of lik e h a bi t th e

’
, , , ,
d u cts of ol d a nd y ou ng l ea v es alik e ret a i n th e dye v e ry p ers i s te ntly

th e c onc o m i ta n t i n th e se c a s e s b e i ng th e s m a ll a m ou nt o f p ros e n
ch m a a r ou n d t he d u c ts o r th e s m a ll a m ou nt o f d e p os i t i n i t or
y , ,
b oth More c onc lu s i ve yet i s th e e v id enc e w h i ch m eets u s w h en w e

.
turn from v e ry s u c cu l e nt l e a v e s t o v ery s u c cu l e nt a x es Th e t en d er .
y ou ng sh oots ofK l ei ni a a n te eu p hor bi u m or E u p hor bi a M a u r i ta ni ca

-
, ,
w h i c h for m any i n ch e s o f th ei r l e ngth s h a v e s ca r c e ly a ny li gn eou s

fibres s h ow u s s ca rcely a ny es cap e of th e c ol ou re d liqu id from th e
,
v es s el s o f the m e du ll a ry s h ea th S o t oo i s i t w i th S tap el ia
.
, ,
E uf on i a a pl a nt of a n oth er o rd e r h a v i ng s oft s w oll e n a x es

, ,
A nd .
th e n w e h a v e a r ep e ti ti o n o f th e lik e c o nne x i o n of fa ct s t hr ou gh out

th e Ca c ta cece th e m os t s u c c u l ent s h ow i ng u s th e s m a l l e s t p erm s
a bili ty o f th e v es s el s I n t w o s p e c i es of R hip s a l i s i n t w o s p e c i es o f
.
,
Cer eu s a n d i n t w o s p ec i es of M a m m i l l a ri a w h i ch I h a v e t ri e d I
, , ,
h a v e fo und th i s s o M a m m i l l a r i a g r a ci l i s m ay be na m e d a s e x
.
em l if i n t h e r el a t i on u nd e r i ts e x t r e m e form I nto o ne o f th es e
p y g .
s m all s ph ero id a l m a s s es th e dye as c e nd s t hr o u gh th e l a rg e b u ndl es

,
5 72 A P P E ND IX 0 .
of s pi ra l or a nnul ar du cts or c ells p a rt i ally u ni te d i nto s uch d u cts

, .
c ol o u ri ng t h e m d eeply a n d l ea v i ng th e fee bly m a rk e d s h e a th o f

,
-
p ros enchy m a tog eth er wi th th e s u rrou ndi ng w a tery c ellu lar ti s s u e

, ,
p e rfe ctly u ncolou re d .
Th e m os t c o nc lu s i v e e v id e nce h o w e v e r i s furni s h e d by t h o se , ,
Ca c ta cece i n w h i ch th e tra ns i ti on from s u c cul ent to d ens e ti s s u e

t a k e s pl a ce v a ri a bl y a c c ordi ng as l oc a l c i rc u m s t a n ces d et e rm i ne
,
.
Op u n ti a yi eld s goo d e x a m pl es If a pi e c e o f i t i n c lu di ng o ne of
.
th e j o i nts a t w h i ch w oo d i s b e gi nni ng to fo rm he a ll ow e d to a b s orb ,
a c ol ou re d liqu id th e liqu id runni n g u p th e i rre gu l a r bu ndl es of

, ,
vess els a nd i nt o m a ny of thei r m i nu te ra m ificati ons i s re stri ct ed t o ,
th es e w h ere t h ey p as s th rou gh th e p a re nch y m a form i ng th e m as s of

th e s t em but nea r th e j oi nt s th e h a rd e ne d t i s s u e a rou nd th e v es s els
i s c o l o u re d I n o n e of t hes e fl es h y g ro wth s w e g et c l ear e v id e nce
.
t h a t th e e s ca p e of th e dye h a s no i m m e di a t e d ep e nd en c e o n t h e ag e
o f th e v es s el s s i nc e i n p a rts of th e s te m th a t a re a lik e i n a e s om e
, , g ,
o f t h e v ess e l s ret a i n th e i r c o nt e nts w h il e oth ers d o n ot Na y w e .

,
e ve n fi nd t h a t th e
yo unger v es s els a re m ore p e rvi ou s t h a n th e old er
o n es if ro u n d th e y o u ng e r o n es th e re i s a for m a ti on o f w o o d
,
.
Thus th e n i s c o nfi rm e d th e i nfe re nce b efore d ra w n th at i n o rdi

, , ,
na ry s t em s th e s t a i ni ng o f th e w o o d by a n as c e ndi ng c ol ou r e d liq u id
i s d ue not t o th e p as s ag e of th e c o l ou re d liq u id u p th e s u b s ta nc e o f
,
th e w oo d bu t t o th e p e rm e a bili ty of i ts d u ct s a nd s u ch of i t s pi tt e d
,
c e ll s a s a re u ni t e d i nt o i rreg u l ar ca n a l s A nd th e fact s sh ow i ng .
th i s a t th e s a m e ti m e i nd i c a t e w i th t ol e ra bl e c l e arne s s t h e p roce s s
,
by whi ch wood i s form e d W ha t i n th es e c as es i s s e e n t o t ak e pl a ce

.
w i th a dy e m ay be fa i rly p re s u m e d t o ta k e pla c e w i th s a p
,
W h ere .
t h e dye e x u d es bu t s l ow ly w e m a y i nfer th a t th e s a p e x u d es bu t,
s l o w ly a n d i t i s a fa i r i nfe r enc e th a t w h ere th e d e l e a k s rapidly ou t

y
o f th e v es s el s th e s a
p d o e s t he s a m e
,
I nfe rri ng th u s th a t w h ere .
, ,
e v e r th ere i s a con s id e ra bl e form a ti on of w oo d th e re i s a con s id era bl e
es c a p e o f th e s a
p w e s ee i n th e o ne th e res ul t of th e oth er
, The .
th i c k eni ng o f t h e p ros e nch y m a i s p rop ort i ona te t o th e qu a nti ty of

nu t ri t iv e liqu id p as s i ng i nt o i t ; a nd th i s nu t ri ti ve liq u id p a s s e s
i nto i t from th e v ess els d u cts a nd i rregula r ca nals i t s u rro u nd s
, , .
B ut a n obj ec ti o n i s m a d e t o s u ch e x p e ri m ents as the forego i ng ,
a nd to a ll th e i nfere nc es d ra w n from th e m I t i s s a id th a t p o rti o ns .
o f pl a nts c u t o f f a nd th us t re a t e d ha v e th ei r p h y s i ol ogi ca l a cti o ns ,
a rre s t e d or s o ch a ng e d as m a re n d r th e re u lt m i l di ng ; a nd i t
, y e s s s e a
i s s a id th a t w h e n d e t a c he d s h o ots a nd l ea v es h a v e the i r c u t e nd s
pl a ce d i n s olu ti ons th e o p en m o u ths of th e i r v es s els a nd d u cts ar e
,
di rec tly p res ent ed w ith th e li qu id s t o be a b s orb e d wh i ch d o es not ,
h a pp e n i n th ei r na tu ral s t at es Furthe r m a ki ng th es e o bj ecti ons .

,
l ook s eri ou s i t i s alleg ed th at wh en s ol uti ons are a b s orb ed through

,
t h e roots q u i te d i fi ere nt re s u l ts a re o b t a i ne d : th e a b s orb e d m a tte rs

,
a re found i n th e ti s s u es a nd not i n th e v es s el s C l ea rly w ere the e x .

,
CI RC U L A TI ON A ND F ORMA T I ON OF WOOD I N P L A N TS .
5 73
er i m ent s yi e ldi ng th ese a d v ers e resu lts c ond uct e d i n u n o bje ct i o n

p
a bl e w a y s th e co nc lu s i o n i m li e d b t h em w ould ne g a t i v e t h e co n
, p y
e lu s i o ns a b o ve d ra w n B ut t hes e e x p e ri m e nt s are no l es s o bje eti on
.
a bl e t h a n tho s e t o w h i ch t h e y a re o pp os e d S u ch m i ne ra l m a tters .
a s s a lts o f i ro n s o l u t i o ns o f w h i ch h a v e i n s o m e c as e s b e e n s u ppli e d
,
t o th e ro o ts for th e i r a b s orpti on a re o b v i ou s ly s o u nlik e t he m a t

,
t ers ordi na rily a b s orb e d t ha t th ey a re lik ely t o i nt e rfe re fat ally

,
w i th t h e p h y s i ol og i ca l a c t i o ns I f e x p e ri m ents of thi s ki nd a re
.
m a d e by i m m e rs i ng t he roo t s i n a dye th e re i s b es id es th e d if , ,
fi c u l ty th a t th e m i n era l m ord a nt c o nt a i n e d by th e dye i s i nj u ri o u s

t o t h e pl a nt t h e fu rth e r di fli cu l ty th a t th e c o l ou ri ng m a tt e r b e i ng
, ,
s e i ze d by th e s u b s t a nces for w h i ch i t h as a n a f fini ty i s l eft b ehi nd ,
i n th e fi rs t l a y ers of ro ot ti s s u es p a s s e d th ro u gh a nd th a t th e ,
d ecolo ri ze d w a t er p a ss i ng u p i nto th e pl ant i s not tracea ble T 0 .
be c oncl u s iv e th en a n e x p eri m ent o n a b s orp ti o n th roug h r oots

, ,
m u st b e m a d e w i th s om e s olu ti o n w hi ch w ill no t s e ri o u s ly i nt e r
fe re w i th th e pl a nt s v i t a l p ro ces ses a nd wh i ch w ill not h a v e i ts
’
dis t i nc ti v e el em e nt l eft b eh i nd T o fulfil th ese re qu i rem e nts I .
a d o t e d th e fo ll o w i ng m eth o d
p H a v i ng i m b e d d e.d a w ell s o a k e d -
b ro a d b ea n i n m oi s t s a nd c onta i ne d i n a n in v ert e d co ne of c a rd
-
,
b o a rd w i th i ts a p ex c u t o ff for th e ra di cl e t o c om e th rou gh — h a v i ng
pl a ce d thi s i n a w id e m outh ed dw arf bottl e p artly fill ed w i th w at er
-
, ,
s o th a t t h e p r otru di ng ra di cl e dipp e d i nt o th e w a t er— a nd h a v i ng
w a i t e d u ntil th e y ou ng b e a n h a d a s h oot s o m e th ree o r m ore i nch es

hi gh a nd a c lus te r of s e c o nda ry rootl ets from a n i nch to a n i nch
,
a nd a h a l f l o ng— I s u ppli e d for i ts a b s orp ti o n a s i m pl e d ecoc t i o n o f

-
l ogw oo d w hi ch b e i ng a veg etal m a tter w as not lik ely to d o i t m u ch

, , ,
h a r m a nd w h i ch b e i ng w i th o ut a m ord a nt w ould not l e a v e i ts s us

, , ,
p end e d col our i n th e firs t t i s s u es p a ss ed through T o a v oid a ny .
p os s i bl e i nj ury I did not rem o v e th e pl a nt from the bottl e but

, ,
s li h tly ra i s i ng th e c o n e o u t o f i t s ne ck I p o u re d a wa y t h e w a t e r
g ,
thr ou g h th e c re vi c e a nd th en p ou re d i n t h e l o gw ood d ec o ct i on s o
th a t t h e re cou ld h a v e bee n no br ok e n e nd o r a b ra d e d s u rfa c e of a
root l et t h ro u h w h i ch th e d eco ct i o n m i gh t ent e r B e i n g p re p a r e d
g .
w i th s o m e ch l o ri d e o f t i n as a m o rda nt I cu t off a ft e r s om e th r ee , ,
h ou rs o ne of th e l o w es t l e a v e s e x p ecti ng th a t th e a ppli ca t i on of th e
, ,
m o rd a nt t o th e c u t s u rfa c e w o u ld b ri ng ou t th e ch a ra ct eri s t i c c o l o u r
i f th e l og woo d d eco cti on h a d ri s en t o th at h e ight I g ot n o re .
a c t i o n h o w e v er
, B ut a ft er e igh t h ou rs I fou nd on c u t ti ng o ff
.
,
a noth er l ea f th a t th e v es s els of i ts p e ti o l e w ere m a d e v i s i bl e a s d a rk

, ‘
s t rea k s by th e c o l ou r w i th wh i ch th ey w e r e ch a rg e d— a c ol o u r di ffer
i ng as w as to be e x p ect e d from th a t o f th e l og wo od d ec oc ti on
, , ,
w h i ch s p ont a neou sl y ch a ng e s e v en by s i m pl e e xp os u re I t w as th en .
to o l a te i n th e d a y t o p u rs u e th e o bs erv a ti ons ; bu t ne x t m o rni ng

th e v e s s el s o f th e w h ol e pl a nt as fa r a s th e p eti ol es of i ts h igh es t
,
u nfold e d l e a v es w e re full o f th e col ou ri ng m a tt er ; a nd o n a pplyi ng

,
-
c h l o rid e of t i n t o th e c ut s u rfa c es th e v es s el s a ss um e d th a t p urpli s h

,
5 74 A P P E ND I X 0 .
re d wh i ch th i s m d a nt p rodu c es wh en di rectly m ixe d wi th th e l og

or
w o o d d e coct i on S u b s e qu ent ly w hen on e o f t h e cot y l ed o ns w a s c u t

.
,
op en by P rof Oli v er t o w h o m . i n c o m p a ny w i th D r H o o k e r I
, , .
,
s h ow e d th e s p ec i m e n w e fou nd tha t th e w h ol e o f i t s v as cu l a r s y s t em
,
w a s fill e d wi th th e d e c o ct i on w h i ch e v erywh ere g a ve t he ch a ra ct eri s ti c

,
r ea ct i o n .A nd i t b eca m e m anifes t th a t th e liqu id a b s orb e d throu gh

th e rootlets i n th e c e ntra l v ess el s of w hi ch i t w a s s i m ilarly t ra c ea bl e
, ,
h a d p art of i t p as s e d di rectly u p th e v ess el s of th e a x i s w h il e p a rt o f ,
i t ha d p as se d th ro ugh o th er v e ss e ls i nto th e c otyle d on o ut of w h i ch , ,
no d ou bt t h e liqu id o rdi na r ily s o c arri e d re t u rns c h a rg e d w i th a

,
s upply of th e s to re d nu tri m e nt I h a v e s i nce o b ta i ne d a v erifi ca ti o n

.
by v a ryi ng th e m eth od D iggi ng up s om e y oung pl a nts (Ma rigolds

.
h a pp e ne d t o a fford th e b e s t c h o i ce) w i th l a rge m as s es of s oil rou nd

th e m pl a c i ng th em i n wa te r s o as g ra d u a lly t o d e ta c h th e s oil w i th
, ,
o u t i nj u ri ng t h e ro otl ets pl a nt i ng th em afres h i n a fl ow er ot full

p
-
,
o f w a s h e d s a nd a nd th e n a ft er a fe w d a y s w a t eri ng t h em w i th a
, , ,
l ogw o o d d e co cti on I found as b efore th at i n l es s tha n t we nty

, , ,
fou r h ou rs th e c olo uri ng m a tter h a d ru n up i nt o th e v ess el s of th e

-
l ea v es T hough the rea cti on p rodu ced by t he m o rd a nt w a s not s o

.
s tro ng as b e fore i t w a s m a rk e d e no ugh t o be q u i t e u nqu e s ti o na bl e

, .
A s th es e e x p eri m ents w e re s o c o nd u ct e d t ha t t here w as no ac

c ess t o th e v ess els e x c ep t th rou gh t h e na tu ra l c h a nnel s a nd as th e ,
v i ta l a cti ons o f the pl a nts w ere s o littl e i nte rfere d w i th tha t a t the
e nd of t w e nty fo u r h o u rs t h e y s h ow e d no t ra c es of di s tu rb a nce I
-
,
th i nk th e res ults m u s t be h eld conc l us iv e .
T a ki ng i t th e n a s a fa ct t ha t i n pl a nts p o ss es s i ng th em th e v ess els

, ,
a nd d u c ts a re th e c h a n nel s th rough w h i ch s a
p i s d i s tri bu t ed w e c om e ,
no w t o th e fu rt her u es ti on— W h a t d et erm i ne s t h e v a ryi ng p erm e a

q
bi l i ty of th e w a lls of th e v ess el s a nd d ucts a nd th e c ons e u e nt v ary
q ,
i ng form a t i on of w o od T o thi s qu es ti o n I b eli e v e th e t ru e re ly i s

p
-
The e x p os ure o f t he p art s t o i nt erm i tte nt m e cha ni cal s t ra i ns a c tu a l ,
o r p ot e nti a l or b o th, By a ctua l s tra i ns I of c ou rs e m e a n thos e
.
w hi ch t h e pla nt e x p e ri e nc es i n t he c ou rs e of i ts i ndi v id ual li fe Bv .
p ot e nti al s trai ns I m ea n thos e wh i ch the form a tti tud e a nd ci rcu m , ,
s ta nc es co m m on t o i ts ki nd i nv olv e a nd w hi c h i ts i nh eri te d s tru e ,
t u re i s a d a p te d t o m e et I n pl a nt s w ith s te m s p eti ol es a nd l ea v es
.
, , ,
h a v i ng t ol e ra bly c ons ta nt a tti tu d e s th e i nc rea s i ng p oros i ty o f th e ,
t u bes a nd cons e qu e nt d e p os i t of d ens e t i ss u e t a k es pl a c e i n a nti cip a

t i on of th e s tra i ns t o w h i ch th e p a rt s o f the i ndi v id u a l a re li a bl e bu t
,
t a k es pl a ce a t p arts w hi ch h a v e b e en h a bi tu ally s u bj e ct to s u c h
s tra i ns i n a nc est ra l i ndi v id u a l s B ut th ough i n s uch pl a nts th e .
t e nd enc y t o rep ea t th a t di s trib u ti on of d e nse ti s s u e c a u s e d by

m e ch a ni ca l a ct i o ns o n p as t e nera ti o ns g o e s o n i rres e c ti v e o f th e
g p ,
m ech a ni c a l a cti on s to w hi c h the d e v el o pi ng i ndi v i d u a l i s s u bj e ct

,
th es e di re c t a c ti ons w hil e th e y g re a tly a i d th e as su m pt i o n of th e
,
typi ca l s tru ct u re a re th e s ol e ca u s es of th os e d evi a ti ons i n th e rel a

,
CIRC UL AT I ON A N D FOR MA T ION OF WOOD I N P L A NTS .
5 75
t iv e th i ck eni ngs of p a rts w h i ch di sti ngui sh th e i ndi v id ual from oth ers
o f i ts ki nd A n d th e n i n c ert a i n i rr egu l a rly grow i ng pl a nt s s u c h as
.
, ,
C a ctu se s and E u p horbi as w h e re th e s tra i ns fa ll on p arts th a t d o

,
no t c orres p o n d i n s ucc e s s iv e i ndi v id u al s w e di s t i nc tly t ra c e a di rec t

,
re l a ti o n b et w e e n th e d eg ree s of s tra i n a nd t h e ra t es o f th e s e c h a ng es
w h i c h r e s u lt i n d e ns e t i s s u e I w ill not occupy s p a ce i n d e ta i li ng

.
t he e vi d e nc e of th i s r el a ti on w hi c h i s cons pi cu ou s i n th e ord e rs
,
na m e d bu t w i ll p ass t o t h e qu e s ti on— W h a t a re th e p h y s i c al p roc ess es

,
by w hi ch i nte rm i ttent m ech a ni c al s tra i ns p rodu c e t hi s d ep os i t of

re s i s t a nt s u bs t a nc e a t pl a c es w h e re i t i s need e d t o m eet the s tra i ns ?
W e h a v e not to s eek far for a n a ns w er If a t runk a b ough a .
, ,
s h oo t or a p et i ol e i s b e nt by a gu s t of w i n d th e s u b s ta nc e o f i ts
, , ,
c o n v e x s id e i s s u bj ect t o l o ngi tu di na l t e ns i on : th e s u b s t a nc e of i ts
c o nc a v e s id e b e i ng a t th e s a m e ti m e c o m p re ss ed Thi s i s th e p ri .
m a ry m e ch a ni c al e f fect T h e re i s h ow e v er a s econd a ry m e c ha ni
.
, ,
c al e f fe ct w h i ch h ere ch i efly c once rn s u s

,
Th a t be nd by w hi ch t h e .
t i s s u es of th e c o n v e x s id e a re s tret ch e d a ls o p ro du ce s l at era l c om ,
p re ss i o n of th em B u ttoni ng on a t ight gl o v e a nd th en cl o s i ng th e
.
h a nd w i l l m ak e th i s ne c e s s i ty cl ea r : th e l e a the r w h il e i t i s s tra i ne d
, ,
a l o ng th e b a ck s o f t he fi ng ers p res s es w i th c o ns id e ra bl e fo rc e on th e
,
k nuc kl e s .I t i s d e m o ns tra bl e th a t th e t e ns i ons of th e oute r l ay er

o f a m ass m a d e c o n v e x by b e ndi ng m u s t by com p os i t i o n of forc e s, , ,
p ro d u ce at e v e ry p o i nt a resu lta nt a t right a ngles to th e la y er be

ne ath i t ; th a t s i m il a rly th e j o i nt te ns i o ns o f t h e s e t w o l a y e rs m u s t
, ,
t hro w a p ress u re on th e ne x t d e ep er la y e r ; a nd s o 011 H e nc e if .

,
a t s om e li ttl e di s t a nc e b e nea th t h e s u rfa ce of a s t e m tw ig or l ea f , ,
s t a lk th e re e x i s t l o ngi tu di na l tu b es th ese t ubes m u st be s qu eez e d

, ,
e a ch t i m e th e s id e of t h e b ra n ch th e y a re pl a c e d o n beco m e s c o n v e x .
Modi fyi ng th e illu s tr a ti o n j u s t d ra w n fro m t he c lench e d h a nd w il l

m a k e th i s cl e ar W h en on forc ibly g ras pi ng s om et hi ng th e ski n i s
.
, ,
d ra w n t ightly o v er th e b a ck of th e h a nd th e w h iteni ng of the ,
k nu ckl es sh o ws h o w the bl o od is e x p el l ed from th e v ess els bel ow

th e s u rfa ce by th e p re s s ur e of th e tighte ne d s ki n I f th e n th e s a p .
, ,
v ess els m u s t be th u s com p ress ed wh a t will h app en t o th e liquid th ey

,
c onta i n I t w ill m o v e a w a y a l ong th e li nes o f l e as t res i s t ance .
P a rt a nd p ro b ably th e g reat er p art w ill es c a p e l e ngt h w ay s fr om the

, ,
pl a ce o f g re a te s t p res su re : s om e of i t b ei ng e x p ell e d d ow nw a rd s ,
a n d s om e o f i t u p w a rd s B u t a t th e s a m e t im e p a rt of I t will be
.
, ,
lik ely t o ooze throu gh th e wa lls of th e tubes If the se w all s a re s o .
p e rfect as to p erm it the p as s ag e of liquid only by os m os e i t m a y ,
s t i l l be i nfe rre d th a t th e os m os e w ill i nc reas e u nd er p res s u re ; a n d

p ro ba bly under re current p ress u re th e pla c es a t w hi ch th e os m o t i c
, ,
c u rre nt p as s es m os t r e a dily w ill b e co m e m ore a n d m ore p e rm e a bl e ,
u n til th e y e v e n tu a lly form p o res A t a ny rate i t i s m ani fes t t hat

.
w h ere p ores and s li t s e x i s t w h eth e r thu s fo rm e d or form e d i n a ny

,
oth er w a y the es ca p e o f s a p i nt o th e a dj a ce nt t i ss u e a t e a ch b e n d
,
5 76 A P P E ND I X 0 .
w ill b ecom e eas y a nd ra pi d W h at furt h er m ust h app en

. Wh en
th e b ra nch or s h o ot re c oils th e v e sse ls o n th e s id e th at w as c on v e x
, ,
b ei ng reli e v e d from p re s s u re w il l te nd t o res u m e t he i r p re vi ou s

,
di a m et ers ; and w ill be h elp ed t o d o thi s by th e el as ti ci t y of t h e

s u rro u ndi ng t i s s u e as w e ll as by t h os e s pi r a l a nnu l a r a nd a lli e d
, , ,
s t ruc t u res w h i c h t h e y c o nt a i n B u t thi s re s u m p ti on of th e i r p re v i

.
o u s di a m eters m u s t c a u s e a n i m m e di a t e ru sh o f sa b a ck i nt o t h em
p .
W he nc e w ill i t com e Not t o a ny c ons id era ble e x te nt from the s ur

ro undi ng t i s s u e s i nt o w h i c h p a rt o f i t h as b ee n s qu e ez e d s ee i ng th a t
,
th e res i s t a nc e t o th e re tu rn of liqu id t h ro u gh s m a l l pores w ill be

grea ter th a n th e res i s ta nce to i ts retu rn al ong th e v ess el s th em s el v e s .
Ma ni fes tly th e s a p wh i ch w a s th rus t up a nd d o wn th e v es s els from

th e pl a ce of c om p re s s i on w i ll return— th e q ua nti t i es r etu rni ng from
a b o v e a nd from b el ow v a ryi ng a s w e s h a ll h e r e aft er s ee a c c ordi ng
, ,
to c i rcu m s ta nc es B u t t h i s i s no t a ll
. From s om e s id e a g rea te r
.
qu a nti ty m u s t c om e b a ck th a n w as s ent a w a y ; fo r th e a m ount that

h as e s c ap e d o ut of t h e t u b e i nt o the p ro s enchym a h as t o be re pl a c e d .
Th u s d uri ng t h e ti m e w h e n t h e s id e of th e bra nch o r t wig b ecom es

c o n ca v e m o re s a p re t u r ns from a b o v e o r b e l o w th a n w as e x p ell e d
,
u p w a rd s o r d o w nw a rd s d u ri ng t h e p re v i o u s c o m p ress i o n T he .
re fill e d v es s el s w h en t h e n e x t b e nd re nd e rs th e i r s id e c o n v e x a ga i n
, ,
h a v e p a rt of th e i r c ont ents force d th roug h t h e i r p a ri et es a nd a re ,
ag a i n refill e d i n th e s a m e w a y Th e re i s t hu s s e t u p a d ra u g ht of s a p
.
to th e pl a c e w here t h es e i nte rm i tt e nt s t ra i ns a re g oi ng o n a n e xu d a ,
t i on p r opo rt i ona te t o th e fre qu enc y a nd i nte ns i ty o f t he s tra i ns ,
a n d a p ro p o rt i o n a te nu t ri ti on or th i ck e ni ng of t h e w o o d c e ll s ~
fitti ng t hem t o res i st th e s tra i ns A rud e id ea of thi s a cti o n m a y

.
be o b t a i ne d by gras pi ng i n one h and a d a m p s p onge h a v i ng i ts ,
l o wer end i n w a ter w h il e h oldi ng a pi e c e o f blott i ng p a p er i n

,
-
c ont a c t w i th i ts u pp er e nd a nd th en , gi v i ng th e s p onge rep e a te d

s q u e eze s . A t e a c h s qu e ez e s o m e o f t he w a t er w ill be s ent i nto
t h e bl otti ng p a pe r ; a t eac h rel a x a ti o n th e s p ong e w ill refill from
-
b el ow to gi v e a not her p orti on of i ts contents to the blotti ng

,
p a p er w hen a g ai n s q ueez ed .
B ut ho w d oes thi s e x pla nati on a pply to roots ? If the form a t i o n

o f w o o d i s d ue t o i nt erm i tte nt t ra ns v ers e s t ra i ns s u c h as a re , p ro
d u c e d i n th e a eri al p a rts of up righ t pl a nts by t he w i nd h o w d o e s i t ,
ha pp e n th a t w o o dy m a tt er i s d ep os i te d i n roots w h ere th e re a re no
,
l at era l oscill a ti o ns no t ra ns v ers e s t ra i ns

, T he a ns wer i s th a t ,
l o ng i tu di nal s t ra i ns also a re c a p a bl e of ca u s i ng th e e ffe cts d es cribed .
I t i s t ru e th a t p e rfe ct l y s t ra igh t fi bres u ni t e d i nt o a bu ndl e a nd p u ll e d

l e ngt h way s w o uld not e x ert on o ne a noth e r a ny la teral p res s u re a nd ,
w ould not l ate rally com p res s a ny s i m i l a rly s t ra ig ht c a na ls ru nni ng

-
a l o ng w i th t hem . B u t i f th e fibres u ni te d i nto a b undl e are v ari ou s ly

b ent or t wi s te d th e y c annot be l ongi tu di nally st ra i ned w i th o ut c om
,
p res s i ng o ne a noth e r a nd s t ru ct u res i m be dde d i n them It nee d s .

CIR CU L A T I ON A ND FOR MA T ION OF WOOD I N P L A NT S .
5 77
bu t to w a tc h a w et rop e d ra w n tight by a c a p s tan, to s ee th at a n

a c ti o n li ke t h a t w h i c h s q u e ez e s t h e w a te r ou t of i ts s t ra nd s , w ill
s q u eez e th e s a o u t o f th e v es s el s of a r o o t i nt o th e s u rrou ndi ng

p
t i s s u e , as ofte n as t h e ro ot i s p u ll e d b y th e s w ayi ng of t h e pl a nt i t
b el o ng s t o . H e re , to o , as b e fore , th e v es s el s w ill refill w h e n t h e
p ull i nt erm i ts ; a nd s o, i n th e roots as i n the branches , thi s ru d e
p u m pi ng p roc ess w ill p rod uc e a growth of h ard t i s s u e p rop ort i on
a t e t o t h e s t res s t o be borne .
T hes e c onclu s i o ns a re s u p port e d by th e e vid enc e wh i ch e x c ep

t i o na l c as es s u pply . If i nt erm i tt e nt m ech a ni cal s tra i ns th u s c a us e
t h e form a ti on of wo o d w h ere w o o d i s fou nd , t h e n w h e re i t i s not
fou nd , t he re s h ou ld be a n a b s e nc e of i nt e rm itte nt m e c ha ni c al st ra i ns .
T he re i s s u ch a n a bs e nce Vas c ul a r pl a nts c h a ra ct eri ze d by l i ttl e

.
o r no d ep os i t o f d e ns e s ub s ta nc e , a re th o s e h a v i ng v e s s el s s o c o n
d i t i o ne d th a t no c on s id era bl e p ress u re s a re b orne by t h em . Th e

m o re s u c c u l e nt a p et i ol e o r l ea f b e c om e s , th e m ore d o th e e f fe ct s o f
t ra ns vers e s t ra i ns fal l on i ts ou ter l a y ers of c ells I ts m e ch a ni c a l s u p
.
p ort i s chi efly d eri ve d from th e a bili ty of th es e m i nu t e ves i cles , full

o f liq u id , to res i st b urs ti ng a nd te a ri ng u n d e r th e c om p res s i o ns a nd
t ensi ons th e y a re e x p os e d t o . A nd j u s t as fas t a s this ch a nge from

a thi n l eaf o r foot s t a lk to a t hi c k o ne e nt a il s i n c reas i ng s t res s on th e
-
s up erfi c i al t i s s ue , s o fas t d oes i t di m i ni s h th e s tres s on t h e i nt e rn a lly
s ea t e d v as cul a r t i ss u e .T h e s u c cul ent l e a f c a nnot b e s w ay e d a bou t

by th e w i nd as m u ch a s a n o rd i na ry l eaf ; a nd su ch s m all bend s as
c a n be gi ve n to i t a n d i ts foot s t a lk a re p re v e nt e d from a f fec ti ng
-
i n a ny c o ns id e ra bl e d eg re e th e tubes ru nni ng th rou gh i t s i nt eri or .
H e nc e the re t enti v e n ess of t he v ess el s i n th e s e fl e shy l ea v es , a s s h ow n

by th e s m a l l e x u d at i on o f dye ; a nd h ence th e s m all th i ck e ni ng of
t h e i r s u rr ou ndi ng p ros e nchy m a by w o ody d ep os i t . S till m ore c on
s i c u ous l y i s th i s c o nn e x i o n of fa ct s s h o wn w h en, from t h e s oft th i c k
p
l e a ves b efo re na m ed a nd s u ch o th ers as th os e of E chever ia , R oc hea ,
P er es / vi a , w e t u rn t o th e th i ck l ea v es th a t h a v e s tro ng e x o s k e l e t o ns
-
.
Ga s ter i a s er v es a s a n ill u s t rat i o n .The l eath e ry or h orny s ki n h ere

e v id e ntly b e a r s th e e nt i re w e ight o f th e l e a f, a nd i s s o s ti f f as to pre
v e nt a ny o s ci lla ti on H ere , t he n, th e v e ss els ru nni ng i ns id e a re pro
.
t e c t ed fr om a ll m e ch a ni c a l s t re s s ; a nd a c c o rd i ngly w e fi nd th at th e
c e ll s s u rrou n d i ng th e m a re n ot a pp re c i a bly thi c k e ne d .
E qu a lly c l e a r, a nd m ore s t rik i ng b eca u s e m ore ob vi ou sly e x cep

ti onal , i s th e e v i d e nc e g i ve n by s u c cu l e nt s t e m s w h i ch a re l ea fle s s .
S tap el i a E uf oni a , h a v i ng s o ft p roc u m be nt a x es not li a bl e t o be

b e nt b a ck wa rd s a nd forw ar ds i n a ny c ons id e ra ble d egree by th e
w i n d h a s , ra m ifyi ng th rou gh i ts t i s su e , v ess e ls th at a ll o w bu t a n e x
,
t re m el y s l ow es ca p e o f dye a nd h a v e u nth i ck e n e d s h ea th s .S u ch of
th e E u ph orbi a s as h a v e a c q u i r e d th e fl es h y c h a ra c te r w h il e reta i ni ng
t h e a rbores c ent gro wt h , lik e E up hor bi a Ca na ri ensi s, t ea ch u s th e s a m e
tru th i n a no th e r w a y .I n t h em t he form at i on o f w ood a round th e
83
5 78 A P P E N D IX 0 .
v ess els is i nc onspi cu ous wh e re th e i nterm itt ent stra i ns a re but s li ght ;
bu t i t i s c ons pi cu ou s at th os e j oi nts o n w h i ch l at era l os cillati ons o f
t h e a tt a c h e d bra nch es th ro w g re a t e x t ens i ons a nd com p ress i ons of
t i ss u e Th rou gh out th e Ca c tacece w e fi nd va ri e d e x a m pl es o f t he
.
a l l e ge d r el a t i o n M a m m i l l a r i a fu rni s h es a v ery m a rk e d o ne T h e
. .
s u bs t a n c e of o n e o f t h es e g l o bu l ar m as s e s res t i ng o n th e g rou nd , ,
a d m i ts o f n o be ndi ng from s id e t o s id e ; a nd a cc o rdi ngly i ts l a rg e
b undles of s pi ra l a nd a nnula r v es s els or p a rti ally unite d cells h a v e

,
-
,
v ery fe ebly m ark e d s hea ths not a t a l l th i c k ene d I n s uch typ es as

-
.
Cereu s a nd Op u n ti a w e s e e as i n th e E u ph orbi as t h at wh ere li ttl e

, ,
s tres s fa lls o n th e v e ss el s li ttl e d ep os i t t a k es pl a c e a ro u nd t h e m ;

,
w h il e th e re i s m u ch d ep o s i t whe re t he re i s m u ch s t ress H e re l e t m e .
a dd a c o nfir m a ti o n o bta i ne d s i nce w ri ti ng t h e a b o v e A ft er obs e rv .
i ng a m ong t he C a ctus e s th e v ery m a ni fes t rel a t i on b et we e n s tra i n

a nd t h e fo rm a t i on of w oo d I i nq u i re d o f Mr C ro uch er th e i nt e ll i
,
.
,
g e nt forem a n o f th e C a ctus h ou s e a t Ke w wh ether he fou nd t hi s

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re l a ti on a c o ns ta nt one H e repli e d th at h e dl d a nd th a t h e h a d
.
,
fre q u e ntly te s t e d i t by a rt ifi ci a lly s u bj e cti ng p a rts of t h e m t o s t ra i ns .
Ne gle cti ng at t h e ti m e t o i nqui re h o w h e h a d d o ne thi s i t after wa rd s ,
o cc urre d t o m e t ha t i f h e h a d s o d o ne i t a s t o c a u s e c o ns ta nt s tra i ns
,
th e o b s er ve d re sul t wou ld not t ell i n fa vou r of th e fo reg oi ng i nt e r
p reta ti on S ub s e qu ently h owe v er I l ea rne d th at h e ha d p rod uc e d

.
, ,
t he s t ra i ns by pl a c i ng th e pl a nt s i n i ncli ne d a tti tu d es — a m eth o d

w hi c h by p erm i tti ng o s c ill a t i o ns of t he s tra i ne d j oi nts al l ow ed t h e
, ,
s t ra i ns t o i nterm i t A nd the n m a ki ng t he p roof concl u s i v e Mr

.
, , .
C rou c her v ol u nteere d th e s ta te m ent th a t w h ere h e h a d p roduc e d

c ons ta nt s tra i ns by t yi ng n o fo rm a ti o n o f w o o d t o o k pl ace
, .
A b e rra nt g row th s o f a nothe r c l as s di s pl a y th e s a m e rel a ti o ns

o f p he no m e na T a k e fi rs t th e u nd e rg rou nd s t em s s u c h as th e
.
,
P o ta t o a nd the A rt i ch ok e The v e s s el s wh i ch ru n th rou gh th es e

.
,
s l o wly t a k e u
p t he dye w i th out l et ti ng i t p ass t o a ny c o ns id e ra bl e
e x te nt i nto th e s u rrou ndi ng t i ss u es * Only after an i nte rv al o f
.
’
m a ny h ou rs d o e s t he p ros enchy m a b ecom e s ta i ne d i n s om e pl a ces .
H ere as b e fore a n a bs e nc e of rapid e x ud ati on a c com p a ni es a n

, ,
a bs e nce o f w oo dy d ep o s i t
; a nd b o t h th es e g o a l ong w i th t h e a h
s ence of i nt e rm i tte nt s tra i ns T a k e a ga i n t h e fl esh y roo ts
. Th e .
T urni p th e C a rrot a nd th e B e etroot h a v e v ess el s th a t re t a i n v e ry

, , ,
p ers i s tently th e col ou re d liqu id s t hey t ak e u p A nd di ffe ri ng i n thi s .

,
a s t hese ro ots d o fro m o rdi na ry ro ots w e s ee t h a t th ey a l s o d if fe r

, ,
from the m i n not b e i ng w oo dy and i n not be i ng a p pre c i a bly s u b

,
Those wh o repeat t he s e e x peri m ents m u s t b e p r ep a red for gre a t i rregu ~
l a ri ti e s i n the ra t es of a b s orpt i on Su cc ul ent s t ru c tures i n g enera l a b sorb

.
m uc h m ore s l o wl
y tha n others , a nd s om et i m e s wi l l s ca rc el y t a k e u p t h e dye
at al l. The d i ffe rences bet wee n d i ffe rent s t ru cture s , a nd th e s a m e s tru c tu re
a t di f fe rent ti m es , p rob a bl y d e p e nd o n t h e de re e s i n wh i c h th e ti s s ues a re
g
cha rge d wi th li q ui d and t he ra t es a t wh i ch t h e a re l os i n i t b e v a o ra ti on
y g y p .
CIRC U L A T ION A ND FORMA TI ON OF WOOD IN P L A NT S 5 79 .
j c et to th e l m ech a n c a
i l actions I n th es e cas es as i n the
us u a .
,
o th ers p a rts th a t o rdi na rily b e c om e d e ns e d evi a t e fro m th i s t

, yp ,
i cal cha ra cter wh en th ey a re not exp os e d t o th e s e force s wh i c h

p ro du ce d ens e t i s su e by i nc reas i ng t he e x tra vas a ti o n of s ap .
T o com pl et e t he p roof t h a t s u ch a rela t i on e xi s ts l et m e a dd t h e ,
re s u l ts of s om e e x p eri m e nts on e q u a l a nd s i m ila rly d e v el op e d p a rt s -

,
k ep t res p ec tively a t res t a nd i n m oti on I ha v e t es t ed th e effe cts o n .
l a rge p et i ol es o n herb ac eou s s hoots a nd on w oo dy sh oots I f t w o

, , .
s u ch p et i o l es as t h o s e of Rhu b a rb w i th th e i r l e a v es a tt a ch e d h a v e
, ,
th e i r c ut e nd s i ns e rt e d i n b ottl es of dye a nd th e o ne be b ent b a c k ,
w ard s a nd forw a rds w hi l e th e oth e r re m a i ns m oti onl ess t here ari s es , ,
a fte r th e l a p s e o f a n h o u r s c a rc ely a n di f
f re nce i n th e s t a te s o f
y e ,
t h e i r v es s els : a c e rta i n p rop ort i on of th e s e a re i n b oth c as e s c h a rg e d

w i th th e dye a n d li ttl e e x u d a t i o n h as been p rod u ce d by t h e m oti o n
, .
H ere h o w e v e r i t i s to be o b s erv e d th a t th e c a u s e s of e x u d at i on a re
, ,
s c arc e ly op era t i v e ; t he v as c u l a r b u ndl e s a re di s t ri bu t e d a ll t hrou gh
th e m as s of th e p eti ol e wh i c h i s form e d of s oft w a t e ry ti s s u e ; a nd

,
t h ey a re t h erefo re not s o c i rcu m s ta nce d a s t o be e ffe ctu a lly c om

, ,
p res s ed by t he be nd s I n h erba c eou s s tem s s u ch as th os e of t he

.
,
Jerus a le m A rt i c hok e a nd of th e Fox glov e a n effect s c a rc ely m ore ,
d ec id e d i s p ro d u c ed ; a nd here t oo w hen we s eek a rea s on w e find , , ,
i t i n t h e non ful fil m e nt o fth e m ech a ni c al co ndi t i ons ; for th e v as cul a r

-
b undles a re not s o s ea t e d betw e en a to ugh l ay e r of b a rk a nd a s olid

c o re as t o b e c o m p ress e d a t e a c h b e nd W h en how eve r w e c om e .
, ,
t o e xp e ri m e nt u p on w oody s h oots w e m e et w i th c ons pi c u ou s e ffe c ts

, ,
t h oug h by no m e a ns u ni fo rm ly I n s om e cas es o s c ill a ti ons p ro du c e

.
i m m ens e a m ount s of e x u d a ti on— p a rallel t ra ns v ers e s ecti ons o f th e

c o m p are d sh oo ts s h o w i ng tha t w h e re i n th e one t ha t h as b e en a t ,
re s t th e re a r e s p ots o f co l ou r ro u n d but a fe w pi tt e d d u c ts i n th e
, ,
o ne th a t h as b e e n k e pt i n m ot i o n th e s u bs t a n ce o f th e w oo d i s s oa k e d
a l m os t u ni fo rm ly thro ug h w i th dye I n oth er ca s es , es p ec i a lly where

.
th ere i s m u ch u ndi ffe re nt i at e d t i s s u e rem a i ni ng th e e x u d a ti o n i s no t ,
v e ry m a rk e d T he differe nc e a pp ea rs t o d ep end on th e qu a nt ity of

.
liq uid c o nta ine d in th e s h oot If i ts s u b s ta nce i s rel a ti v ely d ry th e

.
,
e x u d a ti o n i s g rea t ; bu t i t i s c om p a ra t i vely s m a ll i f a l l t h e ti s s u e s
a re fu lly c h a rg e d w i th s a p T hi s c o ntras t of res ul ts i s one w hi ch

.
c ont em pl a t i o n o f th e m ech a ni ca l a ct i o ns w ill l ea d u s t o e x p e c t .
A nd n o w w i th th e s e fa cts t o a i d ou r i nt e rp ret ati on l et u s re

, ,
tu rn t o o rdi na ry s t em s I f t h e u pp e r e nd o f a gro w i ng s h oot t h e

.
,
p ro s ench ym a of w h i c h i s but li ttl e th i ck ene d be a ll ow e d to i m bi be ,
t h e dye t h e v esse l s o f i t s m e d ulla ry s h e a th a l o ne b ecom e c h a rg ed ;

,
a nd fro m th e m th ere t a k es p l a ce bu t a s l ow o oz i ng If a lik e e x .
eri m e nt b e t ri e d w i th a l o wer p a rt of t h e s h oo t w h e r e th e w oo d i n
p ,
c ou rs e o f form a t i o n h as i t s i nn er b o u nd ary m a rk e d bu t not i t s o u t e r

b o und a ry w e find tha t th e p i tte d d ucts a nd m ore esp eci ally th e
, ,
i nn er o ne s c om e i nto pl a y A nd th en l ow e r s till w here th e w o od

,
.
,
58 0 A P P E ND I X 0 .
h as i ts p eri ph ery d efine d a nd i ts hi s tologi cal ch ara cters d e cid e d ,
th e a pp eara nces s h ow t ha t t he t i ss u e fo rm i ng i ts ou te r s urfa ce

b egi ns t o t ak e a l e a di ng p art i n th e tra ns m i s s i o n of liqu id W h at .
no w i s th e e xpl a na ti o n of th e s e ch a ng es m e c h a n i c a lly c o ns id e re d ,
I n th e y ou ng s oft p a rt of t h e s h e e t a s i n al l no rm a l a nd a b norm a l
,
g rowth s th at h a ve not form e d w ood th e ch a nnels for th e p as s ag e of

,
sa are th e s pi ra l a nnu l a r fe nes t rat ed o r r e ti c u l a t e d v es s el s Th es e

p , , ,
.
v e s s el s h ere i nclu d e d i n th e b u ndl e s of t he m e du ll a ry sh e a th a re

, , ,
i n com m o n w i th t h e t i ss u es a rou nd t hem s u bj ect by th e b en di ngs , ,
o f the s h o o t t o s ligh t i nt e rm i tt ent c o m p ress i o n s a nd e s p e ci a lly

, , ,
t he ou term os t of th em are thu s force d t o gi v e th e p ros ench y m a

,
a n e x t ra s u pply of nu tri t iv e liqu id Th e t hi ck e ni ng o f th e p ro

.
s ench m a s p rea di ng l a t e rally as w ell a s ou twa rd s fro m e ach bu ndl e

y ,
o f th e m e d u ll a ry s h e a t h g oe s o n u ntil i t m ee ts th e th i c k e ni ngs t h a t
,
S p re a d from th e oth e r bu ndl es ; a nd t h e re i s s o fo rm e d a n i rre g ul a r

c y l i n d e r o f h a r d ene d ti s s ue s u rrou ndi ng t h e m e d u ll a a nd t h e v as
’
c ul ar b u ndl e s o f i ts S h e a th A s s oon as th i s ha pp ens th e se v a s cu l ar

.
,
b u ndles b ecom e to a c o ns id erable e x te nt S hi eld ed from th e effe cts

, ,
of t ra ns v ers e s t ra i ns s i nc e th e t ens i o ns a n d com p res s i ons chi efly

,
fa ll on th e d ev elopi ng w oo d ou ts id e o f th em C l ea rly too t h e .

, ,
greates t s tres s m u st be felt by th e outer l ay er of the d ev el opi ng

w oo d : b ei ng furth er re m ov e d from t h e neut ra l a x i s i t m u s t be ,
s u bj ect t o s e v ere r s tra i ns a t e a ch be nd ; a n d lyi ng b etw ee n th e
b ark a nd th e lay er of wood firs t fo rm e d i t m ust be m os t exp os e d ,
t o l at eral com p res s i o ns A m o ng th e el o ng a t e d c ell s of thi s oute r

.
l ay e r s om e uni te t o form th e pi tte d d u ct s B ei ng a s w e s ee

, .
, ,
b etter c i rcu m stance d m ech a ni ca lly the y b e com e gre at er ca rri ers
,
o f s a p th a n th e ori i na l v es s el s a nd i n c o ns e q u e nce o f t hi s a s w ell

g , , ,
a s i n cons e qu e nce of t he i r rel a ti v e p r o x i m i ty b ec o m e th e s ou rc e s ,
of nu tri ti o n t o th e s till m ore e x te r na l l a y e rs of w ood c e ll s Th e -

.
s a m e c a u s es a nd th e s a m e ef fec ts h old wi t h e a ch ne w i nd u ra te d
c oa t d ep o s i t e d rou n d th e p re v i ou s ly i nd ura t ed co a ts .
Thi s d es c rip ti on m ay be thou ght t o g o far to w ard s j us ti fyi ng the

curre nt v i e w s res p e c ti ng th e c ou rs e t a k e n by th e s a p B ut th e
“
j ustifi cati on i s more a pp arent th an real I n the first place the i m .

,
pli c a ti on h ere i s th at th e s a p c arryi ng functi on i s a t firs t di s ch arge d

-
e nti rely by th e v esse ls o f th e m e d u ll a ry S h e a th a nd th a t t h ey c ea s e ,
t o di s ch a rg e th i s funct i o n o nly a s fa s t as th ey a re rel at i vel y i nc ap a c i

t at e d by t he i r m ec h a ni c a l c i rcu m s t a nce s A nd th e s e cond i m pli ca.
t i on i s th a t i t i s not th e w oo d i t s elf bu t th e m ore o r l e s s cont i nu ous

, ,
c a nal s form e d i n i t w h i ch a re th e s u b s e qu e nt s a
p di s t rib u t o rs Thi s
-
, .
,
t h ou gh re a dily m a d e cl ear by m i c ros c opi c e x am i na ti on of t he l a rg e
p i tte d d u c ts i n a p arti a lly l i g nifi e d S h oot th a t h a s a bs orb e d th e dy e ,
1 8 l e s s m a ni fes tly t ru e of th e p erip h era l l a y er o f s a c arryi ng t i s s u e

p
-
fi nally form ed B ut i t i s re a lly true here Fo r th i s la y er t hough

. .
,
nom i na lly a l a y er of w o od i s p ra cti c a lly a l ay er o f i nos cul a ti ng

,
CI RC U L A TION A N D FORMA T ION OF WOOD I N P L A NT S .
581
v es s els It i s form e d out of i rregula r lines a nd networks of elon

.
g a te d pi tte d c ells o bliqu ely u ni te d by th ei r e nd s E x am i nati on of

, .
t h em aft er a b s o rp ti on of a dy e sh ow s t ha t i t i s o nly al ong t he c on

,
t i nu o u s c ha nnels t h ey u ni t e t o fo rm t ha t th e c urrent h as p ass ed .
B ut th e ess e nti ally va s cul a r ch a ra ct e r of t hi s o ut er a nd l a t es t form e d -
l ay e r of th e a lburnu m i s best s ee n i n th e fa c t t ha t the v as cu la r sys

t e m s o f ne w a x es t a k e t h e i r ri s e from i t a nd form w i th i t c o nt i nu o u s
,
c a na l s . If a sh o ot of l as t y ea r i n w hi ch g rowth i s rec o m m e nci ng b e ,
c u t l e ng th wa y s a ft e r i t h as i m bib e d a d e c l ea r p r oof i s o b t a i n e d
y ,
t ha t t h e p as s a g e of the dye i nt o a la t era l bu d t a k e s pl a c e fro m thi s

o u t erm os t l ay er o f pi t te d c e l l s a nd th at th e c ha nn e l s t a k e n by th e
,
dy e t h rou gh t h e ne w t i s s u e a re c o m p os e d of c ell s th a t p as s through

m o difi e d form s i nt o t h e S pi ra l v e s s el s of t h e n ew m e d ulla ry s h e a th .
T hi s t ra ns i t i on m ay be s t ill m ore cl ea rly t ra c ed i n a t e rm i na l bu d

t ha t c o nti nu es th e li ne of la s t y ear s s h oot A lo ng i tu di na l s e ct i o n
’
.
o f th i s s h o w s th a t th e ve s s el s o f th e new m e d ull a ry s h ea th d o not
o b t a i n t he i r s a
p from t h e v e s s el s of l a s t y e ar s s h e a th
( w h i c h
’
as ,
s h o w n by th e n o n a b s orp t i on of dy e h a v e b e com e i na c t i v e
-
, ) b ut ,
t h a t t h e i r s uppl i es a re o b ta i ne d from th o s e i nos cul a ti ng ca na ls

form e d out of las t y ea r s ou t erm o s t l ay e r of p ros enchy m a a nd th a t
’
,
b e twe e n th e com p o ne nt c ells of th i s a nd t ho s e of th e new v as cul ar

*
s y s t em th e re a re a ll g ra d a ti ons of s t ru c tu re .
It m ay be a dde d h e re tha t, on cons i de ri ng the m e ch a ni c a l a cti ons t ha t

m u s t go on, we a re e na bl e d i n s o m e m ea s ur e t o u nde rs t a nd b ot h h ow s uch i nos
cu l a t i ng c h a nn e l s a r e i ni t i a t e d , a nd h o w t h e st r uct u res o f the i r c om p onent
ce l l s a re ex pl i ca bl e . Wh a t m u st ha ppen to one of the s e el onga ted p ros en
c hym a-ce l l s i f, i n t he cou rs e of i t s d e v e l op m ent , i t i s s ub e c t to i nterm i ttent j
c o m pre s s i ons ? I t s s q u e e ze d o ut li q u i d wh il e p a rt ia l ly e s ca pi ng l a t era l l y,
-
w i l l m ore l a rgel y e s ca p e u p w a rds a nd do wnw a rds ; a nd w hi l e re p ea te d

l a t eral e sca p e w i l l t end to form l a t era l ch a nnel s com m uni ca ti ng wi th
j
l a t e ra l l y-a d ace nt c el l s , r ep ea te d l ongi tu di na l e s c a pe wi l l t end to form
c h a nn e l s co m m u n i ca t i ng w i t h l on g i t u di na l l y-a d a c e ntjc e l l s — so p rod uc i ng
co n ti nu ou s t ho ugh i rregul a r l ongi tud i na l c a na l s . Mea nwhil e e a ch c el l i nto
a nd ou t of whi ch t he nu t ri t i v e l i q u i d i s from ti m e t o ti m e s q u eeze d
t h rough s m a l l openi ngs in i t s w a l l s , ca nnot thi ck e n i ntern a l l y i n a n
e v en m a nne r : dep os i t i on wi l l b e i nte rfered wi th by t h e p a s s a g e of t he
cu rre nt s t h rough t h e p ore s .The ru s h t o or from ea ch p ore wil l t end
t o m a i nta i n a funnel s ha p e d d ep re ss i on i n th e dep os i t a round ; a nd t he
-
j
op eni ng from cel l to ce l l w i l l s o a c q ui r e u s t t h at sh a pe wh i ch th e m i cros c op e
s h o ws u p — t wo h ol l ow cone s wi th t h ei r a p i ce s m eet i ng a t t h e p oi nt whe re
the c ell m em b ra ne s a re i n cont ac t

-
Moreov er; a s confi rm i ng thi s i nter
.
p rot a t i on, i t m ay he re m a rk e d th a t we a re th us s u pp l i e d wi th a rea s on

for t he di fference s of s h a pe b et wee n t h e s e p a s s a ge s from one p i tted c el l
t o a no th er, a nd th e a na l ogou s p a s s a ge s th a t ex i st b e twee n c e l l s o the r
wi se form e d a nd oth e r wi s e c onditi one d I n t he cel l s of th e m edul l a , a nd
.
ot h e rs w h i ch a re b u t l it t l e e x p o s ed t o com p re s s i on, the p a ss a g e s a re se v e

ra l l y form e d m ore l ik e a t u b e w i th t wo t r um p e t m outh s , on e i n e a ch ce l l
-
.
j
Thi s i s us t th e form wh i ch m i ght b e exp e cte d wh ere th e nut ri t i v e ui d fl
asse s from ce l l t o ce l l i n m ode ra t e c u rrent s , a nd n ot by th e v i ol ent r u s h e s
p
ca u sed by i nterm i tte nt p res s ure s Of c ou rs e i t i s not m ea nt tha t i n e a ch
.
5 32 A P P E ND I X o .
I t i s not th e a i m of th e foreg oi ng re a s oni ng t o Sh ow th at m ech a ni

c a l acti o ns a re t h e s o l e c a u s es o f t h e fo rm a t i o n of d e ns e t i s s u e i n
p l a nt s .D ens e t i ss u e i s i n m a ny c as es for m e d w h e re n o s u c h c a u s es
h a v e c o m e i nt o pl a y— a s , for e x a m pl e , i n th orns a nd i n th e s h ells o f

nu t s . H e r e th e n a tu ra l s el ect i o n o f v a ri a ti o ns c a n a l one h a v e Op e
r ate d . I t i s m a ni fes t, t o o, th a t e v en th os e s u pp ort i ng s t ru ctu re s th e
bu ildi ng u p o f w hi ch i s a b o v e as c r ib e d t o i nt erm i tt e nt s tra i ns , m a y,
i n th e i ndi v idu a l pla nt o f a s p e c i e s th at ordi narily h as th em , be d e
v el o e d t o a g re a t e xt e nt w h e n i nt e rm i tt e nt s t r a i ns a re p r e v e nt e d
p .
W e s e e thi s i n t rees th at a re a rt ifici ally s u pp ort ed by na ili ng t o

w alls ; a n d w e a ls o s e e a ki nd r ed fa c t i n na tu ra l c li m b ers Thou g h
.
i n th e s e c as e s t h e form a t i on o f w oo d i s o bvi ou s ly l es s th a n i t w ou ld
h e w ere th e s t em a nd b ra nch es h a bi tu ally m o v e d a bou t by th e wi nd , i t
ne v e rt h e l e s s g oes o n . C l ea rly th e t end e nc y of t he pl a nt t o rep e a t th e
s t ru c tu re o f i t s t yp e i n th e o n e cas e t h e s t ru c tu r e of i ts S p ec i es ; a nd
(
i n th e o th er c a s e th a t of th e ord e r from wh i ch i t h as di v e rg e d i n be
c o m i ng a c li m b e r i s h e re a lm o s t t h e s o le c a u s e o f w oo d fo rm a ti on
) -
.
B ut th ou gh i n pla nt s s o c i rcum s t ance d i nt e rm i tt e nt m e ch a ni c a l

s tra i ns h a v e li t tl e o r no di re ct s h a r e , i t m a y s t ill be t ru e , a nd I b e li e v e
i s t ru e , th a t i nt erm i tt ent m ech a ni c al s t ra i ns a re th e ori gi na l c au s e ;

for, a s b efore hi nt e d , th e t ypi c a l s tru c ture wh i ch t h e i ndi vidu a l t hu s
rep ea ts i rres p ec t i v e o fi ts o w n c o ndi ti o ns , i s i n terp r e t a bl e as a typi cal
s tru ctu re th a t i s i ts el f th e p ro d u c t o f t h es e a c t i o ns a nd re a c ti o ns be
t ween th e pl a nt a nd i ts e n v i ro nm e nt . Gra nt th e i nh erit a n ce offunc

t i o na ll y p ro d u c e d m o dific a ti o ns ; g ra nt t ha t na tu ral s el ecti on w ill
-
a lw a ys c c - o p e ra t e i n s u ch w a
y as t o fa v our th o s e i ndivid u a ls a nd
fa m ili es i n wh i c h fun ct i on ally p ro du c e d m o dific a ti o ns h a v e pro
-
re s s e d m o s t a d va nt a g eou s ly ; a nd i t w ill foll o w th at th i s m ec h a ni

g
c a lly - c a us e d form a ti o n of d e ns e s u b s t a nc e , a c c u m ul a t i ng from en
g
e ra t i o n t o
g enera t i o n by th e s u rvi val of th e fi ttes t , w ill re s u lt i n a n
o rg ani c h a bi t o f fo rm i ng d e ns e t i ss u e a t th e re qu i re d pl a c es The
.
d ep os it ari s i ng from e x ud a ti on a t th e pl a ce s of g re at es t s trai n, re

c u rri ng fro m ge nera ti on to ge ne rat i o n a t th e s a m e plac es w ill ,
c o m e to be rep r od u ce d i n a nti c ip a t i o n of s t ra i n, a nd w ill c o nti nu e
t o be re p ro d u c e d for a lo ng ti m e a ft er a c h a ng e d h a bi t o f t h e
S p e c i es p re v e nts th e s tra i n— e v e ntu ally , h ow e v er, d ec re as i ng b o th
,
t h ro ug h fu nc t i ona l i na ct i v i ty a nd na t ura l s el ecti on, t o th e p o i nt

a t w hi ch i t i s i n e qu il i b ri um wi th th e re u i rem ent
q .
i ndi vidua l c el l t hes e s t ru ct ure s a re det erm i ne d by th e s e m ech ani c al a cti ons .
T he fac ts cl e a rl y nega ti v e a ny s uch concl u s i on, s howi ng u s , as th ey i n m a ny

c a se s d o, t h a t t h e s e s t r u c tu re s a re a s s um ed i n a d v a nce of th ese m e c h a ni ca l
a c ti ons Th e im pli c a ti on i s tha t s u ch m e ch ani ca l act i ons i ni ti a te d m odi fi
.
,
ca t i ons t h a t h a v e, wi t h t h e a i d of n a t ur al s el ec ti on been a ccu m u l a t ed fro m

,
ge ne ra t i on to gen e ra t ion ; unti l, i n c on for m i ty wi th ord i na ry e m b ryol og i ca l

l a ws , the c el l s o f t h e p a r ts e x po s e d to s uch a c ti ons a s s u m e t h e s e S p ec i a l
s tru c t ures i rres e c ti v e of t h e a c ti ons — th e a c t i ons h ow ev e r s ti ll serv i n
p , , g to
a i d a nd c om l e t e t he a s s um
p p ti on of th e i nh eri te d typ e
.
CI RC U L A T I ON A ND F OR MA TI ON OF WOOD I N P L A NT S .
5 83
A noth er s i de of th e ge ne ra l qu e s ti on m a y no w be cons id ere d W e .
h a v e s ee n h ow , by i nte rm itt e nt p re ss u re s o n c a pilla ry v ess els a nd

du cts a nd i nos cu la ti ng c a nals , t he re m u st be p rod uc e d a draught of
s a t o w a rd s t h e p o i nt o f c o m p ress i o n t o repl a c e th e s a
p p s qu eez ed ou t .
B ut w e h a v e s t il l t o i nqu i re w h a t w ill b e th e effect o n t h e di s tri bu

ti o n o f s a p thro u gh ou t the pl a nt as a wh ol e . It was c o nclu d e d tha t
o u t o f th e c o m p re s s e d v ess el s t he g re a t e r p a rt o f t h e liqu id w ou ld
e s ca p e l o ngi tu di na lly — t he l o ngi t u d i na l res i s t a nc e t o m o ve m e nt
b e i ng le as t I n e v e ry c as e the p ro ba bili ti es a re i nfinity t o one ag a i ns t

.
t h e r e s i s t a n ces b e i ng e qu a l u p w a rds a nd d o wn wa rds . A l w a y s t h e n,

,
m ore s a w ill be e x p e ll e d i n o ne di rec ti on th a n i n t h e o th er Bu t i n

p .
wh i ch e v e r di re c t i on le as t s ap i s e x p e ll e d , from t ha t s am e di recti o n
m o s t s a p w ill re tu rn wh en th e v es s els a re re li e v e d fro m pr ess u rcfi t he
force w h i ch i s p ow erfu l i n a rre s t i ng th e b a c k c u rre nt i n th a t di recti o n
b e i ng th e s a m e forc e w h i ch i s p o w erful in p rod u c i ng a forw a rd cu r
re nt . Ordi naril y , th e m ore a b und a nt s upply of liqu i d b e i ng from be
l o w , th ere w ill re s ul t a n u p w a rd cu rr ent A t ea ch b e nd a p orti on o f
.
t h e c ont e nts wil l be s qu eez ed o u t th rou gh th e s id e s o f th e v es s el s— a

p ort i on w ill be s qu e ez e d d ow nw a rds , re v ers i ng th e cu rrent as ce ndi ng
fro m th e root s , but s oon s topp e d by i ts re s i s ta nc e w hi l e a l a rg e r po r
t i o n w ill be s que e z e d u p w a rd s t ow a rd s th e e x tre m iti es of th e v es s el s ,
w he re c o ns um pt i o n a nd loss a re m os t ra pid . A t ea ch rec oil th e v es
s el s w ill be re pl eni s h e d , ch i efly by th e re p r es s e d up w a rd c u rrent a nd
a t t h e ne x t b e n d m ore o f i t w il l b e t hrus t o nw a rd s th a n ba ck w a rd s .
H enc e w e h a ve e v e ryw h e re i n a cti on a k i nd of ru d e forc e p um p , -
w ork e d by th e w i nd ; a nd w e s e e h ow s ap m ay thu s be ra i s e d t o a
h eigh t fa r b e y ond th at t o w hi ch i t cou ld be ra i s e d by ca pill a ry ac
t i o n, a id ed by o s m os e a nd e v a p o ra t i on .
Thu s fa r, h ow e v er, th e a rgu m ent p ro ce e d s on th e as s u m p ti on tha t

there i s liqu id e n ou gh t o r epl eni sh e v ery ti m e th e v ess els s u bj ec t
i p
t o th s ro c es s . B u t s u pp os e t h e s u pply f a il s — s upp o s e th e root s
h a v e e x h a u st e d th e s u rrou nd ing s to c k of m o i st ure . E v id e ntly th e

v ess el s thu s rep ea te dly h a v i ng th e i r c onte nt s s qu e ezed ou t i nt o the
s u rro u nd i ng ti ss u e , c a nn ot g o o n refilli ng th e m s el v es from o th e r
v e s s els w i thou t t ending t o em p ty th e v as cu l ar sy s tem On th e o ne .
h a nd , e v a p ora t i on from th e l e a v es c a u s i ng a d raugh t o n th e c a pilla ry

t u bes th at e nd i n th em , c onti nu ally g e ne rat e s a c apill ary t ens i on u p
w a rd s ; wh il e , o n th e oth e r h a nd , th e v es s e ls b el ow , e xp a ndi ng a ft e r
t h e i r s a p h as b e en s qu ee ze d out , p ro d u ce a t e ns i on both u pw a rds
a nd d ow nw ard s to w a r d s th e p o i nt o f l o s s . W ere th e li m iti ng m em
b ra ne s ofth e v e s s el s i m pe rm eabl e, t h e m o v em e nt of s a p w ould , u nd er
th e s e c ondit i ons , s o on b e a rres t e d . B u t th es e m em b ra nes are pe rm e
a bl e a n d th e s u rrou ndi ng ti s s u es re a dily p e rm i t t h e p a ss ag e of a i r .
T h i s s t a t e of t ens i o n th e n, w ill c a u s e a n e nt ra nc e o f a i r i nto th e tu b es ;

th e c olu m ns o f liqu i d th ey c o nt a i n w ill be i nt e rru p t e d by bu bbl es .
I t s eem s , i nd e e d , n ot i m p ro b a bl e th a t th i s e nt ra nc e of a i r m ay t ak e
5 84 A PP E N D I X 0 .
pl a ce e v en w h e n the re i s a go o d s u pply of liquid i f the m echa ni c al ,
s tra i ns a re s o v i ol ent a nd th e e x u d a t i o n s o ra pid th a t th e cu rre nts
c a nn ot re fil l th e h alf e m p t i e d v ess e l s w i th su fii c i e nt ra pidi ty

-
A nd .
i n t hi s cas e th e i nt rudi ng a i r m a y p os s ibly pl ay th e s am e p a rt a s

t ha t c ont a i ne d i n th e a i r c h a m be r of a forc e p u m p — te ndi ng by
- -
,
m od era t i ng th e vi ol e n c e o f t h e j e t s a nd by e qu a li z i ng t he s tra i ns
, ,
t o p re v e nt ru p tu re of th e a pp ara t us Of cou rs e w h e n th e s up ply

.
o f liqu i d b eco m es a d e qu a t e a nd th e s tra i ns not t o o vi o l ent t h e se

, ,
b ub bl es w ill be e xp el l e d as rea dily as they e ntered .
H e re as b efore l et m e a dd the conclu s iv e p roof furni s he d by

, ,
a di re ct e xp eri m ent T o as ce rta i n th e a m ou nt o f th i s p r op u l s i v e

.
a ct i o n I t oo k from th e s a m e t re e a L a u rel t w o e q u a l s h oots a nd

, , , ,
pl a ci ng the m i n th e s a m e dye s u bj e ct e d th e m t o c ondi ti ons tha t

,
w ere a lik e 1 n all res p ects s a v e th a t of m ot i o n : w h il e one r em a i ne d

a t re s t t he o th e r w a s be nt b a c k w a rd s a nd forw a rd s no w by s w i t ch
, ,
i ng a nd no w by s t ra i ni ng w i th th e fi ngers A fter th e l a p s e o f a n
.
h ou r I fou nd th a t th e dye ha d as c e nd e d th e o s cill a ti ng s h oot th ree

,
t i m e s as fa r a s i t h a d as cend e d t h e s t a ti ona ry s h oot— t hi s re s u lt

b ei ng a n a v erag e fro m s e vera l tri a ls S i m il ar tri als b rou ght out .
s i m i l a r effec ts i n oth e r s t ruc tu re s T h e v a ri o us p eti ol es a nd h erb a

.
ce ou s sh o ot s exp eri m e nt e d u p on for th e p u rp os e of a s cert a i ni ng
th e am ou nt of exu d ati on p rodu c e d by t ra ns v ers e s tra i ns s h o we d ,
a l s o t h e a m ou nt o f l ongi t u di na l m o v em e nt I t w as o b s erva bl e .
t h at th e h e ight as ce nd e d by th e dye w as i n a l l c as e s g rea ter w h e re

th ere h a d b e en o s cill a t i o n th a n w h e re th ere h a d b e e n res t— t h e
di fference h o we v er b ei ng m uch l ess m a rk ed i n s u ccul ent s tru e
, ,
tu re s th a n i n w oo dy o n es .
I t ne e d s c a rc ely be s a id th a t thi s m e ch a ni cal a cti o n i s not h e re

a ss ig ne d as t h e s o l e c a u s e o f c i rc u l a ti o n but as a c a u s e c o op era t i ng
,
-
w i th oth e rs a nd h elpi ng oth e rs t o p ro d u ce effe ct s th a t c ould no t

,
oth erw i s e be p ro d u ce d T re es g row i ng i n cons e rv a t ori e s a fford u s

.
a b u nd a nt p roof th a t s a p i s ra i s e d t o c o ns id e ra bl e h ei hts by o th er
g
forc es . T h ou gh i t i s not ori ou s th at t re es s o c i rcu m s ta nced d o not
th riv e unl es s t h ro u gh o p e n s ash e s th ey a re fre q u e ntly s u bj e ct t o
, ,
b re ezes s uffi ci ent t o m ak e thei r p a rts os cilla te yet there i s e v id e ntly ,
a c i rcu l a ti on th a t g oe s o n w i th o u t m e ch a ni c a l a i d T he ca u ses o f .
c i rcul a t i o n are th os e a c ti ons o nl y w h i ch d i s tu rb th e l l qm d e u 1 11 br1 u m

q
i n a pl a nt by p e rm a ne ntly a bs t ra ct i ng w a te r o r s ap fro m s om e p a rt
,
o f i t ; a nd o f t h es e t he fi rs t i s th e a b s o rp ti on o f m a t e ri a l s for th e fo r
m a ti on o f ne w t i s s u e i n g row i ng p a rt s th e s e co nd 1 8 th e l oss by
e v a p ora t i o n m a i nly th ro u gh a d ul t l ea v e s ; a nd the thi rd 1 s th e l o ss by
,
e x t ra v a s a t i o n th ro ugh c o m p res s e d v ess el s

, Only s o far a s i t pro
.
d uces th i s l as t ca n m ech a ni c al s tra i n be rega rd ed a s t ruly a c a us e o f

,
c i rc u l a t i o n A l l the o th er a cti o ns c onc erne d m u s t be cl a s s e d a s a ids

.
t o c i rc ul at i o n— a s fa cili t a ti ng th a t re di s t rib u t i on of liq u id th a t c o n

t i nu a l ly res tores th e e q u ili bri u m co nt i nua l l v di s tu rb e d a nd of th e s e
CI R C U L A TI ON A ND F OR MA TI ON OF WOOD I N P L A NT S .
5 85
ca pil l ary a cti on m ay be na m e d as th e firs t o s m os e a s th e s e cond a nd

, ,
th e p ropu ls i v e e ffect of m e ch a ni c al s tra i ns a s th e th i rd T h e fi rs t .
t w o of t h ese a id s a re d o u b tl es s c a p a bl e by th em s el v e s of p rod u c i ng
a l a rg e p a rt o f th e o b s er v e d res ul t— m ore o f th e o b s e r v e d res u l t th a n
i s a t fi rst s ight m a ni fes t ; for t h ere i s a n i m p o rt a nt i ndi rec t effect

o f os m oti c a c ti o n w h i ch a p ears t o be o verl ook e d Os m o s e d o e s no t
p .
a i d c i rc ul a ti on o nl y by s e tti ng u
p w i th i n th e pla nt exc ha ng e c urre nt s
, ,
b et w ee n th e m ore d ens e a nd th e l es s d ens e s olu ti ons i n di fferent p a rt s

o f i t ; bu t i t a id s c i rcul a t i o n m u ch m ore by p ro du c i ng d i s te nt i on
o f th e pl a nt as a w h ol e . I n c o ns eq u e nc e o f th e a v era g e c o nt ras t i n
d e ns i ty b et we en th e w a ter ou ts id e o f th e pl a nt a nd th e s a p i ns id e of
i t t h e cons ta nt t end ency i s for th e p l a nt t o a bs orb a qu a nti ty i n e x c es s
,
o f i t s c a pa c i ty a n d s o t o p ro d u c e di s te nt i o n a nd erecti o n of i t s
,
t i ss u es . I t i s b e c a u s e o f t hi s th a t t he d roopi ng pl a nt r a i s e s i ts e lf
w h e n w a t e re d for c apill ary a ct i o n a l one co uld only refill i t s t i s s u es
w i t hou t c h a ngi ng th e i r a tti t ud e s .A nd i t i s b e c a us e o f th i s th a t
j u i cy pla nts w i th c olla p s ibl e s t ru ctu re s bleed s o ra pidly w h en cu t not ,
o nly from th e c u t s u rfa ce o f th e ro ot e d p a rt bu t fro m t h e cu t s u r

,
fa c e of th e d e ta ch e d p a rt th e el a st i c t i ss u es t e ndi ng t o p re s s o u t t he
-
liquid wh i ch di ste nd s th em . A nd m a ni fes tly i f os m os e s erv es thu s

t o m a i nt a i n a s t ate of d i s t enti o n th rou gh out a pl a nt i t i ndi rectly fu r ,
t he rs c i rc u la t i on ; s i nce i m m e di a t ely e v a p ora ti o n or g row th a t a ny

p art by a b s tra cti ng liq uid from th e neig hb ou ri ng ti s su e s begi ns to
, ,
di m i ni sh th e liqu id p res s ure w i th in s u ch ti ssu es the di s t end e d s t rue

,
t u res th rou gh ou t t h e res t of t h e pl a nt thrus t th e i r liq u id c onte nts t o

w ard s th e pl ace of di m i ni sh e d p re s s u re Thi s i ndee d m a y v e ry pos
.
, ,
s ibl y be th e m os t ef fi ci e nt of the ag enc i es a t w ork Rem em beri ng .
ho w grea t i s t he di s t enti on p ro d u ci ble by o s m oti c a bs orp ti on— g re at

e nou gh to b urs t a bl a d d e r — i t i s c l ea r t h a t th e force wi th w h i ch th e
di s t end e d tis su es of a pla nt u rge forwa rd th e s a p t o pl a ces o f c on
s u m p t i on i s p ro b a bly v ery g re a t
,
. W e m us t th ere fore rega rd the a id
w h i c h m e ch a ni c a l s t ra i ns gi ve as b ei ng one of s e vera l Os cilla ti ons .
h elp di re c tly t o res tore a ny d i s turb e d li qu id e qui lib ri u m ; a nd t h ey

a l s o h el i ndi r ectly by fa c il i ta ti ng th e redi s t n buti o n c a u s e d by c a p i l
p ,
l a ry a cti on a nd th e p roces s j u st d es crib ed ; but i n th e a bs e nce of

os c ill a t i o ns t h e e qu i l ib riu m m a y s till be res tore d th ou gh l es s ra pidly
,
a nd w i th i n na rrow e r li m i ts of di s t a nce .
O ne h alf of th e p ro bl em o f th e c i rcul ati on h ow e v er h as b een left

, ,
ou t of s igh t . T h u s fa r ou r i nqu i ry h a s b e en h ow th e as cendi ng cu r

,
r e nt o f s a
p i s p r o d uc e d
. Th e r e re m a i ns th e ra t i o na l e of th e d es c e nd
i ng cu rre nt . W h at forc es c au s e i t a nd th rough w h at ti ss u es i t ta k es

,
pl a c e a re qu esti ons to w hi c h no s ati sfa ct ory a ns w e rs h a v e be en

,
g i v en. T h a t th e d es ce nt i s d u e t o gr a v i t a t i o n as s om e a ll eg e , ,
i s di ffi c u lt t o c onc e i v e s i nc e as gra v i t a ti on a ct s e qu ally o n a ll

, ,
liqu i d c ol u m ns c ont a i ne d i n th e s t e m i t i s n ot e a s y t o s e e w hy
,
i t sh ou ld p rod uce d o wnw a rd m o v e m e nt s i n s om e wh il e p e rm i tti ng

586 A PP E N D I X C .
u
pw a rd m v
o e m e nts i nd ee d th ere e x i s te d d es c end
i n o th e rs — l
u n ess , ,
i ng tu b es t o o w id e t o a d m i t of m u ch c a pill a ry a ct i on w h i ch th ere ,
d o not More o v e r g ra vi ta ti o n i s cl e a rly i nad e qu a te to c a u s e c u r

.
,
re nt s t o w a rd s t h e roo ts ou t of b ra n ch es th a t d ro op t o th e gro u n d .
H ere th e g ra v i t a t i on o f th e c onta i ne d liqu id c olu m ns m u s t nearly

ba l a nc e t h a t of t h e c o nnect e d c ol u m ns i n th e s t em l ea v i ng no ,
a pp re c i a bl e fo rce t o c a u s e m o ti o n Nor d oes th ere s e em m u ch .
ro ba b il l t y i n t h e as s um p t i o n th a t th e rou t e o f t h e d es c e ndi ng s a
p p
i s th rou gh th e c a m bi u m lay e r s i nc e e x p e ri m e nts o n th e a b s orp ti on
,
o f d es p ro v e th a t s i m pl e cell u l a r t i ss u e i s a v e ry b a d c o nd u c tor
y
o f liqu ids : th ei r m o v e m e nt th ro u gh i t d oe s no t t a k e pl ac e wi t h o ne
fifti e th of th e ra pidi ty w i th w hi ch i t t ak es pla c e throu gh v ess els *

.
Of c ou rs e th e d efenc e for th es e hyp oth e s e s i s th a t th ere m us t be ,
a d o w nw ard cu rre nt w h i c h m u s t h a v e a cou rs e a nd a cau s e ; a nd th e

,
v ery natu ral as s u m p ti on h as been th a t th e c ours e a nd th e cau s e m u s t

be o th e r t ha n t h os e wh i ch p ro d u ce t he as c e ndi ng cu rrent N e ve r .
t he l es s th e re i s a n a l te rna ti v e s upp os i ti o n t o w hi ch t he foregoi ng

c o ns id e ra ti o ns i ntro d u ce u s I t i s q u i te p o s s ibl e fo r th e s a m e v as cu l a r
.
s y s t e m t o s e r v e a s a ch a nn el fo r m o v e m e nt i n o pp o s i te d i re c ti o ns
a t di f ferent ti m es W e ha v e a m ong a ni m al s w ell k no wn cas es i n

.
-
w h i ch th e bl o od v es s el s ca rry a c u rre nt firs t i n o ne di re c ti o n a nd

-
t h en a ft er a bri e f p a u s e i n the re v e rs e di re cti on

, ,
A nd th ere s e e m s .
a n a p r i or i p r o b a bili ty th a t l o w ly o rg a ni ze d a s th ey a re pl ants a re
, ,
m ore lik ely t o h a v e di st ri buti ng a ppli a nc e s of th i s i m p erfec t ki n d

t ha n t o h a v e t w o s ets of ch a nnel s for tw o s i m ul t a ne ous c urre nt s If .
,
l e d by thi s s u s pi ci o n w e i nqu i re wh e th er a m ong th e forc es w h i c h

,
u ni te t o p ro d u ce m o v e m e nts o f s a t h e re a re a n v i t i n i nt er
p y a r a o , s or
m i s s i o ns c a p a bl e of d et e rm i ni ng th e c u rr ents i n d i f fere nt di rec ti o ns ,
w e qu i ckly di s co v er t h a t t here a re s u ch a nd th a t the h yp oth e s i s o f ,
a n a lt erna ti ng m o ti on of t h e s a n o w c e nt ri fu g al a nd n o w c e nt ri
p ,
p etal throug h the s a m e v ess el s h as goo d w arra nt W h at a re the

, , .
s e v era l fo rc es a t work ? Fi rs t m ay be s et d o w n th a t te nd ency

e x i s ti ng i n e v e ry p a rt of a pl a nt t o e x p a nd i nt o i ts typi ca l fo rm a nd ,
t o a bs o r b nu tri t i v e liqu id s i n d oi ng th i s Th e res ul ti ng c o m p eti t i o n.
So m e e xce pti ons

to thi s occur i n p l a nts th a t h a e retrorD gra ded i n th e v
ch a ra c t e r o f th e i r t i s s ue s to wa rds th e s i m p l e r e g e ta l t yp e s Cc rt a i n e ry v . v
v
s uc c ul ent l e a e s , s u ch a s t ho s e of S em
p m zm m , i n wh i ch the ce l l ul a r ti s s ue
'
v
i s i m m ens el y de el ope d i n com p a r i son wi th th e a s c ul a r t i s s ue , s ee m t o v
v x
h a e re s um ed t o a c ons ide ra bl e e tent w h a t w e m us t reg a rd a s th e p ri m i t i e v
form of e ge ta l c i rcul a ti on— s i m p l e ab s or pt i on fro m c e l l t o c el l
v Th e s e, .
v
when t h ey h a e l os t m uch o f t h ei r wa te r, w i l l t a k e u p t he dye to s om e di s
t a nce t hrough t he i r g e nera l s u bs ta n c e, or ra the r th ro ug h i ts i nte rs t i ce s , e e n v
neg l ec t i ng t h e v
e s se l s A t oth e r t im e s , i n th e s a m e l ea es th e e s s e l s w i l l
. v v
v
b eco m e c h a rg e d whil e com pa ra t i el y l i t tl e a bs orpt i on t a k es p l a ce t h rough
t h e ce l l ul a r t i ss u e v x
E en i n th e s e e c ep ti ona l ca s e s , h owe e r, t h e m o e m e nt
. v v
t hrou gh ce l l ul ar t i s s ue i s noth i ng l i k e a s fa s t a s th e m o em ent t hrou h
g v
v e s s el s .
CI RC U L A T I ON A N D FORMA TION OF WOOD I N P L A NT S .
5 87
for sa
p w ill , o th er th i ngs b ei ng e qu al , c a u s e cu rr e nts t o w a rd s th e
m o s t ra pi dly gr o w i ng p a rt s — t o w a rd s u nfoldi ng s h o o t s a n d l ea v es
-
.
but n o t t o w a rd s a d u lt l e a v es . Ne x t w e ha v e e v a p ora ti o n, a cti ng

m ore o n th e a d u lt l ea v es t h a n o n t h os e w hi c h a re i n th e bu d ,
o r but p a rt i a lly d e v e l o e d Th i s e v a p o ra t i o n i s b o th regul a rly
p .
a n d i rregu l a rly i nt erm i t t e nt . D ep e ndi ng ch i efly on t he a ct i o n '
o f th e s u n, i t i s , i n fi ne w ea th e r, grea tly ch eck e d or w hol ly

a rre s t e d e ve ry e v e ni ng ; a nd i n c l o u dy w e a th er m u s t be m u c h
r eta rd e d d u ri ng th e d a y . Fu rth er, e v ery h y g rom et ri c va ri a ti on,

a s w ell as e ve ry v ari a t i o n i n t h e m o v em ent o f t h e a i r, m u s t
v a ry th e e v a p ora ti on Thi s ch i ef a ct i o n, therefore , whi ch , by c on
.
t i nu a l l y e m ptyi ng t h e e nd s of t h e c a pilla ry t u bes , m a k es u p w a rd

c u rre nts p oss i bl e i s on e w h i c h i nt erm i t s e v e ry nigh t , a nd e v ery d a
,
y
i s s t rong o r fe e bl e as c i rc u m s t a nc es d e t e rm i ne . The n, i n th e th i rd
pla c e, w e h a v e thi s ru d e pum pi ng p ro c es s a b o v e d es cri be d g oi ng
’
o n w i th g reat e r v ig o u r w h en t h e w i nd i s v i o l e nt , a nd w i t h l es s
v igo u r w h en i t i s gentl e — d ra w i ng li qu id towa r ds di ffe re nt

p a rts a ccordi ng to the i r d egr e es o f o s c ill a ti on, a nd fr om d i ffe
r e nt p a rt s a c c o rdi ng a s th ey c a n m o s t rea dily furni s h i t . A nd
n ow l et u s as k w h a t m u s t res u l t u nd er c h a ngi ng condi t i ons fro m
t h es e v ari ou s ly - c o nfl i c t i ng a nd c ons pi ri ng forces . I Vh e n a w a rm

s u ns h i n e , ca u s i ng ra pid e v a p ora t i on , i s e m p t y i ng th e v e s s e l s o f th e
l ea v es , th e osm oti c a nd c apillary a c ti ons t h at refill t hem w ill be

c o nt i nu a lly a id e d by t h e p u m pi ng a ct i o n of t h e s w a yi ng p eti o l e s ,
t w i gs , a nd b ra nch e s , p ro vid e d th ei r o s c il l a ti o ns a re m o d e ra te U n d e r .
th es e c o ndi ti o ns th e curr e nt of s a p , m o v i ng i n th e di re c t i o n of l e as t
res i s t a nc e , w ill s et t o w a rd s th e l ea v es . Bu t w h at w ill h app e n w he n
t h e s u n s e t s ? Ther e i s no w n othi ng t o d et erm i ne c u rre nts e i th er
u p w a rd s o r d ow nw a rd s , e x c ep t th e re l a t i v e ra t es of g ro w t h i n th e
p a rt s a nd th e rel ati v e d em a nd s s et u p by th e o s c il l a t i ons ; a nd th e
o sc i l l a t i o ns a ct i ng al on e , w ill d ra w s a p t o th e os c ill a t i n g p a rts as
m u ch from a b o v e as from b el o w . If th e res i s ta nc e t o b e o v erc om e

b y a c u rre nt s ett i ng b a ck from th e l ea v es i s l e s s t ha n t he res i s t a nc e
t o be o v e rcom e by a c u rre nt s et ti ng u p from th e ro ots , th e n a
c u rre nt w i l l s e t b a c k from th e l e a v e s . No w i t i s , I t h i nk , t ol era bly
m a ni fes t th a t i n t h e s w a yi ng tw ig s a nd m i nor bra n ch es , l ess fo rc e
w ill b e re q u i re d t o o v erc om e th e i ne rti a o f th e s h ort c ol u m n s of
liqu id b e tw e en th em a n d the l ea v e s th a n t o o v e rco m e t he i nert 1 a of

th e l o ng c olu m ns b et wee n th em a nd t he r oot s . H e nc e d uri ng th e
nigh t , as a l s o a t o th e r t i m e s wh e n e v a pora t i on i s n ot g o i ng o n, th e
s ap w ill b e d r a w n o u t o f t h e l e a v es i n t o t h e a dj a ce n t s u
pp o rt i ng
p arts : a nd th ei r nut riti on w ill be i nc re as ed If t he w i nd 1 s s trong
.
e n ou gh t o p r o du ce a s w a yi ng o f t h e th i c k e r b ra n c h es , th e b
ack
c u rre n t w i l l e x t e n d t o t h e m a l s o ; a n d a f u r th er s t re n
g t h e ni ng w ill
re s u lt f r o m t h e i r a b s o rp ti o n o f t h e e l a b o r a t e d s a
p . A n d w h e n t h e
rea t b ra nch es a nd th e s te m a re b ent b a c k w ard s a nd for w a rd s by

a
g
588 A P P E N D IX 0 .
ga le , th ey t oo will sh a re i n th e nu t r i ti o n
firs t s ight s eem
. It m a
y at
t h a t t h es e p a rt s be i ng n ea re r t o t h e roots t ha n t o t h e l ea v e s w ill
, ,
d ra w thei r s uppli es from the roots o nly Bu t th e qu a nt it y whi c h the .
r o ot s c a n fu rni s h i s i ns u f fi c i ent to m eet s o grea t a d e m a nd Und er .
t h e condi ti ons d es c rib e d t he e x u d a ti on of s a p fro m t h e v es sel s w ill

,
b e v ery g rea t a nd th e d ra ught of liqu id re qu i re d to refill t he m not

, ,
s a ti sfi ed by t h a t w h i c h th e ro o t fi bre s c a n t a k e i n w ill e x te nd to th e
-
,
lea v es Th us s ap will fl o w to th e s everal p a rt s a c c o rdi ng to the i r

.
p
r e s ec t i ve d e g re es o f a c t i v i t y — t o t he l ea v e s wh il e lig ht and h e a t
e na bl e th em t o di s c h arg e th e i r fu n ct i o ns a n d b ac k t o th e t wigs , ,
bra nch es s t e m a nd ro ots wh en th es e b ec om e a cti v e a nd t he l ea v es

, ,
i na cti v e o r w h en th ei r a cti vi ty d om i nates o v er tha t of th e l e a v es

,
.
A nd thi s di s t ri bu ti o n of nu tri m ent v a ryi ng w i th th e varyi ng ,
a ct i v i ti es of t h e p a rt s i s j u s t s u ch a di s trib u ti o n as w e k no w m u s t
,
b e req u i re d t o k e ep u p th e o rga ni c bal a nce .
T o t hi s e x pl a na t i o n i t m ay be obj ect e d th a t i t d oes not a c c ou nt

for t h e d ow n w a rd c u rre nt of s a p i n pl a nts tha t a re s h e l t ere d The .
s t em a nd ro ots of a d ra w i ng ro o m Ge ra ni um d i s pl a y a t hi ck e ni ng
-
w hi ch i m pli es th a t n u tri t i v e m a tt ers ha v e d es ce nd ed from th e l ea v es ,
a l th o u gh th ere a r e no ne o f th e s e os cill a t i ons by w h i c h th e s a p i s s a id
t o be d ra w n d ow nwa r ds as w e l l as u p w ard s Th e reply i s t ha t th e .

,
s tem a nd ro ots te nd t o r ep ea t th e i r t y pi c a l s tru ctu re s a nd th a t t h e ,
a bs orp t i o n of s a p for t h e form a t i on o f t h ei r r es p e ct i v e d e ns e t i ss u e s ,
i s h ere th e forc e wh i c h d e term i ne s t h e d es ce nt I nd e ed i t m us t be .
b orne i n m i nd th a t th e m e cha ni ca l s tra i ns a nd th e p u m pi ng p roc ess

w hi c h t h ey k e ep u p a s w ell as t he di s te nti on c a u s e d by os m os e d o
, ,
not i n th e m s el v e s p r o d u ce a cu r re nt e ith er u p w a rd s o r d o w nw a rds :

t he y s i m ply h e lp t o m o ve th e s a p t ow a rd s t h a t pl a c e w h e re the re i s
t he m os t rapid a bs tra c t i o n o f i t — th e plac e t ow a rd s w h i c h i ts m oti o n
i s l eas t re s i s t ed Wh eth er th ere i s os cillati on o r wh e ther th ere i s
.
no t t h e phy s i ol og i ca l d em a n d s of th e di f
,
fere nt p a rts of th e pl a nt
d et erm i ne th e di recti on of t he current ; a nd a l l w hi ch the os cilla t i ons
a nd th e di s t enti o n d o i s t o fa c ili t a te t h e s u pply o f th es e d e m a nd s .
Jus t as m uch th erefore i n a pla nt a t res t as i n a pl a nt i n m oti on

, , ,
t h e c u rre nt w ill s et d ow n w a rd s w hen th e fu nc ti o n of th e l ea v es i s

a rres te d a n d w h en t h ere i s not hi ng to res i s t t ha t a b s tra cti o n of s a
, p
c a us e d by th e t end enc y o f th e s te m a nd ro ot t i ss u es to as s um e t h e i r -
t ypi c a l s tru c tu re s T o w hi ch a d m i ss i on h ow e v e r i t m u s t be a dd e d
.
, ,
t h a t s i nc e thi s typi c al s tru c tu re as s u m e d t h ough i m p e rfe ctly as ,
s u m e d by the h o t h ou s e pl a nt i s i ts e l f i nt erp ret a bl e as th e i n

,
-
,
h e ri t e d e ffec t of e x ternal m e ch a ni c a l a cti ons on i ts a nc es tors w e ,
m a y s t ill c o n s id er t h e c u rre nt s et up by t h e as s um p t i o n of th e
typi c al s tru ct u r e t o b e i ndi rectly d u e t o s uch a cti o ns .
I nteresti ng e v id e nce of a nother o rd er h e re d em a nds noti ce I n .
t h e c o u rs e o f e xp eri m e nts o n th e a b s orpt i on o f dy es by l ea v e s i t ,
happ ened th a t i n m a ki ng s ecti o ns p ara ll el t o th e pl a ne of a l e af w i th ,

CIR C U L AT I ON A ND F OR MA TI ON OF WOOD I N P L A NT S
’
.
5 89
th e v i e w of s ep a ra ti ng i ts m iddl e l a y er co ntai ni ng the v ess els I c am , e

u p on s om e s tru ctu res t ha t w ere ne w t o m e Th es e s tru ctu res w here .
,
t he y a re p res ent fo rm th e t e r m i na ti ons of the vas c u l a r s y st e m

, Th e y .
a r e m as s e s of i rre g u l a r a nd i m p e rfe c tly u ni te d fi bro u s c ell s s u c h as ,
t h o se out of w h i c h v e s se ls a re d e v el op e d ; a nd t h ey a re s om e ti m es
s l e nd er s om et i m e s b ulky — u s u all y h o we ve r b e i ng m ore or l es s cl u b
, , ,
s h a p ed I n tra ns v ers e s ect i ons of l ea ves th e i r di s ti nc ti v e cha ra ct e rs

.
a re not s h o wn : th ey a re ta k en for t h e s m a ll er v e i ns I t i s o nly by .
c a re fu lly s li c i ng a w a y th e s u rfa c e of a l e af u ntil w e c om e d o w n
t o th a t p a rt w hi ch c o nta i ns th e m th a t w e g et a ny id e a of th e i r ,
n a t u re Fig 1 rep res ents a s p e ci m en t a k en from a l e af of E u

. .
p h or bi a neri ifol i a O cc u pyi n

g o ne of th e i nt e rs p a c e s o f th e u l ti
.
m a te v e no u s net w ork i t c o n s i s ts o f a s p i ra lly li ne d d u ct o r s et o f

,
-
d u cts whi ch c onne cts w i th th e neigh b ouri ng v ei n a clus t er of h alf

,
re ti cu l a t e d h a l f s c al a ri fo r m c ells Th es e c ell s h a v e pro e ct i ons m a ny

,
-
j .
,
o f t h e m ta p e ri ng th a t i ns ert th em s el v es i nt o th e a dj a ce nt i nt e r cell
,
u l a r s p a c es t hu s p r o d u ci ng a n e xte ns iv e s urfa ce o f c o nt a ct b e t w e e n
,
th e org a n a nd th e i m b e ddi ng ti s s u es A fur th e r tra i t i s th at th e e n .

,
s h e a th i ng p ros e nchy m a is e i th er bu t li ttl e d e v el op e d o r wh olly a h
s e nt ; a n d c o ns e q u e ntly th i s e x p a n d e d v as cu la r s t ru c tu re e sp ec i a l l y ,
a t i ts e nd com es i m m e di a tely i n c o nt a ct w i th th e t i s s u es c o nc e r ne d
,
i n as s i m il a ti o n T h e l e af of E up hor bi a nerzifol i a i s a v ery fles hy

'
o ne ; a nd i n i t th es e o rg a ns a r e d i s trib u te d th r ou g h a com p a ct ,
t h ou gh w a t ery c el lu l ar m as s B ut i n a ny l e a f of th e ordi na ry typ e

,
.
w h i ch p os s ess es th em th e y l i e i n th e netw ork p are nchy m a c om p os i ng

,
i ts l o w er l a y er ; a nd w h e re v e r t h ey oc cur i n th i s l a y er i ts c ell s u ni t e
t o e ncl o s e t h em Thi s a rra nge m e nt i s sh ow n i n fig 2 rep res enti ng
. .
,
a s a m pl e from th e C a o ut c h ou c l e af a s s ee n w i th th e u pp er p art o f
-
,
i t s e n v elop e re m o v e d ; a nd i t i s sh own s ti l l m ore cl early i n a s a m pl e

from t he l e a f o f P a na cc L essom z fig 3 Figures 4 a nd 5 rep res ent
' ’
. .
, ,
w i th ou t th e i r s h e a th s oth er s u ch o rg a ns from th e l ea v es o f P a na x
,
L ess om i a nd Cl u sza fl a va S om e rel a ti on s eem s t o e x i s t b etw e en

' ’
th e i r fo rm s a nd th e th i c kn es s es o f t he l a y ers i n w hi c h th e y l i e .
C e rt a i n v e ry th i ck l e a v es s u ch a s th os e of Ul u si a fl a va h a v e th e m
, ,
l ess abu nd a ntly di st rib ut e d th a n i s u su al bu t m ore m a s s i v e W h ere , .
t h e p are nchy m a i s d e v el op e d not t o s o gr e at a n e x trem e th o u gh ,
s til l l a rg e ly as i n th e l ea v e s of H o lly A u cu ba Ca m el l i a t h ey are

, , , ,
no t s o bulky ; a nd i n t h i nn e r l e a v es lik e th os e o f P riv e t E ld er , , ,
& c th ey b e c om e l onger a n d l e s s c o ns pi cu ou s ly c lu b s h a p e d S om e
.
,
-
.
a d a p t a t i ons to th e i r res p e c ti v e p o s i t i o ns s eem i m pl i e d by t h es e m o di
fi cat i ons ; a nd w e m a y na tu ra l l y e x p e c t th at i n m a ny th i n l e a v e s
th es e fre e e nd s b e c om ing s till na rrow er lo s e th e di s ti nc ti v e a nd
, ,
s u g g e s t i v e c h ara ct ers p o ss ess e d by th o se s h o w n i n th e di a gr am s .
Rel a t i ons of th i s ki nd a re no t regul ar h ow e v er I n va ri ous o th er ,

.
ge nera m em b e rs of wh i ch I h a v e e x am i ned as R i ms Vi bu rnu m

, , , ,
Gri sel i m a B rezvi a B otryod endron P er es kia th e v ari ati ons i n th e

'
-
, , , ,
590 A P P E ND IX 0 .
b u lk form of th es e s tru ctures are not di re ctly d eterm i ned by

a nd
th e s pa ces w h i ch th e l e a v es a ll o w : o b vi ou sly th e re a re o ther m o di

fyi ng c au s e s I t s hould be a dd e d tha t w hil e t h e s e e x p a nd e d free
.
e x trem iti e s g ra du a te i nto t a p eri ng fre e

xtrem i ti es not differi ng e ,
from ordi na ry v e ss els th ey a ls o p a s s i ns e ns i bly i nt o th e o rdi na ry

,
i nos cul ati ons Oc ca s i onally a long w i th nu m e rous fre e e ndi ngs
.
, ,
t h e re o cc u r l o op s ; a nd from s u ch 100ps th e re a re t ra ns i ti o ns t o
t h e u lti m a t e m esh es of th e v e i ns .
T h es e o rg a ns a re by no m ea ns c om m on t o a l l l e a v es I n m a ny .
th at a ffo rd a m pl e s p a c es for th em t hey a re not t o b e fou nd S o far .
as I h a v e o bs erv e d th e y a re a b s e nt from th e th i c k l e a v e s o f pl a nts

,
w hi ch fo rm v e ry li ttl e wo od I n S em p er m vu m i n E chever i a i n
’
.
, ,
B r y op hy l l u m t hey do not a ppea r t o e x i s t ; a nd I h a ve be en u na bl e

,
t o di s c o v e r th em i n K a l a nchoé r o tu ndifol i a i n K l ei nia a n te eup hor ,

-
bzu m a nd fi cozdes i n th e s e v e ra l s p ec i es of Cra ss u l a a nd i n oth er

' ’
, ,
s u ccu l e nt pl a nts I t m ay be a dd e d t ha t th ey a re not a b s ol utely

.
c o nfi ne d to l e a v es but occu r i n s t e m s t h a t h a v e a s s u m e d the fu nc

,
t i ons of l ea ves A t l eas t I ha v e fou nd i n th e g reen p a re nchy m a

.
,
o f Op u n ti a o rga ns th a t a re a na lo go u s t hou gh m u ch m ore ru d ely

,
a nd i rre gu l arly fo rm e d I n oth er p a rts t oo t hat h a v e u su rp e d

.
, ,
t h e l ea f func t i o n th ey occu r a s i n th e phyll od es of the A u s tra li a n

-
, ,
A c a ci as Th es e h a v e t h em a b und a ntly d e v el op e d ; a nd i t i s i nt eres t

.
i ng t o o b s erv e t hat h ere wh e re th e t wo v e rti ca lly pl a ce d s u rfa ces

,
-
o f t he fl a tte ne d ou t p e ti ol e a re e qu a lly a d a p t e d t o t he a s s i m il a tiv e

-
fu nc ti o n t here e xi s t t wo l a y e rs of t hese e xp a nd e d v a s cul ar term i na

,
t i ons one a ppli e d t o th e i nner s u rfa ce of e a ch l a y e r of p a re nchy m a

, .
C ons id e ri ng t he s tructures a nd p os i ti ons of th e s e orga ns a s w ell ,
a s t h e na tu res of t he pl a nts p oss es s i ng t he m m a y w e no t form a ,
s h rew d s u s pi c i o n res p e c t i ng th e i r func t i on I s i t not p ro b a bl e t ha t

t hey fac ili ta te a b s orp ti on of t he j u i ces c a rri e d b a ck from th e l eaf fo r
t h e nu t ri ti on of th e s te m a nd roots i T he y a re a dm i ra bly a d ap t e d
’
for p erform i ng thi s o fli ce Th ei r c om p onent fib rou s c ell s h a v i ng

.
,
a ngl es i ns i nu a t e d b et wee n th e cells o f th e p a re nch y m a are s h a p e d ,
j us t a s they sh ou ld be for taki ng u p i ts c ont e nts ; a nd th e a bs ence

o f s h ea thi ng ti s s u e b et w een t hem a n d the p a renchy m a fac ili ta t es t he
p a ss ag e o f t he ela b orated liqu id s Moreo v er th ere i s th e fa c t th at .
t he y a re alli e d t o o rg a ns w h i c h o bv i ous ly h a v e a b s o rb e nt fu n c ti o ns .
I a m ind e bte d t o D r H ook er fo r p oi nti ng out th e figu re s of tw o

.
s u ch o rg a ns i n th e I c ones A na t om i cze of L i nk One o f th em i s .
from th e e nd of a di cotyl ed o nous root fibre a nd th e oth er i s fro m -

,
t h e p roth a ll us o f a y oung Fe rn I n e a ch ca s e a cl u s ter o f fib rou s .
c ell s s e a t ed a t a pl a c e from w h i ch li u id h a s t o be d ra w n i s c on
, q ,
ne cte d by v es s el s w i t h th e p a rt s t o w h i c h li u id h as t o b e c a rri e d
q .
Th e re ca n s c a rc ely be a d ou b t th en t ha t i n both c as e s a b s orp ti o n , ,
i s e ffe ct e d th r ough th em I h a v e m e t w i th a nothe r s u ch orga n

. ,
m ore el a b orat ely c onstructed b ut e v id e ntly a d a p te d to th e s am e ,

CI RCU L A TI ON A ND FORMA T ION OF WOOD I N P L A NTS .
591
o fli c e, i n th e c o m m on T urnip ro ot
-
A s s h ow n by th e end vi e w
. .
a nd l o ngi t u di nal s e ct i o n i n fi s 6
g a nd 7, th i s o rg a n c ons i s ts of
.
ri n s o f fe nes tra t e d c e ll s , a rra ng e d w i t h v a ryi ng d e r e e s of re g u

g g
l a ri ty i nt o a funnel , o rdi na rily h a v i ng i ts a p e x di re c t e d t ow a rds t he
c e nt ra l m as s of t h e T u rnip , w i t h w h i c h i t h as , i n s om e c as es a t l eas t ,
a t ra c ea bl e conn e x i on by a c a n a l P res e nt i ng a s i t d o es a n e x te rna l
.
p o rou s s u rfa ce t erm i nati ng o ne of t h e bra nch es of the v a s cul ar s ys

t em , e a ch of th es e org ans i s w e ll fi tt e d for t a ki ng u p w i th ra pidi t y
th e nu t ri m e nt l a id by i n th e T u rni p root , a nd u s e d by th e pl a nt
-
w h e n i t s e nds u p i ts fl ow e r s talk -
Nor d o es e v e n thi s e x h a us t th e
.
a na l o gi e s. Th e c otyle d ons of t h e y oung b e an, e x p eri m ent e d u p o n

as b e fo r e d es c rib e d , fu rni s h e d o th er e x a m pl e s o f s u ch s tru ctu res ,
e x a ctly i n th e pl a ces w h er e, if th ey a re a b sorb e nts , w e m ight e x
p eet t o find t hem A m id the b ranchi ngs a nd i nos c ul ati ons of th e

.
v as cula r l ayer runni ng th rough th e m as s of nutri m ent d ep o s i te d i n

e a ch co t yl e d on , th ere a re c o ns pi cu ous fre e t erm i na ti ons th a t a re clu b
s ha p e d , a nd p rov e t o b e c o m p os e d , lik e th os e i n l ea v es , of i rr e gul arly
form e d a nd c lu st ere d fib rou s cell s ; a nd s om e of t h em , di v e rg i ng

from th e pl a n e of th e v a s cu l a r lay er, dip d o wn i nto th e m as s of
s ta rch a n d a lb u m e n w h i ch t he y o ung pl a nt h as t o u t ili z e , a nd
w h i ch t h es e s t ru ctu res c a n h a v e no ot h er fu ncti o n bu t t o t ak e u p .
B es id es b e i ng s o w ell fitt e d fo r a b s orp ti on, a nd b es id es b e i ng

s i m il a r t o org a ns w h i ch w e c a nno t d ou b t a re a bs orb e nt s , th es e v as
c u l a r t e rm i na t i ons i n l e a v es a fi ord u s ye t a noth er e v id en c e o f th ei r
fu nct i ons . Th ey a re s ea te d i n a ti s s u e s o a rra ng e d as s p eci a lly t o

fa c ili t a te the a b s t ra cti o n of li qu id Th e ce nt rip et a l m o v em e nt o f
.
th e s a p m u s t be s et u p by a forc e t h a t i s c om p a ra ti v ely fe eble , s i nc e,

th e p a ri e t e s of t h e d ucts b ei ng p o r ou s , a i r w ill e nt er i f t he t ens i o n
o n t he c onta i ne d c ol um ns b ec om e s c o ns id era bl e H ence i t i s nee d ful
.
th a t th e e xit of s ap from the l ea v es sh ou ld m eet w i th v ery littl e re s i s t

a nc e. N ow w er e i t not for a n a dj u s tm ent p res ently t o be d es cri be d ,
i t w ould m e et w i th g re a t re s i s ta nc e, notwi ths ta ndi ng th e p ecu li a r
fitnes s of th ese o rg a ns to t ak e i t i n L i qu id c a nnot be d ra w n ou t
.
o f a ny c l os e d ca v i ty w i tho u t p ro d u ci ng a c olla p s e o f th e c a v i ty s
’
s id es ; a n d if i ts s id es a re no t r e a dily c oll a p s i bl e , t h e re m u s t b e a c o r
res p o ndi ng res i s t a nc e t o t h e a bs t ra cti o n o f liqu id fro m i t C l ea rly

.
th e lik e m u s t h a pp e n i f th e liqu id i s t o b e d ra w n ou t of a ti s s u e
w hi ch c a nnot e i th e r di m i n i s h i n bu lk b odily o r a llo w i t s com p o ne nts
i ndi v idu ally t o di m i ni sh i n bu lk I n a n ordi nary l eaf, th e u pp er
.
l ay e r of p arenchym a, form e d as i t i s of clos ely p a ck e d c ells th at a re

-
w i th ou t i nt ers p ac es , a nd a re e v e ry w h ere h eld fa s t w i thi n th e i r fram e

w o rk of v e i ns , c a n ne i th er c o ntra c t ea s ily as a m as s , n or a ll o w i t s s ep
a ra t e c ells to d o s o ! ui te o therw i se i s i t w i th th e n etw ork p aren
.
-
c h ym a b el o w T he l o ng c ells of t hi s , u ni te d m erely by t h e i r e nd s
“
a nd h a vi ng th ei r fl e x i bl e s id es s u rro u nd e d by a i r, m a y s e v era lly h a v e
th ei r c onte nts cons id e ra bly i nc rea s e d a nd d ecreas e d w i th out offe ri ng

5 92 A P P E ND I X 0 .
a pp re ci a bl e res i s t a n ces ; a n d th e n et w ork ti s s ue w h i ch th e y fo rm w ill -

,
a t th e s a m e t i m e be c a p a bl e of u nd e rg o i ng s li g h t e xpa ns i o ns a nd c o n
,
t ra c t i o ns of i ts t hi c kn es s I n t hi s l a y er o cc u r t h es e o rg a ns tha t a re s o
.
o b v i ou s ly fi tt e d for a bs o rp ti on H ere w e fi nd th em i n di re ct co m m u
.
ni ca t i o n wi th i ts s y s tem of c oll a p s ibl e c e lls Th e p ro ba bili ty a pp ea rs .
t o be th a t w h e n t he c u rr ent s ets i nto th e l e a f i t p as s e s th rou g h th e

, ,
v ess els a nd th e i r s h ea ths ch i efly i nto th e upp e r l ay er of c ell s (thi s

u pp er l a y e r h a v i ng a l a rg er s u rfa c e o f c o nta c t w i th th e v e i ns t ha n th e
lo wer lay e r a nd b ei ng t h e s ea t of m ore a cti v e proc es s es ) ; a nd that

,
t h e j u i c es o f th e u pp er l ay er enri c h e d by t h e as s i m il a te d m a t ters
, ,
p as s i nt o t he net work p a renchym a w h i ch s e rves as a res e rvoi r fro m

-
,
w h i c h th ey a re from ti m e t o t i m e d ra w n fo r th e nu t ri ti o n of t h e re st
o f t he pl a nt w h e n th e a ct i o ns d e t erm i ne th e d o w n wa rd c u rre nt
, .
Sh ould i t be a s k e d w h a t h a pp e ns w here the a b s orb ent s i ns t ea d of ,
b e i ng i ns e rt e d i n a netwo rk p a renc hym a a re as i n th e l ea ves of

-
, ,
E up hor bi a nerzzfol i a i ns e rt e d i n a s olid p are nch y m a t h e re ply i s

' '
, , ,
th a t s u c h a p a rench y m a th ou gh not fu rni s h e d w i th s y s t em a ti c ally

,
a rra ng ed a i r c h am b e rs n e v e rt h e l ess c o nt a i ns a i r i n i ts i nte r c ell ul ar

-
,
s p a ces ; a nd t ha t w h e n th e re occ u rs a d ra u g ht u p on i t s c ont ents ,
th e e xp a ns i o n o f th i s a i r a nd th e ent ra nce of m ore from w i th ou t ,
qui ckly s u pply th e pl a ce of th e a bst ra ct e d liqu id .
If th e n retu rni ng t o th e g e ne ra l a rgu m e nt w e concl u d e th a t

, ,
th es e e x pa nd e d t e rm i na ti ons of th e v a s c ul a r sy s t em i n l ea v e s a re a h
s o rbe nt o rg a ns w e fi nd a fu rt h e r c o nfi rm a ti o n o f th e vi e w s s et fo rth
,
res e c ti ng th e a l t erna t i n m o v e m e nt of th e s a l ng t h e s a m e c h a n
p g p a o
n el s. Th es e s po ngi ol es of t h e l ea v es lik e th e s p o ng i ol e s of th e roots
, ,
be i ng a ppli a nces by w hi ch liq u id i s t a k e n u p t o be c a rri e d i nto t he

m as s o f t h e pl a nt w e a re o blig e d t o re g a rd t h e v ess el s t ha t e nd i n
,
th es e s pongi ol e s of th e l ea v es as b ei ng t he ch a nnel s o f t he d o w n
c u rre nt wh e ne ve r i t i s p ro d u ce d I f t h e el a bor a t e d s ap i s a bs tra c te d
.
from th e l e a v es by th es e a b s orb e nts t h en w e h a v e no a l te rna ti v e ,
b ut t o s upp os e th a t h a v i ng e nte re d th e v as cul a r s y s t em th e e l a b

, ,
o ra t e d s a d e s ce nd s th rou h i t A nd s e ei ng h ow by th e h el p of
p g .
,
t h e s e s p e c i al t erm i na ti ons i t b ec om e s p os s i bl e fo r the s a m e v e s s el s

,
t o c a rry b a c k a qu a li ty o f s a p u nlik e th a t w h i ch t he y b ri ng u p w e ,
a re e na bl e d t o u n d e rs t a nd t ol e ra bly w ell h o w th i s rhy th m i ca l
m o v em ent p rod u c es a d o w nw a r d t ra ns fe r o f m a te ri a l s fo r g row t h .
T h e s e v e ra l li nes o f a rgu m ent m ay no w be b roug h t t og eth e r ;

a n d a l o ng w i th th e m m a
y be w o v e n up s u c h e vi d e nc e s as re m a i n .
L e t m e firs t p oi nt o ut t h e v ari ety o f q u e s ti o ns t o whi ch t he

hyp oth es i s s up pli es a ns w ers .
I t i s req ui re d t o a c c o u nt for th e as cent of s a p to a h eig ht be y on d

th at t o wh i ch c a pill a ry a c t i on c a n ra i s e i t Thi s as c e nt i s a c cou nte d .
fo r by t h e p rop u ls i v e a c ti on o f t ra ns v ers e s t ra i ns j o i ned w i th t h a t of ,
o s m ot i c di s te nti on A c a u se h as to be as s ig ne d for th at ri s e of s a p
.
CI RCU L A TI ON A ND F ORMA TI ON OF WOOD I N P L A NTS .
5 93
w hi c h, i n th e s p ri ng , wh il e yet th ere i s no c ons id era bl e e va p orati on

t o a i d i t, g o es o n w i th a p o w er w h i ch c a pill a ri ty d oes n ot e x pl a i n .
T h e c o-o p era ti on of th e s am e t w o ag e nc i es i s ass igna bl e for th i s re s ul t

*
als o . Th e c i rc u m st a nc e that v ess el s and d ucts h ere cont a i n s a p a nd
t h er e c onta i n a i r, a nd a t th e s am e pl a c e c onta i n a t diffe rent s eas ons
n e w a i r a nd no w s a p i s a fa ct c a lli ng fo r e x pla na t i o n A n e x pl a na .
t i on i s furni s h e d by th es e m e ch a ni c al a ct i o ns w hi ch i nv ol ve t he e u
t ra nce or e x p u ls i on of a i r a cc ordi ng t o t h e s upply of liqu id That .
v ess els a nd d u cts wh i ch w ere o rigi na lly a cti ve s ap ca rri ers g o c o m -
l e tel y o ut o f us e, a nd h a v e t h ei r fu nc t i on di s charg ed b
p y oth er
v ess els or du cts , i s a n a nom a ly th at h as t o be s ol v e d A g a i n, w e .
a re s uppli e d w i th a s ol ut i o n : th es e d es ert e d v e ss el s a n d d uc ts are
t h os e wh i ch , by th e form i
a t on of d e ns e ti ssu e ou ts id e of th em ,
b ec om e s o c i rcum s t a nce d that th ey c a nnot be c om press ed a s th ey

o rigi na lly w e r e .A ch a nne l h as t o be fo u nd for th e d o w nwa rd
c u rrent o f s a p wh i ch o n a ny ot her hyp ot he s i s th a n th e foreg oi ng
, , ,
m u s t be a c h a nne l s ep a ra t e fro m t h a t ta k e n by th e u p w a rd
c u rr ent ; a nd ye t no g oo d e v id e n c e of a s e p ara t e ch a nnel h a s b e e n
p o i nte d out H ere h ow e ver t he difficulty dis app ea rs s i nce o ne
.
, , ,
c h a nnel s u fli ce s fo r t h e c u rrent a l terna ti ng u p w a rd s a nd d o w nw a rds
a c c ordi ng t o th e c ondi t i ons More o v er th e re has t o be fou nd a

.
fo rc e p ro d u ci ng or fa c ilit a ti ng th e d o wnw a r d c urr ent c ap a bl e e v e n ,
o f d ra w i ng s a p o u t o f d ro opi ng b ra nc h e s ; a nd n o such for ce i s
fort h co m i ng T h e hyp othes i s s et fort h di s p e ns es w i th thi s nec e ss i ty ;

.
u n d er t h e re cu rri ng c h a ng e o f c ondi ti ons th e s a m e d i s t e nt i o n a nd

,
o s c il l a ti on wh i ch be fore ra i se d th e s a p t o t h e pl ac es of co ns u m p t i o n ,
no w b ri ng i t d o w n t o th e pl ac es o f c on s u m p ti on A ph y s i c a l .
p ro ces s h as to b e p oi nt e d ou t by w hi ch th e m ateri a l th at form s

d ens e t i ssu e i s d ep os i te d a t th e pl a ce s wh ere i t i s w a nte d ra th er ,
t h a n a t o th er pl a c e s Thi s ph y s i ca l p roc ess t h e hyp oth es i s i n

.
d i c a te s. I t i s r e qu i s it e t o fi nd a n e x pl a na t i on of th e fa ct th at ,
w h e n pl a nt s ordi na rily s w a y e d a b ou t by t h e w i nd a re g row n i nd oors ,
th e for m a t i on of w o o d i s so m u ch di m i ni sh e d th a t t h ey b e com e a h
norm a lly s l e nd er Of thi s a n e x pl a na ti on i s s u ppli ed Ye t a furth e r
. .
It s eem s prob b l e h owe v e r t h a t o s m oti c di s te nti on i s h ere e s pe ci a l l y

a .
, , ,
th e m ore i m port a nt o f th e t wo fa c tors The ri s i ng of t h e s a p i n s p ri ng m a y

.
i ndi re ctl y res u l t, l i k e t h e s p rou ting of th e s ee d, from t h e t ra n s form a ti on o f

s t a rc h i nto s u ga r . D uri ng g erm i n at i on, t hi s ch a n g e of a n oxy hyd ro-ca rb on -
from a n i nsol ubl e i nt o a so l u b l e form , l e a ds to ra pi d e ndos m os e ; con

s eq u e ntl y t o g re a t d i s t ent i o n o f th e s ee d ; an d th er efore to a for ce wh i ch
th rus t s t h e c on t a i ne d l i q u i ds i nto t he p l um ul e a nd r a di cl e , a nd gi v e s t he m
p o we r t o di s pl a ce t h e s oi l i n t he i r w a y : i t s e ts up a n a ct i v e i nte rn a l m ov e
m ent wh en ne i th er e v a p ora t i on n or th e c h ang e w hi ch l igh t p rod u c es ca n be
o p era t i v e . A nd s i m i l a rl y, i f i n th e s p ri ng , th e s t a rch s to re d u p i n t h e roots
,
o f a t r ee p a s s e s i nt o t he fo rm o f s u g a r, t h e unu s u al o s m o ti c a b s orp t i on t h a t
a r i se s w il l ca u s e an u nu s ua l di s t enti on
—a di s te nt i on w hi ch , b ei ng re s i s te d by
t h e t ou gh b a rk of t h e root s and s tem , will res ul t i n a powerful u p ward th rust

o f th e conta i ned l i q ui d .
84
5 94 A P P E ND IX 0
.
fa ct t o be i nterp ret ed i s th at i n the s am e i ndivi dual pla nt h om ol

,
o ou s p a rt s w h i ch ac c or di ng t o th e typ e o f th e pl a nt s h o u ld b e
g , , ,
e q u a l ly w o o dy b e com e m uch th i ck e r o ne th a n a not h er if s u bj ec t

,
t o g re a t er m ech a ni c a l s tress A nd of th is t oo a n i nte rp reta ti o n i s

.
s i m il a rly a fford e d .
Now th e s u ffi c i e ncy of th e as s ig ne d a cti ons to a cc ou nt for s o m a ny

p h enom ena not oth e rwi s e e x pla i ne d w ou ld b e s trong evid enc e th at ,
t h e ra t i o n al e i s t h e t ru e o ne e v e n w e re i t of a p u rely hyp oth eti c al

,
ki nd H ow s t rong th en b e com es the reas on for b eli evi ng i t th e

.
, ,
t ru e o ne w h en w e rem em b e r th a t t h e a c ti ons all eg e d d e m o ns trably

o o n i n th e w a a s s e rt e d T h ey are e v e r op era t i ng b efore ou r
g y .
e y es ; a nd th a t t h ey p ro d u ce th e e f fects i n qu es ti on i s a c onclu s i on
d ed u cible from m e ch a ni cal p ri ncipl es a c onclus i on est a bli shed by ,
i ndu cti on a nd a concl us i on v eri fie d by e x p eri m ent Thes e th ree

,
.
o rd ers o f p roof m a y be b ri efly s um m e d u p as foll ow s .
Th a t pl a nts w h i ch h a v e to ra i s e t h em s el v e s a bove th e e arth s s ur

’
fa c e a nd t o wi ths ta nd t h e a c t i ons of th e w i nd m us t h a v e a p ow er of
, ,
d e v el opi ng s upp ort i ng s tru cture i s a n ap r i or i concl us i on w hi ch m ay

,
b e s a fely d ra w n It i s a n e qually s afe a p r i or i c onclu s i on th a t i f

.
,
th e s u pp orti ng s tru ctu re e i th er as a wh ole o r i n a ny of i ts p a rts h as

, ,
t o a d a p t i ts elf to t h e p a rt i cul ar s t ra i ns wh i ch th e i ndividu al pl a nt

i s s u bj e ct t o by i ts p a rti cul ar c i rcum s t a nc e s th ere m u s t be a t w ork ,
s o m e p ro c es s b w h i ch th e s t re ngt h of th e s upp orti ng s t ru ctu re i s

y
e v e ry wh e r e b rou ght i nt o e qu ilib ri u m wi th t h e forc es i t h as t o b ea r .
T hough t he typi c a l dis tri buti on of supp ort i ng s tru cture i n e a ch ki nd

o f pl a nt m a
y be e x pl a i ne d tel e ol ogi c ally by thos e w h om t el eologi c al
e x pl a na ti o ns s a t i s fy a nd th ough oth er wis e thi s typi c al di s trib uti on
m ay be as crib e d t o na tu ral s el ec ti o n a cti ng a p a rt from an di rectly

y
a d a p t i v e p roces s ;
y et i t i s m a ni fes t tha t t h os e d ep a rtures from th e
typi ca l di s tri buti on w h i c h fi t th e p a rt s of ea ch pl a nt t o th ei r sp eci a l
c o ndi ti o ns a re e x pli ca bl e ne i th e r t el eol ogi cally n or by n a tu ra l s e l e c
t i on . W e a re th erefore c om p elled to a d m i t that if i n e a ch pl ant

, , ,
th ere goes o n a b al a nci ng of th e p a rt i cul a r s t ra i ns by th e p a rt i c u l a r

s t re n t h s th ere m u s t be a ph y s i c al or phy s i c o ch em i c al p roce s s by
g
-
,
w hi c h the a dj us t m ents of th e t wo a re effec te d Mea nwhil e w e a re .
e q u a lly c o m p e ll e d t o a d m i t a p r i or i th a t th e m ec h a ni c al a c t i ons to
‘
, ,
be res i s t e d t hem s elv es a ffec t th e i nt ernal ti ss u es i n s uch w a y s as to

,
furth e r th e i ncrea s e of t ha t d ens e s u b s ta nc e by wh i ch th e y a re re

s i s ted I t i s d em ons t ra bl e th at b e ndi ng th e p eti ol es s h o ots a nd
.
, ,
s tem s m u s t c o m p re s s th e v es se l s b enea th t h e i r s u rfa ces a nd i nc r eas e

,
t h e e x u d a t i o n of nut ri ti v e m a tte rs from th em a nd m u s t d o thi s a c t ,
i vel y i n p rop orti on as th e b end s a re gre at a nd fre qu ent ; s o t h a t

whil e o n th e one h a nd i t i s a neces s ary d e d u cti on th a t i f th e p a rt s of
, , ,
e ac h
pl a nt a re t o be s e v erally s treng th e ne d a c cordi ng to th e s e v e ra l
s t ra i ns th ere m u s t b e s om e di rec t c onne x i o n b e t w e e n s t ra i ns a nd
,
streng th s i t i s on t h e o th er h a nd a ne c ess a ry d ed ucti on from

, , ,
CIRCU L A T ION A ND F ORMA T ION OF WOOD I N P L A NT S .
5 95
m e ch an ca i l princ ipl es th at th e s tra nsi do ac t in su ch w a ys as to ai d

th e i ncreas e of th e s tre ngths H o w a lik e c orres p ond enc e b et we en
.
t w o a p ri or i a rg um ents h ol ds i n t h e c as e of th e c i rc ul at i on nee d s ,
not to b e s ho wn i n d et a il . It w ill s u ffi c e t o re m i nd th e rea d er

t h a t w h il e t h e ra i s i ng of s a p to h eights b ey o nd th e li m i t of ca p
i l l a ri ty i m pli es s om e forc e to e ffec t i t w e h a v e i n th e os m oti c ,
di s te nti on a nd th e i nterm itt ent c om p re s s i ons c aus e d by t ra ns vers e

s t ra i ns fo rces w h i ch u nd er th e c o ndi ti ons c a nnot bu t t end t o
, , ,
ef fe ct i t ; a nd s i m i larly wi th th e r e qu i rem ent for a d ow nw a rd

c urre nt a nd th e p ro d u ct i on of a d o w nw a rd cur re nt
, .
A m o ng th e i nd uc tive p roofs w e find a ki nd re d ag ree m ent D i fi e .
re nt i ndi vidu als o f t h e s a m e s p e c i es a n d d if ,

fere nt p a rts o f th e s a m e
i ndi v idua l d o s trength en i n d i fferent d egre es ; and t here i s a clea rly
,
t ra c e a bl e conne xi on b et ween t he i r st rength e ni ngs a nd th e i nt e rm i t

te nt s tra i ns th ey a re e x p os e d t o Thi s e v id ence d eri ve d from cont ras ts
.
,
b et we en g rowt hs on t he s am e pl a nt o r on pla nts o f th e s a m e typ e i s ,
e nforc e d by e vid e nce d eri v e d from c o nt ras ts b et w e e n pl a nts of d i f fe

re nt t yp e s T h e d efic i e nc y o f w oody t i s s ue w hi ch w e s e e i n pl a nts
.
c a ll e d s u cc ul e nt i s a cco m p a ni e d by a b ulki ne ss o f th e p a rt s wh i ch
,
pre v ents a ny c ons id era ble os c illati ons ; a nd t hi s ch a ra cter i s a ls o

h a bi tu ally accom p a ni e d by a d w a rfed grow th W he n lea ving th es e
.
,
re l a t i o ns as di s pl a y e d e x t ern a lly w e e x am i ne t h em i nt erna l ly w e

, ,
fi nd th e fa cts u ni ti ng t o s h o w by th e i r a greem e nts a nd di fferenc es

, ,
th a t b etw e en th e com p re ss i on of th e s a p c a na ls a nd th e p rod u c ti o n

-
o f w o o d t h e re i s a di re ct rel a ti on W e h a v e t he facts t ha t i n ea ch
. ,
pla nt a nd i n e v ery ne w p a rt of e a ch pla nt th e form a ti o n of s a p

, ,
c a na l s p r ec e d e s th e fo rm a ti o n o f w oo d ; th a t t h e d e p os i t o f w oo d y
m a tt er w h en i t b e g i ns t a k es pl a c e a ro u n d th es e s ap ca n a ls a nd
-
, ,
,
a ft erw a r ds a rou n d th e n e w s a p c a n a l s s uc c e s s i v ely d e v e l o p e d ; th at

-
h i f t i f w d n d t h c a n a l s t a k es pl a c e w h e re t he
t s a r o u e s a
p
-
or m a o n o o o
a n d th a t t he a m o unt
c oa ts of th e c a n al s a re d em ons tra bly p erm ea bl e ,
o f w o o d fo r m a ti o n i s p rop o rti ona te t o t h e p e

- rm e a bili t y A nd t h en .
t h at t h e p e r m e a bili t y a n d e x t r a v as a t i o n o f s a
p o c c u r w h ere ver i n ,
th e n i di v id u a l o r i n t h e t yp e ,
t h e re a re i n t e r m i tt e n t c o m p res s io ns ,
i s p ro v e d alik e by ordi na ry cas es a nd by e xc eptl o na l c as es In .
l f w t h t t h d e p os i t o f w o o d r oun d th e
h
t e o n e c as s o c as e s e s e e a e
v es s els be gi ns to t ak e pl a c e wh en th ey c om e i nto p os iti ons th a t

h
j
s u b e ct t h e m t o i n t e rm i tt e nt c o m p r e s s i on s w
l
l e 1 t c e as e s w h e n
,
A n d 1 n th e oth er c l as s
t h ey b e c om e sh i eld e d from com p re s s i ons .
o f c as es w h e r e from th e b eg i nn i ng th e v es ,
s e ls a r e s h i e ld e d fr om
, ,
c om p ress i o n b y s u rr o u n di n g fl e s h y t i ss ue t h e r e i s a p e rm a nent
,
a bs en c e of w o o d form a ti on
- .
T o w hi ch c om pl et e a gr e e m ent b et w ee n th e d e d u cti v e a nd 1 ndu c

ti v e i nfe re nc e s h as t o b e a dd e d th e di rect p roo f s u ppli e d by e xp e ri
m ents . I t i s pu t b ey on d d ou bt by e x p erim e nt th a t th e liq u i ds a h
a rt s th rou gh th ei r
s orb e d by pl ants a re d i st rib u t e d to th e i r diffe re t p
n
5 96 A PP E N D I X c .
v ess els— at firs t by th e s pi ral or alli e d v ess els o ri g i nall y d ev el op e d,

a nd t h en by th e b ette r pla ce d d u cts fo rm e d l a t e r B y exp e ri m ent
-
.
i t i s d e m o ns tra te d th a t th e i nt erm i tte nt c om p r ess i ons c a u s e d by o s

c il l a t i o ns u rg e t h e s a a l o ng th e v ess el s a nd d u cts A n d i t i s als o e x
p .
er i m e nt all y p ro ve d t h a t t h e s a m e i nte rm i t te nt c o m pres s i o ns pro du c e

p
e x u d a ti o n o f s a
p from v es s el s a nd d u c ts i nto th e s u rroundi ng t i s s u e .
Th a t th e p roce ss e s h e re d es crib e d , a ct i ng th rou g h a ll p as t t im e,

h a v e s uffi ce d of th em s el v es t o d ev el op e th e s u pp ort i ng a nd d i st ri but .
i ng s tru ctu res o f pl a nts , i s no t al l e ge d Wh at s h a re th e n atural

.
s e l e ct i o n of v a ri a ti o ns d i s ti n u i s h e d as s onta n eou s , h as h a d i n e s ta b
g p
l i shi ng th e m , i s a qu es ti o n w hi ch rem a i ns to be di s cu s s ed W h e th er .
ac ti ng a l on e n atu ra l s e l e c ti on wou ld h a ve s u fli c e d t o e vo l v e t h es e
v as cu la r a nd r es i s ti ng ti ss u e s , I d o not p rofes s to s ay T ha t i t h as .
b ee n a c o op erati ng ca use , I t ak e to be s elf e v id ent : i t m us t al l al ong

- -
h a ve furt h e red th e a ct i on o f a ny o t h er ca us e , by p re s e rvi ng th e i n

d i vi du al s on w hi ch s u ch oth e r cau s e h a d a ct e d m os t fa v ou ra bly .
S e ei ng, h o w e v er, th e c onclus i ve p ro of wh i ch w e h a v e th a t a noth er

c a u se h as b e e n i n a c ti on— c e rt a i nly o n i ndi vidu al s , a n d , i n al l p ro ba
bi l i ty, by i nh eri t anc e o n ra c es — w e m ay m os t ph il os oph i cally as crib e

th e g en es i s of th es e i nt erna l s tru ctur es t o thi s ca us e , a nd reg a r d
n a tu ra l s e l e c ti on as ha vi ng h e re pl a y e d th e p a rt o f a n acc el era t or .
E XP L ANA TION OF P L A TE .
Fig 1 A b s orb ent organ from th e l e af of E up hor bia ner i zfol i a

. . .
Th e clus ter of fib rou s c ells form i ng o ne of th e t erm i nati ons of th e

v as cul ar sy s te m i s h ere i m b e dd ed i n a s olid p arenchy m a .
Fig 2 A s tru ctu re of a nal o g ous ki nd from the l eaf of F i cu s

. .
el a s ti ca H ere th e exp a nd e d t e rm i na ti o ns of th e v es s el s a re i m
.
b e dd e d i n the net work p a rench ym a the cells of w hi ch u ni te t o ,
form e nv el op e s for th em .
Fig 3 S ho ws o n a l arg er s cal e one of t hes e a b sorb ents from

. .
th e lea f o f P a na x L es som i I n t hi s figu re is cl ea rly s een th e w ay

’
i n w hi ch t he c ell s of th e ne twork p a re nch ym a u ni te i nto a cl os ely

fitti n cas e for th e s pi ral c ell s
g .
Fig 4 Repres ents a m uch m ore m ass i v e a b s orb e nt from the

. .
s am e l e af th e s urrou ndi ng t i s s u es be i ng o m i tte d

, .
Fig 5 S i m il arly rep re s ents w i th ou t i ts s h eath a n a b s orb ent

. .
, ,
from th e l eaf of Cl u si a fl a va .
Fig 6 E nd vi ew o f a n a b s orb ent orga n from th e root o f a

. .
T u rnip I t i s ta k en from th e o ut erm os t l a y e r o f v es s el s

. I ts .
funne l s h ap e d i nt eri or i s d ra w n as i t p res ents i t s el f w hen l ook e d

-
a t from t he ou ts id e o f th i s l a y er i ts n arro w e nd b ei ng di rect e d

,
t o wa rd s th e ce nt re of t h e T urnip .
Fig 7 A l ongi tudi nal s ecti on th rough th e a xi s of a noth er s uch

. .
o rg a n s h o w i ng i ts a nnuli o f re ti c u l a te d c el l s wh en c u t th rou gh
, .
T h e cell ul ar ti s s u e wh i ch fil ls th e i nte ri or is supp os e d to be rem ov e d .

CIRC U L A TI ON A ND F ORMA T ION OF WOOD I N P L A NTS .
597
Fig 8 A les s d e v elop e d a b s orb ent s h ow i ng i ts a pp ro x i m ate

. .
-
,
c o nne xi on w i th a d u c t. In the i r s i m pl es t form s th es e st ruct u res

,
c o ns i st o f o nly t w o fe n es t ra ted c ell s wi th t h e i r e nd s b e nt rou nd

,
s o as t o m e e t S u ch typ e s o c c u r i n the ce ntral m as s of th e T urni p

.
,
w h ere th e vas cu l ar s y s tem i s rel ativ ely i m p erfect . B e sid es th e

com p ara ti v el y regul a r form s of th es e a b s o rb en ts , th ere a re fo rm s
com p o s e d o f a m o rph ou s m a ss es o f fenes t ra t e d c ells .I t sho uld

be a d d ed b oth th e regul ar a nd i rreg ul ar k i nd s are v ery vari
tha t
a bl e i n th e i r nu m b ers : i n s om e t urni p s th ey a re a b u nd a nt a nd i n
,
oth ers s c a rcely t o be fo und . P oss ibly th ei r p rese nc e d ep ends on

598 A PP E N DIX 0
.
th e a
ge of th e T u rni p .Judgi ng from th e p eri od du ri ng whi ch
m i nv e s t ig a t i ons w e re m a d e n a m e ly w i nte r a nd e a rly s p ri ng I
y , ,
sus p e ct th a t th ey are d e v el op e d o nly i n p re p a ra ti o n for s endi ng
u t h e fl o w er s t alk
p
-
.
5
L e t m e a dd th a t e xp erI m e nts on c i rc u l at i on i n pl a nts m a d e

d uri ng th e s tat e of i na cti v i ty wh en i t i s t o be p res u m e d tha t th e
,
v es s els a nd t i ss u es c ont a i n but li ttl e sa p a re m uch m ore s u c

,
c es s ful th a n th os e m a d e i n t h e s um m e r . I t w ou ld s e em tha t
w h e n th e t i s s u es a r e fully ch a rg e d wi th s a p th e t a ki ng up of
dy e s i s com p a ra ti v ely s l ow a nd the a b ove d es cri be d effe cts a re
-
n ot s o e a s ily d e m o ns t ra bl e .
[ A n e xp ert w ri tes i g th i s e s s ay
concern n I h a v e not
a tt e m p t e d t o a nnot a t e c ri ti c a lly th i s p a p e r T here i s n o d oubt
.
th a t m a ny of y ou r c o ncl us i o ns a re p e rfe c tly s ou nd p a rt i c u l arly

,
t ho s e rel a t i ng t o t he p as sa g e of c ru d e s ap th rough t he ca vi ti es of
th e el em ents of th e w o od th ou gh th e opi ni on th a t t h e a ct ua l
“
p a ss age w a s th rough th e w alls very g enera lly h eld t i ll a b out 1 2

y ea rs
A P P E ND IX D .
ON THE ORI GIN OF THE VE RTE BRATE TYP E .
[ When s tud yi ng the devel op m en t of the ver tebr a te s kel eton, there
occu rr ed to m e the fol l owi ng idea r espec ti ng the p ossi bl e or ig i n f
o the
no tochor d . I wa s l ed to om i t the few p a g es of A p p endi x i n
even tu a l l y
whi ch I ha d exp res sed thi s id ea , beca u se i t was u nsup por ted by devel op
m en ta l evid ence . The d evel op m en ta l evid ence recen tl y di scover ed , how
ever , ha s l ed P r ofessor H d eckel a nd othe r s to a na l og ou s vi ews resp ec ti ng
the afi l i a ti on of the Vert e b r a t a on the Mollu s c oida H a vi ng for ta

.
na ts l y pr eser ved a p r oof of the supp res sed pa g es , I a m a bl e n ow to

a d d them . Wi th the om i ssi on of a s up erfl u ou s p a ra g rap h, they a re
rep r i n ted ver ba ti m fr om thi s pr oof, whi ch d a tes ba ck to the a u tu m n
of 1 8 6 5 , a t whi c h ti m e the chap ter on T he S hap es of Ver tebr a te

S kel etons was wri tten — D ecem b er,
The genera l argum ent conta i ne d i n C h a p X VI of P a rt I V , I

. . .
h a ve th ou ght i t u nd es i ra bl e to i m pli c a te w i th a ny c on cep t i on m ore

s p e cul a t i v e th a n th os e ess enti al t o i t ; a nd t o a vo i d s o i m pli ca ti ng
i t, I tra ns fer t o t hi s pl a c e a n hyp oth es i s resp ect i ng th e d eri v a ti o n

-
o f th e rudi m e nt a ry v ert eb ra t e s t ru c tu re , w h i ch a pp e ars t o m e
w orth c ons id e ri ng .
A m ong t h os e m ollus co id a ni m al s w i th wh i ch th e l ow e s t v e rt e
b ra t e a ni m al has s u ndry t ra i ts i n c om m on, i t v ery g enerally h app e ns
th a t w hile t h e a d u l t i s s t a ti ona ry t h e l a r v a i s lo c om oti v e Th e .
l oc om oti on o f th e l a r v a i s e fi e ct e d by the u ndu l a ti ons of a ta il I n .
s h a p e a nd m o v e m e nt o n e o f t h es e y ou ng A s c idi a ns i s no t a l t og eth e r
u nlik e a T a dp o l e . A n d a s t he t a il of th e T a dp ol e d i sa pp e a rs
w h en i ts fu nc t i o n c om es t o be fulfil l e d by lim b s ; s o t h e A s c idi a n
la rv a s t ail di sapp ears w h en fi x ati on of th e la r v a re nders i t u s eles s
’
.
T h i s di s app e a ra nc e of th e ta il h ow e v er, i s not w i th ou t e x c ep ti on

, .
Th e A pp endi c u l a r i a i s a n A s c idi a n w h i ch reta i ns i ts t ail thr ough

o u t li fe ; a nd by i ts a i d c o nt i nu e s th r ou gh ou t li fe t o s w i m a b o u t .
Ne w thi s t ail of th e A pp endic u l a ri a h a s a v e ry s u gg es ti v e s t ru ctur e .
I t i s l ong, t ap eri ng t o a p oi nt, a nd fl att ene d From e nd t o end

.
th ere ru ns a m i d-ri b, whi ch a pp ears t o be a n im b e dd e d g el a ti nous

rod, not unlik e a notoch ord . E xtendi ng al ong th e t wo s id es of
5 99
6 00 A P P E ND I X D .
th i s m i d -
ri b, a re b u ndl es of m u s cul a r fibres ; a nd i ts t op b ears a
ga ngli ate d ne rvous th rea d gi vi ng off a t i nt e rv als branch es to the
, , ,
m u s cul a r fibres .I n t he A pp endi c u l a r i a t hi s t a il w hi ch i s i nse rt e d ,
a t th e l o w er p a rt o f th e b a c k i s b e nt fo rw a rds s o as not t o be
, ,
a d a p t e d for p rop elli ng th e b o dy of th e a ni m a l h ea d fore m o s t ; but
th e ho m ol og ou s t a il s of th e l a r v a l A s cidi a ns a re di re c t e d ba ck w ards ,
s o a s t o p ro du c e for wa rd m o v e m e nt If w e s u pp os e a typ e lik e the

.
A pp endic u l a ri a i n th e s t ru ctu re a nd i ns e rti on of i ts p erm a nent ta il ,
but re s em bli ng th e l a rv a l form s i n th e di rect i on of i ts t a il i t i s I , ,
th i nk n ot di ffi c ult t o s ee th a t fu n ct i ona l a d ap ta ti on j o i ned w i th

,
na tu ra l s el ec t i on m ight rea dily p ro d u ce a typ e a pp r o x i m a ti ng t o

,
th at w h o se origi n w e a re cons id e ri ng I t i s a fa i r as s u m p t i on.
tha t a n h a bi tu ally l oc om ot i v e creature w ould p rofit by i n

c re as e d p o we r o f l o c om oti o n Thi s gra nt e d i t foll o w s th a t
.
,
s u ch furt h er d e v e l op m ent of th e ta il s t ru c tu re s a s m ig ht ari s e -
from e nha nc e d fu nct i on a nd s u c h better di s tri bu ti on of th em

,
as s p o nt ane ou s v ari a t i on m igh t fro m ti m e t o t i m e i ni t i a te ,
w oul d be p e rp etu at e d Wh a t m u st be the a cc om p a nyi ng ch anges ?

.
Th e m ore vig orous a ct i on of s uch a n a pp end ag e i m pli e s a fi rm er

i ns ert i on i nt o th e b ody ; a nd thi s w ould be e ffecte d by the pro
l on ati on forwa rds of th e c entra l a x i s of th e ta il i nto th e c rea tu re s ’
ba ck A s fas t a s th ere p rogr es s e d th i s fus i on of th e i ncrea s i ng ly

p owerful t a il w i th th e b ody th e b o dy would b egi n t o p a rt ak e of i ts
,
o s c ill a ti o ns ; a nd a t th e s a m e t i m e t h a t th e re s i s ta nt a x i s o f th e t a il
a d v a n c e d a l ong th e d ors a l re i o n i ts a c c o m p a nyi ng m u scu l ar fib re s

g ,
w ould s p rea d o v er th e s id es of th e b o dy : g ra du ally t ak i ng s u ch

m o difie d di recti ons a nd i ns ert i o ns as th ei r ne w c ondi t i ons re nd e re d
m os t a d v a nta g eou s Wi thout fu rth er explanat i on those w ho
.
,
e x a m i ne d ra w i ngs o f th e s tru ct ures d es c rib ed w il l I thi nk , , ,
s e e that i n s u ch a w a a t a il h om ol og ou s w i th t hat of the

y
A pp endi cu l a r i a w ould be lik ely i n the c ours e of that de
, ,
vel O m ent re qu i re d for i ts g rea ter e ffi ci ency g ra du ally to

p ,
e ncro ac h o n t h e b ody until i ts m i d ri b b ecam e th e d ors a l

,
-
a xi s,
i ts g a ngli a te d n erv e t hre ad th e s pi na l c h ord a nd i t s
-
,
m u s c ul a r fi bre s th e m y o co m m a ta S u ch a d e v el op m e nt of a n
.
a pp e nd a g e i nt o a d om i na nt p a rt of t h e o rg a ni s m th ou gh a t fi rs t ,
s ig ht a s t artli ng s upp o s i t i on i s not w i th o ut pl e nty o f p a ra ll els

,
i ns t a nc e the w ay i n whi ch the ce re b ra l g angli a origi nally m ere ,
a dj u nc ts of th e s pi nal ch ord e ventu al ly b ec o m e t he g re a t ce ntres of

,
th e ne rvous s y s te m to w h i ch th e s pi na l c h ord I s qu i t e s u b ordi na te

or i ns t a nc e th e w a i n w h i c h th e li m b s s m a ll a nd i nc ons pi cu ou s i n
y ,
fis hes bec om e i n Man m ass es whi ch t ak en t og eth er ou twe igh th e

, , , , ,
t ru nk . I t m ay be a dd e d th a t th es e fa m ili ar c as e s h a ve a furth er
a pp rop ri a t e ne ss ; fo r t h e y exh ibi t high er d eg rees of th a t s am e
i nc reas i ng d o m i na nc e of t he org ans o f extern al rel ati on whi ch th e ,
hyp oth es is i ts elf i m plies .

ON T HE ORIGIN OF T H E VE RT E BRA T E T Y P E .
6 01
Of c ours e, i f th e rudi m entary v e rt eb ra te app ara tu s th us grew

i nto , a nd s p rea d o ver, a m ollus coid vi s cera l s ys t em , the form ati on
of t he no t o ch ord u nd e r th e a ct i on of a lt erna t i ng t ra ns vers e s tr a i ns ,
did not t ak e pla c e as s u gg es ted i n 2 5 5 bu t i t d oes not th erefore

foll ow th at i ts di ffere nt i a t i on fro m s u rround ing t i ssu es was not
m ech a ni ca lly i ni ti a te d i n th e w a d es c rib e d F or wh a t w as s a i d i n
y .
t ha t s e ct i o n res p ect i ng th e e ffe c ts of l a te ra l b endi ngs o f th e b o dy ,

e qu a l l y a ppli es t o l a t e ra l b endi ngs o f t h e t a il ; a nd as fa s t as t h e
d e velo pi ng t ail encroa c he d on t he b o dy, th e b ody w oul d b e com e

i m pl i ca t e d i n the tra ns ve rs e s tra ins , a nd th e di ffere nt i a ti o n w ould
a d v a nc e forw a rd s u nd er th e i nflu e nc e s o rigi na lly a ll eg e d. O b v i o us ly ,
t oo , t h oug h th e l a t era l m u s cu l a r m as s es w ou ld i n t h i s cas e h a v e a
d ifi ere nt hi s tory ; y et t he s egm e nt a ti on of th e m w ould be e v e nt ually
d et e rm i ne d by th e a ss igne d c a us es For a s fa s t a s th e s t ra ta of
.
c ontra ct il e fib res , d e v el o pi ng s o m e wh a t i n a d v a nc e o f t h e d ors a l
a x is p rea d along the s ides they w ould c om e und er the i nflue nc e

, s ,
o f t h e a lte rna t e fl e xi o ns ; a nd w h il e b
, y s u rvi v al of t h e fi tte s t t he i r
,
p a rts be cam e a dj us te d i n di recti on t hei r s egm ent a ti on w o uld as

, ,
b efore a c co m p a ny the ir i ncreas i ng m ass i ve nes s T he a ctio ns a nd

, .
re a cti ons du e t o l a te ra l u nd u l a ti o ns w o u ld s till th erefore be t h e

, ,
c a u s e s of dif fere nti a ti on wi th wh i ch natura l s el e cti on w o uld c o

,
op e ra te .
A P P E ND IX D 2 .
THE A NN UL OSE TYP E .
THE p rod ucti on o f a s eg m ental s tru cture by undulatory m ov e

m ents s ugg es te d i n A pp endix D as a l s o i n B fi rs t p u bli s h e d i n
, , (
1 8 5 8 ) as e xpl ai ni ng th e v ert e b ra l c ol um n h a s b e en re ce ntly ,
s ugg es t e d by P rof Ko rs ch e l t as t h e c au s e of t ha t s e g m enta ti o n of

.
th e a nnu l os e typ e w hi ch giv es th e na m e t o i t H e es p ou s es .
v ie w wh i ch i s b as ed up on t he a ss um p ti on th at a t first an unsegm ented ,

e l onga t ed a nce s t ra lform wa s p ro duc ed by t e rm i na l growt h, wh e reup on th e
e nti re body beca m e s ep a ra t e d a t onc e i nt o a l a rg e num ber of s egm e nt s by a
re a r ra nge m ent of th e i ndi vi dua l orga ns Thi s a ssum pt i on i s s upp ort ed by

.
t h e cons i de rat i on th a t w i th th e l a tera l s inuous m o v em ent of t h e body, a nd

w i th t h e ri g idi ty o f t he ti s s ue s c au s ed by i ncreas i ng differenti a tion, the
form a t i on of a l te rna ti ng regi ons of grea te r a nd l es s m ot i l i ty was of con
s ide ra bl e a dv a nt ag e t o t he i ndi v idua l , a nd ren dere d p os s ibl e a furt he r el on
ga t i on of th e bo dy .The fi rs t ca u s e for the a pp e a ra nce of m eta m e ri c

s egm ent a ti on woul d then b e s ou gh t i n th e m a nner of l ocom ot i on a nd i n
m echa ni ca l co ndi ti ons Howev e r, thi s l a tte r vi e w i s not s upp orte d i n a ny

.
”
w a y by e m b yo ogy
r l .
( E m bryol ogy of I nver tebr ates , P a rt I, pp 8 4 9 .
I v entu re t o thi k t e co ess i o that t hi s vi e w i s not s up

hn nf n “
”
o rte d i n a ny w ay by e m b ry ol ogy should be j oi ned w i th th e
p
c onfe ss i o n th at i t i s a t vari a nce w i th tha t a b s tra c t em br ol o
il;
y
w hi ch c om preh e nd s th e p ro cess of d e v el o pm ent i n g e nera l T .
a s s u m p ti o n t h a t th ere t ook pl a c e a re a rra ng em e nt o f th e
i ndi v id ual orga ns of a n u ns eg m ent e d e l o ng a t ed a nc e s tra l

,
fo rm , i n s uc h w i s e t ha t t he org a ns , p re v i ously s i ngle , pre s ently

”
b e ca m e m ul tipl e, s o tha t i ns t ea d of o ne o rg a n of each ki nd t here

w ere s u b s ti tute d m a ny org a ns o f ea ch ki nd , i s i nco ns i s te nt w i th
th e g ene ra l l a w of e v ol u ti o n, org a ni c a nd oth er— i m pli e s not
i nteg ra ti on but di s i nteg rati on E very w he re th e a d v a nce i s from
.
m a ny lik e p a rts p e rform i ng lik e func ti ons t o rel a t i v e ly fe w u nlik e

p arts pe rform i ng unl ik e functi ons The high e r fo rm s of the a nnu
.
los e ty pe i ts elf sh o w thi s C om p a re a m y ri ap od a nd a cra b In t he

. .
o ne w e h a v e not only a g rea t nu m b e r o f s i m il a r s eg m ents b eari ng
s i m il a r li m bs , but w e h a v e i n e a c h s e gm ent a dil at a t i o n o f the m a i n

bl ood -vess cl — a r udim enta ry h ea rt— a s woll en p ort i on of th e
nerv e c o rd— a s m a ll g a ngli o n— a nd s o on ; wh e rea s i n th e othe r,
6 02
TH E A NN UL OSE T YP E .
6 03
b es id es rel a ti v ely few s egm ents a nd few li m b s (s undry of th em

e x t re m ely u nlik e th e res t w e h a v e a v a s cul ar s y s t e m con c e ntra t e d
)
i nto a c ent ral h ea rt w i th a rt e ri es a nd a c oncent ra ted ner v ous
s y st em s u c h t ha t th e g re a t ga ngli a i n t h e i nt eg ra t e d c ara p a c e
,
i m m ens ely s ub ordinate the gangli a of th e rem a i ni ng s eg m ents ;

a nd s i m i l a rly w i th the o th er orga ns No w u nless i t b e d eni e d
.
t h a t th e s e h igh es t d ec a p ods h a v e b een e v ol v e d from l o w typ es

ak i n to m ri a p od s i n c om p os i ti o n i t m u s t be a d m i tte d th a t th e
y ,
p rog ress h as be en from a s tri ng of m any lik e s egm ents w ith

s i m ilar s ets o f o rg a ns to a g rou p o f rel at i v ely fe w u nl i k e s egm e nts -
w i th di ss i m il a r s e ts of o rg a ns I f s o w e ca nnot ra ti ona lly d e ny

.
t h at th e p rogr es s h as b ee n of th i s n a t u re u p from th e l ow es t
a nnelid i ns te a d of h a vi ng b e en as P rof Ko rs ch e l t s h yp oth es i s
’
.
, ,
i m pli es of Opp os i te na tu re a t t he b egi nni ng

, .
I n a p rec e di ng p as s a g e a cle a r rec ogni t i on of th e norm a l

co u rs e o f d e v e l op m e nt o c c urs I n Opp os i ng th e vi ew s et fort h i n
. .
2 05 —7 of t hi s w ork P rof Kors ch e l t s ay s

, .
It s eem s sca rcel y fa v oura bl e t o thi s th eory t ha t t h e degree of i nde

“
ndence wh i ch th e i ndi v id a l se g m ents p r ese nt i s co m p a ra t i v el y s l ight

u .
e m ost i m porta nt orga ns (ne v ou s s ys t em r body m us c ul a tu re bl o d asc u

, ,
o -v
l a r s ys te m ) s h ow them sel v es to be si ngl e funda m ent s of th e e nti re b ody ,
a nd a r e a l s o devel op ed a s s u ch e v en t hou gh th ey a l s o e xh i bi t ev i dence s

of m e ta m e i s m r E v e n th e e xcre to ry ca na l s m ay g i v e up th e i r se g m ent a l
.
”
i s ol a tion a nd b ecom e uni ted to one a noth er by m ea ns of l ongi tudin a l cana l s .
(l b p
. .
On turni ng b a c k to 2 06 , th e rea d er w i ll, I thi nk , d e m ur to the

ass ert i on tha t th e i nd ep end en ce i s co m p ara ti v ely slight s ee i n
g
th a t a s i n Ctenodri l u s , a s i ngl e s egm ent s om eti m es b ec om es s ep a
,
rat e an d rep ro d u ces oth e r s eg m e nts t o for m a n e w s eri es I nstea d .
” “
o f a d m i tti ng tha t the m os t i m p o rta nt o rg a ns s h ow t h em
“
”
s elv es to be s i ngl e fu nd a m ents of th e e nt i re b o dy , i t m ay be
h eld , c ont rari wi s e, th a t th e i r origi na l i nd ep e nd enc e i n each s eg
is ly to th e d egree i n v ol ve d by the i r c oop erati on
ked
“
m e nt m as on
as p a rt s o f a c o m p ou nd org a ni s m B u t chi efly I rem a rk tha t

.
w h e n i t i s s aid th a t th e e x c ret o ry c a nal s m a y gi v e u p th e i r s eg

“
”
m e nta l i s ol a t i o n a nd b e c om e u n i t e d by l ong itu di na l c a nals “
.
t h e re i s a c l ea r c onfes s i o n th at t h e i s ola ti on o f th e s e orga ns w as

ori i na l a nd th ei r u ni o n s up eri nd uc e d
— a n i m pli c a ti on th a t th e
g
c o u rs e o f e v o l uti on i s a s I h a v e d e s c ri be d i t a nd a t v a ri a nce w i th ,
th e c ours e of e vo lu t i on as s u m e d by P rof Kors ch el t . .
Ye t a no th e r i nc ongrui ty is i n v ol v e d i n hi s i nte rp re ta ti on He .
w ri t es
“
Ju t s i n th e c onside ra tion of th e t a pe w orm ch a i n w e were i ndu ced b y
as -
'
t h e com pa ris on w i t h uns eg m ented form s to re fer t he e nti re c ha in to a n

u n s e m ent ed i n di vi dua l , a nd , on th e othe r h a nd , to s ee i n t h e p ro gl ot ti s ,
g
no t a com pl e te i ndi vi du a l , but onl y t h e a bs t r i c t e d h inder porti on o f th e
‘ ’
b ody of th e Ce s tode i n the s a m e m a nne r, a nd wi th m u ch m ore re a s on, we

.
a dh ere t o t h e i ndi vi duali ty of th e A nnel i d body (P . .

6 04 A PP E N DIX D 2 .
A nd then on th e p re ce di ng pa g e, re fe rri ng to th e com p os it i on of

th e A nne lid b o dy , h e s a y s T h e m os t na tural c om p a ri s ons a re
th ose w i th th e t ap ew orm ch a i n a nd w i th th e s tro bil a of th e
’
”
S cyp h om e d usae . Now s i nce i t i s h ere a s s u m e d th a t the ta p e
w orm a nd th e s t robil a a re a na l og ous i n com p os i ti on, i t i s i m pli e d
t ha t t he d eta ch e d p roglotti s a nd t he d et ach e d m e d usa are a na l o
g ous ; a nd h ence i f w e a re to reg ard the p roglottis as not a com “
l et e i ndi v id u al bu t only the a b s t ri ct e d h i nd e r p o rt i on of th e b o d

p ”
y
o f t h e C e s tod e , th en w e m u st s i m ila rly re g a rd th e m ed usa as not
a com pl ete i ndi vid u al , bu t o nly th e a b s tri c t e d hi nd e r p ort i on of
t he s t robi la . Thi s c om m i ts u s to th e s trange c oncl u si on th a t

w h e re as i ndividu a l i ty i s as c rib e d t o th e ori gi nal s i m pl e p olyp ,
a nd by - a nd -by to th e p a rt i a lly s e gm ente d s tro b i l a, th ou gh th ese
-
a re w i th out s p e c i a l s ens es a nd w i th only r udim ents of m u s cu l ar
a nd nerv ou s s y s te m s , i ndivid u a li ty i s d e ni e d to th e d etach e d
m edu s a , w hi ch h as o rg a ns of s ens e , a di s ti nct ner vo -m u s cul a r

s y s te m a n d a c o ns id era bl e p o w er o f l oc o m oti on, as w ell a s a
genera tive s y s te m : t ra its w hi c h i n oth er c as es cha racteri ze

'
de v e lop e d i ndividual s . H ere a ls o, th en, th ere s eem s to be a n

i nvers i on of th e ordi nary concep ti on .
T h is c onc ep ti on of th e p rogl otti s a nd th e m edusa i s , I s e e,

a c cep te d by s o m e as te na bl e . B u t i f w e accept i t w e m ust a c c e p t
a ls o a n a nal o g ou s c on c e pti o n, w h i c h w i l l I t h i nk be reg a rd e d a s
u nte na bl e. I t i s th at s uppli e d by th e Ap hid es . From a n egg

p roceeds a s eri es of s ex les s a nd W i ngles s fem ales , and a t the e nd
of th e s eri es th ere c om e w i ng e d m a l es a nd fem a l es w i th resul ti ng
gam i c reproducti on If i ns tea d o f form i ng a di s crete s eri es the

.
i m p erfect fe m al es form e d a c oncrete s e ri es , the m em b ers of whi c h

c ould i ndi vid u ally fe e d w i th ou t b ei ng d e t ac h e d from one an oth er,
as th e s e gm e nts of a ta p ew o rm c a n, t he p ara ll eli s m w ould be

com pl ete ; a nd th e n, a c cordi n t o th e vi ew i n q u es ti on, w e s h ou ld
g
h a ve to re ga rd th e p e rfect m a e s a nd fem a les e v entu all y a ri s i ng,
no t as i n di v id u al s bu t as t erm i na l p o rt i ons o f th e s eri e s , c o nta i n
i ng ge ne rat i v e p ro du cts a nd h a vi ng wi ngs for t he di sp e rs i on of
t he m — l ocom oti v e eg g - b ea ri ng s eg m ents of th e ch a i n . W h oev er
e s pou s es t hi s vi ew m us t h old e i th e r th a t th e fi rst i m pe rfe ct fe m a le
of th e s e ri es w as th e i ndi v id u a l or tha t t he enti re st ri ng of th em
c onst i tu te d th e i ndi v id ua l ( i n conform i ty wi th a vi ew onc e p ro

p ound e d by P rof H uxl ey ) B ut he m us t do m ore th an t hi s
. . .
S i nce th e A p hid es h a v e d es cend e d from s om e w i nge d s p eci es of

th e o rd e r H em ip ter a , h e m u s t h old t h a t a m ong th ose rem ote
a n c es t o rs e ac h p a rt i cul a r fly , m al e or fem al e , w as a n i ndi vi d u a l ;
but th at w h en a b und ant fo o d a nd i nert life l e d t o t he p a rt heno

geneti c h a bit, a n d to ch a i ns of s exl es s form s , th e m al es a nd
fe m al es e v entua lly p rod u c e d a t t he e nd of e a ch ch a i n, th oug h ,
lik e th ei r re m ote a nces t ors , p oss ess e d of p ro crea tive orga ns a nd
wi ngs , a re not i nd i vi du als .
THE A NNU L OS E T YP E . 6 05
[ S om e m em ora n qu esti on h ere dis cuss ed

d a b e ari ng on th e ,
m is l a id a t th e t i m e wh en th e c h a p t e r d e a l i ng w i th i t w as r ev i s e d ,
h a v e be en d is co v e re d i n t i m e for u tili za ti o n i n th is a p p endix ] .
O n e o f m y c ri t i c s s a y s
You h a v e o v ers t a te d th e case in your fa vour : th e a l i m e nt a ry ca na l doe s
s eg m en ta t i on c or re s po ndi ng t o t h a t of t h e ot h e r
no t, as you s u gge s t, s ho w a
o rg a ns i n A nne li ds . E i th e r i t i s a s i m p l e uniform tu b e , or el se i ts di fi e r
e nt i a t i ons (ph a ryn , x m sOp h a g us , c rop, i nte s ti ne) a re q u i t e i n de pe ndent of
”
t h e r e p e ti ti on of th e s om i t e s .
In p res ence of s ta te m ents m a d e i n w ork s of a uth ority thi s o bje c ,
t i on grea tly s urp ri s es m e I m eet w i th th e d e s cript i v e w or d

.
“
m o nili form
”
a p li e d t o th e i nt es t i n e i n s om e A nne li d s a nd
p ,
th e n i n th e T e x t b oo k of C l aus tra ns l ate d a nd e di te d by

-
,
S e dg wi ck i t i s s a i d c onc e rni ng th e ali m e nta ry c a nal i n th e

, ,
A nne l id a :
“
Thi s i s foll o wed b y t h e ga s tri c regi on of th e gut, w hi ch oc cupi es the
r ea t e s t po rt i on of t h e l e ng th o f th e b ody, a nd i s e i th e r re gul ar l y eon
g
s t ri c ted i n co r re s ponde nce wi th t h e s eg m ents , or p os s es s es l a te ra l di v e rti
c ul a.
(P .
A nd a ga i n on p . 3 6 9 it is sa id
“
Th e i nt es ti ne u su a l l y p rese rv es th e s a m e s tructure i n i t s ent i re l ength
a nd i s di v ided by r egul a r cons tri c ti ons i nt o a nu m b er of di v i s i ons or
c ham b ers, w hi ch corre s po nd to t h e s egm ent s a n d di l a t e a gai n i nt o l a te ra l
d i v e rti c u a a nd casc a
”
l .
The li m ent ary c a na l thu s p res ents th e s eg m ent a l ch a ra cter as

a
c le a rly as c o ns i s ts w i th fu lfil m e nt o f i ts fu nc t i on I f th e s uc .
c ess i ve s e gm e nts a r e c oop era t i ng u n i ts o f a com p o u nd a ni m al
h a vi ng but one m onth th en nece s s a ri ly th e gut c a nnot be c om

, , ,
l etel c ut i nto p a rts ea ch a ns w eri ng t o a s egm e nt for th ere

p y , ,
c ou ld be i n th a t cas e n o p as s a g e for t h e fo o d
, , I f th e p orti on of .
t h e i nte s ti ne b el ong i ng t o e a ch s e gm e nt h as a c ons pi cu ou s dil a

ta t i on or h as a c aecu m o n e a c h s id e i t e x h i bi ts th e s e gm e nta l
, ,
c h a rac te r as m u ch as th e p hy s i c a l r e qu i rem e nts p erm i t S o far .
fro m be i ng a t v a ri a nc e w i th th e h yp ot h es i s i ts s t ru cture e x h i bi ts ,
a verifi ca ti o n o f i t .
T h e n e x t o bj e cti on ru ns as foll o w s
Th en aga in the o v a ri es a nd te ste s do not e xhi bi t a corre spondi ng s e g
, ,
i
m enta t on W hen i t i s a l l ow a bl e to s p ea k of ov a ry or tes t i s a t a l l as i n
.
L u m br wus w e fi nd t ha t i n th w av a t m os t t wo of b oth h
"
, r e ca s e o ga ns e e
pa i rs .
It s e em s di s trib uti on of th e g enerativ e org ans i n

to m e tha t th e
a c om p a ra ti v ely d e v el o p e d m e m be r of th e A nn elid typ e i s not th e
-
,
qu es ti on W e h a ve to as k wh at i t i s i n u nde v el op e d m em bers of
.
t h a t t yp e A m o ng t hem th e re p eti t i on o f ge ne ra t i ve p a rts i s i n

.
s o m e cas e s j u s t wh at th e th e ory i m pli es Thu s i n C la u s I re a d .
I n th e m a ri ne C hw top od a th e o va or Sp e rm a tozoa origi nate on ,
t h e b o dy w all from c e ll s of th e p e ri tone a l m em b ra ne e ith er i n

-
,
A P P E ND I X D 2
'
6 06 .
th e a nt e r or i se g m ents l
a one or l
a on g th e wh ol e l ength of th e
b o dy .
”
S o th a t i n th e s e las t c as e s t he re are , i n al l th e s eg m e nt s;
p arts from w hi ch a ri s e g ene ra t i v e p ro d u cts T h e fac t tha t th es e .
p a rt s a r e n ot d efini t e o v ari e s a n d t es te s i s i rrel e v a nt O v a ri es .
a nd t es t e s a re d e v el ope d g ene ra ti v e s t ru ctu res , a nd i n th e ord e r
o f e v ol uti o n a re p re ce d e d by u nd e v el o p e d ones ; a nd th e fa ct th at
t h es e u nd e v el op e d o nes a re fou nd i n li ttle d e v el op e d m em b ers of -
t he t y pe conform s p e rfe c tly t o th e h yp oth es i s [ I m a

y re m a rk i n .
p a s si ng th at h e re i s a g o od i llu s tra ti on of that p roces s of evol u

t i on w hi ch , i n th e a b o v e s p e cu l ati on of P rof Kors c hel t, i s s u p .
p os e d t o be i nverte d : m any di s p ers ed , s im ila r, a nd i nd efinite

p arts , a re i nt egra t e d i nto a few l oc al i ze d a nd d efi nite p art s ]
I n c o nt i nu a ti on th e c ri t i c a b ov e quote d s ay s My p os i ti on
i s th at th e r ep et iti on of s e gm ents i n a n A nn elid i s a ph eno m enon
o f t h e s a m e n a t u re as th e rep et i t i o n of h a i rs i n a Ma m m al or o f
”
s cu t e s i n a Rep t il e , a nd h e p r oc ee d s t o giv e i ns ta n c e s of rep e
t i ti o ns o f org a ns i n o th er typ es , as of t h e re prod u c ti v e s t ru e

t ure s a nd e x c ret ory s y s t e m i n th e y ou ng D og fis h or of t h e
'
o v a ri e s i n A m p hi ox u s The s e e x a m pl es do n ot s ee m to m e
.
rel e v a nt No p a ra ll eli s m e x i s ts betwe en th e rep eti ti on of a p a r

.
t i cul a r o rg a n i n a n a ni m a l, a nd th e rep e ti ti on of a n ent i re clu s te r

o f o rg a ns cons ti tu ti ng a ph y s i ol ogi c a l wh ol e T h e rep eti ti ons of .
th e o v a ri es i n A m p hi oxu s a nd o f th e e x c ret ory s y s t em i n a y ou ng

D og fis h , o c cur wi th ou t th re a t e ni ng to divid e i nt o s i m il a r p a rts
-
t he e nti re org a ni sm B ut th e s e gm e ntal re p et iti ons i n a n a nnu

.
l os e c rea ture i m plic ate th e s tru c tures a t l arg e, an d w ould , if

p ush ed a li ttle further, res u lt i n s ep ara te c reatures T h e s egm ent .
o f a l ow A nnelid c onta i ns a li m ent a ry , v as cul a r, n ervou s , e xcre
t ory , r ep ro du ctiv e , s ens ory a nd l oc om ot iv e o rga ns— al l th e org a ns

r e q u i re d fo r ca rryi ng on l ife s a v e ce rta i n org a ns o f e xt e rn a l
,
r el a ti on w h i ch i ts p o s i ti on e x c lu d es W h en th ere i s s h ow n s om e
.
v erte brate a nim al , or p roto - v ert e b rate a ni m al , that i s di vi s ible

i nto p a rt s e a ch o f wh i ch i s i n g re a t m e as ure p hy s i ologi cally i nd e
p end ent, I s h all feel oblige d to a ba ndo n m y p os iti on .
W hil e thi s a pp endix i s i n h a nd I ha v e rece ived from a noth er
e xp ert w h os e Vl e W i s i n general agreem ent

,
w i th m
y o w n, a l ette r
c onta i ni ng th e foll owi ng p ass a g e
You wi l l see th at D oh rn s th eory wa s th e a nt i th e s i s of your ow n vi e w

“ ’
of v er te brat e s t ru ct u re , na m el y th a t th e v ert ebra3 w ere form ed by t h e seg -
m e nt a t i on, from m e ch a ni ca l ca us e s of a b ody ori gi na l l y s i m p l e Thi s v i e w .
of yo urs h a s b e en c onfi rm ed b y l a te r re sea r ch es , w hi c h h a v e s h o wn th a t

th e m os t p ri m i ti v e form s a l l i e d to th e V e rt eb ra te s , p os s e s s ing th e e ss ent i a l
o rg ans , viz , gi l l s l i t s , notoc h o rd , a nd dors a l ne rv e c ord, a re not se gm en te d
.
-
a ni m a l s , l i k e A nnel i ds and Crw ta cea , but s i m p l e a ni m a l s , h a vi ng a t m os t

t hree re gi ons , not ex a ct l y c orr es p ondi ng to s egm ent s The se p ri m i ti v e
.
u nse gm e nted form s a re A s ci di a n t a dp ol es , B a l m w l oss us , a n d ce rta i n oth e r

g
p ri m i t i v e f orm s T h e
. e m b ry ol ogy : Ve rteb ra te s a ls o prov es t ha t they a re
'
T HE A NN U L OS E T YP E .
6 07
orig i na l l y s i m pl e a nd not s egm ente d a n i m a l s , e s p ec i a l l y th e fa c t th a t th ere

is o ri gi na l l y one p ron e p hri c d uc t o r p ri m i t i v e k i dney .
”
Ne v ert h e le s s th er e sur vi ves a lea ni ng t ow a rd s th e noti on of a
se gm enta l o ri gi n of th e Ver tebr a ta B u t th e rep eti ti ons of org a ns

.
na m e d i n s upp o r t of t hi s n ot i on h a v e 1 t h i nk n o m o re re la t i o n
, ,
t o th e g e nes i s o f t he v ert ebra t e typ e th an t h e m ult ipli ca t i on of

v ert e b rae i n a s na ke h as rel at i on to the g enes i s of t h e vert eb ra l
c o lu m n .
A P P E ND IX E .
THE SHA P E S AN D A R RA NGE ME NTS OF FL OWE RS .
I N P a rt I V , C h a pte r X , u nd e r th e t i tl e o f “ The Sh a p es of
. .
Flo wers , I h a v e, a fter d es cribi ng th ei r s everal ki nds of s ym m etry ,

”
as h a bi tu a lly rel a te d t o t h e i r p o s i t i o ns , m a d e s o m e r em a rk s by w a
y
o f i nt e r r et a ti o n Th e t ru th th a t flo w ers e x h ibi t a ra di al s m m e try
p .
y
w h en th ey a re s o pl a c e d as t o be e qu a lly a ffecte d a ll rou nd by
i nc id e nt forces , h a vi ng b e en e x em plifi ed , a nd a l s o th e truth th a t
t h ey as s um e a bil a t era l s ym m e try w h en t he y a re s o pl a ce d th at
th e i r t w o s id es a re c ondi ti o ne d i n w a y s d i ffe re nt from t h e w ay s i n
w h i ch th ei r u pp e r a nd l o w er p art s a re c ondi ti one d ; I h a v e g o ne
.
o n t o i nq u i re
( i n 2 3 4 ) y
b w h a t c a u s es s u ch m odifi c a t i o ns o f
fo rm a re p rod u ce d I h a v e s ta te d th a t, origi nally , I i ncli ned to

.
as c rib e th em e nti rely t o differences i n th e rel a ti ons of th e p a rt s
t o p hy s i ca l forc es — l ig ht, h ea t, g ra v i t a ti on, e tc . but th at I fou nd

s u n d ry fa cts s t ood i n t h e w a
y o f thi s i nt erp re ta t i on. A nd I h a v e
s a id tha t
“
Mr D arw i n s i n v es tiga t i ons i nto th e fertili z ati on o f
.
’
”
O rch id s l ed m e t o ta k e i nto a ccou nt a n u nnoti c e d a ge ncy . C on
t i nu i ng t o re cogni ze t he phy s i c a l forc es a s fa ct ors h a vi ng s om e
i nflu ence I ha v e conclu d e d that th e m os t i m p ort ant fa ctor i s th e
,
a cti on o f i ns ec ts ; w h i ch , a idi ng m o st th e fert ili za ti o n of th os e
flowe rs wh i ch m os t fac ili ta te th ei r e ntra nce, p ro d uce , i n cou rs e of

g enerati ons, a form of flow er s p eci ally a da pte d to the s p eci al
p os i ti on.
T hough s till a dh eri ng to th i s i nte rp reta ti on, I h a v e s i nce fou nd

rea s on t o th i nk th at th e o ri i nal i nte rp reta t i on co nt a i ns a l a rg er
g
p o rt i on of t ru th tha n I s u pp os e d a t the t i m e wh en I w as l e d t hus t o
re v i s e i t
. W hile s tayi ng a t Mii rre n, i n S wi tz erla nd , i n 1 8 7 2 , I ob
s er v e d s om e m o difica t i o ns i n a s p e ci es of Ge nti a n , w hi ch p ro v e d to m e
t ha t t he a c ti on of i nc id e nt phy s i c a l forces o n fl ow e rs i s , i n s om e cas e s ,

ve ry rapid a nd d ecid e d Th e s p e c i es fu rnish i ng thi s e vid ence w as the
.
Gen ti a na A scl ep i a d ea ; wh i ch I fou nd i n a c op se form e d o f b ush e s tha t

w ere h e re w id e a p a rt a nd th e re c los e t og ethe r . I n s om e pl a ces not
ne a r t o th e b u s h es , t h e i ndi vid ual s of t h e s p eci es gre w v e rti c ally in
ot her pl a c e s , p a rt i al ly s h a d e d , th e i r i ncli ne d s h o ots cu rv e d i n s u ch
6 08
T HE S H A P E S A ND A RRA N GE ME NT S OF F L OWE R S 6 09 .
di re ct i o ns as to g et t h e m o s t light ; a nd i n othe r c a se s th e i r s h oots

w e re l e d t o t a k e di rec t i o ns alm os t or qu i t e h ori zo nt a l T ha t, a l ong .
w i th t h e s e m dific a t i o ns i n t he di re c ti o ns of th ei r s hoots t here w e nt

o ,
a dj u s t m e nts i n t h e a tti tu d e s o f th e i r l ea v es w as a fa c t n ot s p ec i a lly ,
w o rt h y o f r em a rk ; for pl a nts pl a ce d i ns id e th e w i nd o w s of h ou s es
h a bi tu a lly s h o w u s t h a t l ea v es qu i ckly b en d th em s el v es i nt o a tti
tu d es gi v i ng t h e m th e gre a t es t a m ou nt s of ligh t B u t t he fa ct .
w h i ch a tt ra ct e d m y a tt e nt i o n w as th a t th e fl ow e rs c h a ng e d t h e i r,
a tti tu d es i n a n e q u a lly m a rk e d m a nn e r -
The ra di a l di s tribu ti o n .
p as s e d i nt o a bil at e ra l di s tributi on w ith th e g reat es t rea di ne s s .
C om p a ri s o n of th e a nne x e d figures w ill s how th e ch ara ct er o f th i s

c h a ng e .
Figu re I r ep res ents p a rt of a v e rt i cally grow i ng sh o ot T h i s

. .
be l ong e d t o a n i ndi v id ua l gro wi ng u ni m p e d e d by b us hes a nd g etti ng ,
light on al l s id es H ere i t i s ob s er v a bl e th at th e p a i rs of l e a v e s
.
,
pl a c e d a lte rna tely i n di re cti ons tra ns vers e t o one a noth er— one p a i r
p oi nti ng s a y nort h a nd s ou th a nd th e next p a i r p oi nti ng e as t a nd
, , ,
w est— m a i nt a i n t a ki ng th e m i n th e agg reg a t e a ra di a l di s t rib u t i o n

, ,
a n d i t i s a l s o o b s e rv a bl e t h a t th e a l terna t e p a i rs of flo w ers a re
s i m il a rly a rra ng e d .
Figu re 1 1 i s a s k et ch from a s h oot w hi ch l eaned tow a rd s o ne

.
s id e a nd o f w h i c h t h e h igh er p art as i t b e nt m o r e a nd m or e
, , ,
g ot i ts u pp er s id e m ore a n d m or e di ffere ntl y c o ndi ti one d from i ts

l o w er s id e H ere w e find tha t not only th e l ea v es bu t a ls o the
.
,
fl ow ers h a v e a dj u s te d them s el v es t o t h e ch a ng e d condi t i ons Th e

,
.
l ea v e s of th e l ow es t p a i r h a ng out i n th e norm al w a y on th e op ,
p osi ts s id es of t he a x i s s o t hat a pl a ne p as s i ng through th ei r s u r

,
fa c es w ill c ut th e a x i s t ra ns v e rs ely a nd th e i r tw o a x ill ar y fl o w e r

bu d s c a nd a a re s i m il a rly pl a c e d o n opp o s i t e s id es of th e a x i s
'
.
, ,
B u t a t th e oth er p a rt of th e s hoot w e s ee both th at th e l ea v es h a v e ,
a dj u s t e d th e m s e l v es s o th a t t h e i r pl a nes n o l ong er cutt i ng th e a x i s ,
t ra ns v e rs ely k eep a fi t a dj u s tm ent w i th res p e c t t o th e light ; a nd

,
a l s o th a t th e fl o w e rs no l o nge r o n opp os i te s id e s o f th e a x i s h a v e
, ,
b ent rou nd t o th e upp e r s id e as a t a a nd b , .
Fi g ure III s h ow s u s th i s re a rra ngem ent carri e d s till fu rth e r

.
-
.
T h e s h o ot i t r e pres e nt s w as grow i ng i n a di re ct i on ne a rly h o ri z onta l ,
a n d th ere fo re r e c e i vi ng th e lig ht o nly o n o ne s id e A nd h ere .

,
be s id es s eei ng th a t t h e l e a v es h a v e s o a dj u s t e d th em s el v e s th a t th ey
a l l l i e i n a pp ro x i m a t e ly th e s a m e pl a ne w h i ch i s p a ra ll e l t o t h e a x i s
,
i ns t ea d of t ra ns v ers e t o i t w e s ee th a t th e tw o p ai rs of fl ow er bu ds
,
-
h a v e bo th c o m e round t o th e u pp e r s id e o f th e a x i s S o that .
i n th i s s h o ot th e o rigi na l ra di a l s y m m e t ry i n th e arra ngem ent

,
o f l ea v e s a nd fl o w e rs i s c om pl et ely c h a ng e d i nt o a bil a t e ra l
,
s y m m et ry .
Th es e fa cts do not i t i s tru e p ro v e a ny m odific a ti o n i n th e form s

, ,
85
T HE S H A P E S A ND A RRA NGE ME NT S OF F L O WE RS 6 1 1 .
of th e flow ers th em s el v es th ey o nly p ro v e m odifica ti on i n the

g rou pi ng of th e flo w ers Bu t b ey ond sh ow ing as th ey d o conclu
.
,
s i v e l y h o w rea dily a bil a t e ral a rra ng em e nt o f flo w ers i s p ro du c ibl e

,
o u t of a n a rrang e m e nt th a t w as no t bil a t e ra l by th e a c ti o n of ligh t

, ,
et c . th ey gi v e i nc re as ed p rob a bili ty to t h e b eli ef th a t ch a ng e s i n th e

s h a p es o f fl ow e rs a re p r o d u c i bl e by th e s a m e a e n ci es D ou b t l es s
g .
t h i s ch a ng e i n th e a tti t ud e s of th e fl o w er bu ds i s du e t o th e a c ti on o f
-
ligh t on thei r caly c es a nd p edu ncles m ore th an to i ts a cti on on th ei r

u nfo ldi ng c o ro ll as B ut a l ong w i th a n a c ti on s o d e c id e d on th e gro w th
.
of th es e s h ea th i ng a nd s u pp o rt i ng o rg a ns c o nta i ni ng c hl o rop hyl i t i s ,
s c a rc ely p ro b a bl e th a t th ere i s no a cti on on th e gro wth of th e p e t a l s ,
c o nta i ni ng oth er c ol ou r i ng m a tt e r ; co ns id e ri ng t h a t i n b ot h c a s e s
t h e d e v el op m ent of th e c o lou ri ng m a tt er d ep e nd s o n th e a cti on of

light a nd cons id eri ng als o the effe ct of light on p et als fa m ili a rly
, ,
s h o w n by th e i r o p e ni ng a n d cl os i ng A n d i f e v e n bu t a s m a l l
.
e ffe c t i s p ro d u c i bl e o n t h e g ro w th o f th e c oroll a th e n i t i s t o b e ,
e x p e ct e d t h a t ligh t w ill be a n a e nt i n ch a n i ng th e fo rm o f th e
g
g
c orolla w h e n th e a tti t ud e o f th e fl o w er ca u ses i ts p a rts t o b e d i f
“
ferentl y e xp os e d For a s m a ll e ffect o n th e i ndi v idu a l flo w er w ill

.
b ec om e a gr ea t effec t i n t he fl ow e rs of rem ot e d es c e nd a nts ; pro

v id e d th e c ha ng e d a tti t ud es of th e flo w e rs p r es e r v e c o ns id era bl e
c ons t a n cy th ro u gh ou t t h e s u c c e s s i o n o f i ndivid u a l s .
B e th i s as i t m ay h o w e ve r th e fa c ts I h a v e h ere d es crib e d
, , ,
w h i c h I d ou b t n ot oth er o bs e r v e rs h a v e s ee n p a ra ll el e d i n oth er
pla nts are i ns tru cti v e as s h o w i ng h ow q ui ckly c ertai n m et am e r
, ,
ph e s es a re p rodu ce d a nd as i m plyi ng th e e as y e st a bli shm ent of s u ch

,
m e t a m o rp h os es as p e rm a n ent c h a r a c t e rs i n a s p e c i es if t h e m o di fy ,
i ng c ondit i ons b ec om e p erm a ne nt Th e c h anges o f arra nge m e nt

.
I h a v e p oi nt ed ou t do not becom e p erm a ne nt i n thi s s p ec i es b eca us e

,
i ts i n di v id u als are v a ri ous ly a ffe ct e d by th e m o d i fyi ng force s : o n

s o m e th ey d o n ot a c t a t a l l on s om e a li ttl e o n s om e m u ch
, ,
a nd
e v e n o n th e s am e i ndi v id u a l t h e di f fere nt s h oot s are qu i te di fferently

af fec t e d B u t if th e h ab i t of th i s pla nt w e re grea tly c ha nge d — i f
.
,
fo r i ns t a nc e by s p rea di ng i nt o h a bi ta t s yi e ld i ng a bund a nt nu t ri
,
m e nt th e l a nt b eca m e v ery lu xu ri a nt a nd m ult iplyi ng i ts bra n ch es

, p , , ,
re w s h ru b lik e ; i t i s c l ea r th a t b e i ng s h a d e d by o n e a noth e r th e s e
g
-
, ,
bra nch es w ou ld b e h a bi tu a lly c i rcu m s t a nce d i n a w a y lik e th a t whi ch

w e h ere s e e p ro d u ces bila t era l ne ss i n th e di s t ri bu ti o n of th e fl o w e rs ,
if not i n t he fl o w ers th e m s el ve s ; a n d be i ng t hu s p erm a ne ntly

af fe ct e d w ould b e c om e p erm a ne ntly bil at eral
,
A c cu m u l a t i ng by
.
i nherita nc e w hat i s h e re o nly a n i ndi v i du a l p e cu li a ri ty w ould

, ,
b ec om e a p eculi a ri ty of th e s p eci es— a s p e cifi c ch ara cter .

A P P E ND IX F.
P HYS I OL OGICAL (OR CONSTITUTI ONA L ) UNITS .
T H E R E h as re c e ntly com e before m e a fa ct w h i ch has a s igni fi

c a nt bea r i ng o n t h e h yp o th e s i s o f C o ns ti tu t i on a l u n i ts : s er v i ng,
i nd eed , t o gi v e a n app a rently concl us iv e p roof of i ts t ruth B efore .
s t ati ng i t, h ow e v er, I m a
y w i th a d v a nta g e re s t a te t he s e v era l
-
e vid ences a l rea dy ass igne d i n s upp ort of i t .
"
1 Fi rs t com es the a: p ri ori re as on Th es e u ni ts i n th e g erm of

. .
a n o rg a ni s m w hi ch c a u s e d e v e l op m ent i nto a s p e ci a l s t ru c ture ,
c a nnot be chem i ca l u ni ts — ca nno t be s i m ply m ol e c u l e s o f p ro te id
s u bs t a nc e i n o n e o r o th e r o f i ts fo rm s ; s i nce th es e a re n ot s p e c i a l
t o a ny typ e o f c rea t ure bu t c o m m o n t o a ll cre a tu res Nor c a n .
th ey be w ha t w e m a y c all m orp ho l og i ca l uni ts —th e cells or p roto

l as ts ; b eca u s e i n th e ea rly s t a g es o f d e v e l op m e nt t h e cel ls o f
p
o n e org a ni s m a re i ndi s ti n u i s h a bl e fro m
g t h o s e o f o th e rs a nd ,
b e c a u s e w ere c ells th e u ni ts o f com p os iti on th e re c ould be no

i nterp reta ti on o f w h at a re c alle d u ni c ellul a r org a ni s m s — nothi ng
t o a c c o u nt for th e i nnu m e r a bl e v a ri eti es of t h e m H e nc e o f .
,
n ecess i ty t h e s tru c tu ra l e l e m e nts o f w h i c h e a ch org a n i s m i s buil t

, ,
b ei ng ne i ther p rote id m olec u l es nor c ells m u s t be s om ethi ng ,
bet w ee n th em : p ro b a bly s om e com pl ex c om bi n at i on of d iffe re nt

i s om eri c form s of p rote i ds .
2 . T hat uni ts o f s u ch na tu re s are th e p onents o f

e s s ent a i l com
ea ch s p e c i es of o r a ni s m i s s h o w n by th e fa c t th a t i n l o w typ es o f
g ,
c rea t u res li ttl e di fl e re nti a te d i nto s p ec i a l ti s s u e s a n y c ons id era bl e

,
‘
p orti o n of th e b ody wi ll w h en s ep a ra te d b eg i n to as s u m e t h e
, ,
s t ruc t u re p ro p e r t o th e s p e c i es — a t ru th re c e ntl y s h o w n a fres h by
P ro f T H Mo rg a n s e x p e ri m e nts o n t h e re e nera ti o n of P l a na r ia
’
g
. . .
m a c u l a ta
( a l re a dy re fe rre d t o i n
S 2 06 ) sh o w i ng th a t v a ri ou s frag
m e nts c ut o ut d e v el op i nt o n e w i ndi vid u a l s a nd th a t w h e n b e i ng
, ,
to o s m a ll th e y d i e b e fore d oi ng th i s th ere i s a l w a y s a n a bort i v e

,
a tt em p t t o as s u m e t h e s p e cific s tru c t u re .
3 . T hi s t ru th th a t a p orti on o f u ndi ffere nt i a te d t i s s u e , i f a d e

q u a te i n q u a nti ty , ass um es th e s t ru cture of th e typ e , illus trat i ng
612
P H Y SI OL OGICA L ( OR CONS T I T U T I ONA L ) U NI TS .
61 3
as it d oes p ro cli v i ty of the c onsti tuti ona l uni ts t ow ard s th e

th e
s tru c tu re o f th e s p ec i es a lli es i ts e lf w i th t h e p h e no m e na o f b ot h
,
a g a m og enes i s a nd g a m og ene s i s Th e firs t o f t h es e s h o ws u s h o w

.
a fi s s i aro u s l y d et a c h e d p o rt i o n of th e p a r ent a l t i s s u e t a k es o n
p
-
th e s a m e fo rm as th e p arent ; a nd th e s econd s h o ws h ow th os e
s m a ll d et ac h e d p o rti o ns di s t i n i s he d a s s p e rm cell a nd g e rm cel l
- -
a l s o w h e n u ni te d a n d s u ppli e
,
w i th t he ne e dfu l m a t e ri a l s d o th e ,
s a m e t hi ng .
4 . B u t th e s et of ph enom e na foll o wi ng th e u ni o n of s p e rm c ell -
a nd g e rm c ell d i ffe r i n a c ert a i n w a f ro m t h o s e w h i c h foll o w

y
-
w h en a g am m a o r ot h e r u nfe rt ili z e d p o rt i o n of p a re nt a l t i ss u e i s
d et ach e d The i ncom p reh e ns ibl eness o f thi s d ifferenc e as oth e r
.
w i s e c o nt e m pl a t e d , a nd t h e p a rt i a l c om p reh e ns ibl e ne s s o f i t w h e n
j o i ne d w ith t he hyp othes i s of phy s i ol ogi ca l u ni ts , furni s h a
fu rth er s upp o rt for th e hyp oth e s i s .
Th e fa m ili ar t ru th l ea rnt by th e ty ro i n alg e b ra th at a n a p

p a rent s olu ti on w hi ch c onta i ns the u nknow n qu a ntity i s no
s o lu ti on , i s a tru th a t t o be o v e rl o ok e d i n oth e r s p h e r e s t h a n
p
th e al g e bra i c A n il l u s tra t i on i s s uppli e d by th e a ns w e r o nce
.
gi v en i n P a rli am ent to th e qu es ti on W ha t i s a n A rch d ea co n

”
O ne w h o di s ch a rges a r ch idi a c ona l fu ncti ons .B u t s ci e n c e a s
w ell a s d a ily li fe fu rni s h es e x a m pl es W h en i t i s s a id by E ng el
.
m a nn , H en s e n , H e rt w ig , a nd Ma u p as th a t th e es se nti a l end of
“
s e x u a li ty i s rej u ve n e s c ence , th a t i s , th e res t ora t i o n o f g row t h
ene rgy , w e h a ve a noth e r i ns ta nce of a n e x pl a na ti on w h i ch e x pl a i ns

noth i ng . W ha t i s th e ph enom enon t o be e x pl a i ne d ? Th at un
foldi ng o f a n org a ni s m fro m a g erm w h i c h di s play s g row th e nergy -
.
A nd w ha t i s th e e x pl ana ti on ? Th e givi ng of fres h growt h ene rgy -

.
Th e u nknow n qu a nti ty growth energy i s c o nta ine d i n th e

-
e x pl a na ti on p rop os e d T h ere e x i st s no c o n cep ti on of

.
ju ven
e s c en c e s a v e th a t d eri v e d from o b s e rvi ng d e v el opi ng pl a nt s a nd
be p refi x ed , n o i nt e rp re t a ti o n i s th ereby
”
a ni m a l s ; a nd if
“
re
”
gi v e n t o t he u ne xpla i ne d th i ng j u v enes ce nc e .
C o l e rid g e s om ew h ere c om m e nts o n a s ou rce of fall a cy wh 1 cl rh e

— t h e ch a ngi ng of a rel a ti on
O
ll
c a s th e
“
h yp i
os t as s o f a r e l a t i o n
.
i nt o a th i ng Th e plu m be r w h o t ells you th at w a ter ri s es i n a

.
pu m p “ by s u cti on su ppli es a n i ns t ance H a vm g a ss u m e d s uct i o n

.
t o be a n a g ent , h e thi nk s t h at h e u nd e rs ta nd s h ow t e pi s t on d o es h
i ts w ork . S om e of th e e x pl a na ti o ns gi v en of ferti l i z a t i on s u pply
fu rth er i ns t a nces W h e n i t i s s a id th at s e x u al u ni on h a s for i ts
.
e nd
“
t o gi v e i n cre a se d v i g ou r t o a l l th e v i t a l p ro ces s es , it is
h i ch
.
t i tly i pli d t h t v i ou r i s a s o m e th i ng
— a s o m eth i ng w
a c m e a g
c a n b e gi ve n B u t n ow , i n th e fi rs t pl a ce , i t i s o nly by th e h yp o
.
s t a s i s o f a r el a t i o n th at w e are l e d t o th i nk of vig o u r a s a th i ng .
Vig ou r i s a s tate — th at s ta t e of a livi ng b ody wh i ch enabl e s i t t o

61 4 A P P E ND I X F .
g i v e o u t m u ch m oti on . W h a t e n a bl es i t t o d o th i s ? T h e p re s e n c e
i n i t of a bu nd a nt m ol ec ul es c o nt a i ni ng m u ch m ol ecul ar m ot i o n
w h i ch c a n be t ra ns form e d i nt o m ol ar m oti on : t h e t ra ns fo rm a t i o n
bei ng e fl e c t e d by t he fa l li ng of t h ese m ol e cul es i nto th e i r s i m pl e r
a nd r el a t i v ely i ne rt c o m p o ne nts
-
wh i ch a re th e reup on ex c re t e d
, .
E ne rg y cont a i ni ng m a tt e r i s u s e d up a nd m ore e ne rgy o r vig ou r

-
,
c a n be
g i v e n o nly b
y s upplyi ng m ore s u ch m a tte r H ow th e n .
c a n th e u ni o n o f t wo nu c l e i — t h os e o f th e s p erm c ell a nd g erm -
c ell— gi v e vi g o u r ? Onl y a n i nfini t es i m al p ort i on of v i g ou r i n th e

s ens e a b o v e e x pl a i ne d e xi s ts i n e i th e r a nd t h e u ni on of th em
,
l ea v es i t s ti ll i nfi ni t es i m a l A nd th en e v en s uppos i ng th e vi gou r
.
,
t o be a n e nti ty a nd t o be a pp rec i a bl e i n qu a nt i ty h o w c ou ld i t ,
o on p ro d u c i ng t ha t i m m e ns e co m bi na t i o n o f ph y s i ol o gi ca l
g
a cti o ns s ee n i n th e u n foldi ng o f th e g e rm i nto a n o rg a ni s m ? a nd
h o w c ould i t g o o n p ro d u c i ng th e ph y s i ol ogi c al a ct i ons of a n a d ult

o rg a ni s m d uri ng a w h ol e c e nt u ry ?
Ma y w e not th e n s a y th a t th e s e p rop o s ed e xpl a nati o ns l e a ve the

que s ti on w h ere i t w as — a re nom i na l s ol uti ons not real s oluti o ns ? ,
5 . But th e hyp oth es i s i tuti o nal units furni sh es if not a

of c o n s t ,
s a t i s fac t ory a ns w er e t s o m eth i ng i n th e n a tu re of a n a ns w e r— a

y ,
t rue ca us e ; t h a t i s t o s a y a c a us e a c tu a lly k now n t o u s as

,
o p era ti ng i n o th er c a s e s I n § 9 2 i t w as p oi nt e d o ut th at i n
.
p rop o rt i on as u ni ts a re s i m ila r t he re m ay be built u p from them

,
a n a g reg a te wh i ch i s rel a t iv e ly s t a bl e a n d t ha t a l ong w i th i a

g ,
c r ea s i ng di ss i m il a ri ty th e s t a bili t y o f th e a gg reg a t e d ecr e a s es It .
w as i nfe rre d t h a t i f a g rou p of cons ti tu ti ona l u ni ts b el o ngi ng t o

o ne i ndi v id u a l w h i c h h a v e be co m e m ou ld e d i nto rel a t i v e ly e x a ct
c o ng ru i ty wi th t h e o rg a ni s m a nd wi th o ne a noth e r by l ong c o
Op erat i on a re m i ngl e d with s om e b el ongi ng t o a noth er i ndi v id ual

,
w hi ch di ffe re ntly c i rc u m s ta nc e d h as b e com e s om ew h a t di ffe re nt

, ,
i n i ts el f a nd i n i ts u ni ts the n t h e m as s form e d by t he u ni on of
,
th e t w o gr ou p s w ill be rel a ti v ely u ns t a ble — rel a t iv ely m odi fia bl e

by i ncid ent forces W he re as i n e it h er o rg a ni s m no l onger p e r
.
,
petua ll y ch a ng e d i n th e r el a ti o ns of i ts pa rts by grow th th e re i s ,
a n a pp roa ch t o w a r ds e q u ili bri u m b et we en the wh ol e a nd i ts c om

p o n e nt s t h e
, co m p o n e nt s c o n t ri b u t e d by th e t w o t o f o rm a
germ b ei ng s lightly u nlik e one a nother w ill not fo rm a group

, ,
i n a s t a te o f e qu ilib ri u m Th e g ro up t hey form w il l be c a p a bl e

.
o f eas y c h a ng e b i n c id ent fo rc e s ; a nd th e y w ill s o be re nd ered

y
fre e t o foll o w th ei r p ro cli vi ti es tow a rds th e typi c a l form o f th e
s ec i es
p I nfe rri ng t hi s w e m u s t a ls o i nfe r tha t s o l o ng as the s e
.
t w o s e ts of s lightly di ffe re nt u ni ts a re not e x p os e d to a ny c ons t a nt

forces te ndi ng t o c oe rc e th em i nto t he s a m e form th ere w ill c on ,
ti nne t o e x is t i n th e nu c l e i o f al l d es c e nd a nt cells thi s s a m e re l a

t i v e i ns ta bili ty a nd c o ns e q u e nt pl as ti ci ty .
P H YSI OL OGI CA L (OR CONST I T U T I ONA L ) U NI T S .
61 5
S u ch e v id ence as w e h a v e v e rifies thi s i nterp reta ti on T he re i s

.
firs t th e uni v e rsa l fact th a t d e v elopm e nt o f th e g erm begi ns

wh en i t i s e x p os e d t o a n i nc id ent fo rce— h ea t — th e u nd u l a ti ons of
w hi c h , i nc re a s i ng th e o s c illa ti o ns o f t h e m i x e d u ni ts , gi ve t h e m
g rea te r fre ed om to a rra ng e the m s el v es i n c onform ity w i th th ei r
.
typ e . W e s e e t hi s alik e wh e n s p ri ng w arm th m a k es a s e e d

g erm i na te a nd wh en th e w a rm th of a s i tt i ng h en s ets u p org a ni z a
t i o n i n h e r e ggs . H ea t frees th e m ol e cu l es o f i norga ni c m a tt e r
from l o c a l r e s t ra i nts a nd , as w e s ee i n m ol ten m e ta l , l ets t h e m
yi eld t o oth er forces ; a nd s im il arly i n thi s o rg a ni c m atter, t h e
u ni t s a re m a d e fre e t o foll o w th ei r p r o cli v i t i es . Th e n, s e c on dly ,
th er e c o m e s th e e vid e n c e from c o m p a ri s o ns b e tw e en th e effe cts of
m ixi ng c o ns ti tu ti o n a l u ni ts di ffe ri ng i n v a ri o us d egre es L e t th e
.
cl u s ter o f m i x e d u n i ts b e d e ri v e d from a n i m a l s th a t a r e o rdi na lly
di s ti nct Noth i ng h a pp ens Th e uni ts ea ch c ontrib utes t e nd t o

. .
a rra ng e t h em s el v e s a ft e r th e p are nt al t yp e . H enc e a c onfli c t

be t w e en th e t e nd e nc i es t o w a rd s tw o m a rk e dly u nlik e s t ru c tu res ,
a n d no s t ru c t u r e a ri s e s
. S u pp os e th e m i x e d u ni ts c om e from t w o
ki nd re d s peci es f s ay h ors e a nd as s T he s tru ctu res w hi ch th ey
.
r es p e c t i v ely t end t o form , b ei ng i n t h e i r m a i n c h a ra c t ers a lik e ,
t h e re i s s u c h Co op e ra ti on as p ro d u ces a w orki ng org a ni s m bu t a n

o rg a ni s m i n c e rt a i n res p ects i m p e rfec t— a m u l e . S upp o s e , a ga i n,
t he u nits c om e from t w o v a r i et i e s of th e s a m e s p e c i es . A p erfe ct
o rg a n i s m r e s ul ts , a nd , a s s h o wn by Mr . D ar w i n w h en d eta ili ng
t he e ffec ts o f c ros s i ng, a n u nu s u a lly vi g orou s o rg a ni s m .T he
u ni ts b ei ng m o re u nlik e th a n th o s e b e l o ngi ng t o t h e sa m e v a ri e ty ,
t he i ns t a bili ty of t h e g e rm pl as m i s u nu s u a lly g reat , a n d th e

-
tra ns form at i o ns w h i ch c o ns ti t ut e d e v el op m ent a nd a c t i on b e com e

u nu s u a lly a c tiv e . W h en, a s i n ordi nary c as es , th e u ni ts a re s up
pli e d by m em b ers of th e s a m e v a ri ety w h o h a v e not b e en m a d e
ve ry m u ch a lik e by th ei r a ntece de nts , th ere foll o ws th e us u a l
a m o u nt o f o rg an i c v ig ou r . C o m i ng now t o the r es ults of b ree d
i ng i n a n d i n — b re e di ng b et w e e n i n di v id u a ls w h os e c ons t i tu t i o ns
- -
e c o ns ti tu t i o na l uni ts ) h a v e fo r g e n era ti on s bee n g row i ng m o re

( L .
a li k e i n t h e a b s e n c e o f c ro s s i ng w i th ot h e r s t i rp s — w e s e e t h a t
di m i nuti on of org a ni c vi gou r i s di spl ay e d : th e re i s a d ecrea s e i n

t h e r a t e o f p hy s i ol og i ca l c h a ng e . Fi nally , on c om i ng t o a cl o s e r
rel a t i o nsh ip , as i n m arri a g es b e t w ee n cou s i ns , i n w h o m t h e
c o ns ti tu ti o na l u ni ts a r e m ore t h a n c o m m o nly a lik e, w e s e e th e r e
fre qu e ntly foll ow s e i th er ba rr enn es s or th e p rod u c ti o n of fe ebl e

of fs p ri ng .
Al l th es e fa ct s , th e n, are c ongr u ou s w i th th e hyp oth es i s t h at

th e u se o f fe rt ili z a ti on i s th e m i x i ng of u nlik e u ni ts , a nd co u s e
qu ent p ro d u cti on of plas ti city L ea v i ng out c as es i n w hi ch the
.
u nl ik enes s es a re s o gr ea t as w holly t o p re v e nt c oop e ra t i on a m o ng
th e u ni ts , th e d e gre e of v ig ou r, t h a t i s , t h e a c t ivi ty of phys i ol ogi

61 6 A P P E ND IX F .
c a l c ha n ge i s grea t wh ere th e u nlik eness

,
is gre at a nd di m i ni sh es
w i th th e a pp roa ch t ow ard s lik e nes s .
6 . Th e e x i s tenc e of c o ns ti tu ti onal u ni ts s eem s o th erwi s e

ne ces s a ril y i m pli e d I re fer t o th e fac t th at no o rg ani s m i s a
.
,
h om og en eous m ea n bet we e n i ts p a re nts but c o ns is ts o f a m i x t ure

o f p a rts , s om e foll o w i ng o ne p a re nt a nd s om e t h e o th er A m o ng .
illust ra ti ons of th i s th e m o s t c ons pi cu ous a re t hos e yi eld ed by

t h e v a ri ou sly -m i x e d c ol ou rs o f h a i r or fea th ers H ors es , ca t tl e , .
d ogs , c a ts , h e ns , pige ons di spl ay th es e m i x tures : c ol ours i n o ne

p l a ce l ik e th e m o t h er a nd i n a no th er pl a c e lik e th e fa th e r A s th e .
i nte rnal o rg a ns a re i n vi s ibl e , a nd as vi s ibl e o rg a ns h a v e i nd efi ni te

s ha p e s a n d g ra d u a t e i nd efi ni t ely i nto a dj ac e nt o ne s , th e m ixt u re
o f t ra i ts i s e l s e w h ere l es s cons pi cu o u s ; bu t oc cas i o na l m ark e d

'
cas es e sp e c i a lly i n m al fo rm a ti o ns l e a v e n o d ou b t th a t i t p e rv a d es
( )
t h e e nt i re org a ni s m .
T hi s p ecu li a ri ty of tra ns m i ss i on s ee m s necess a ri ly t o i m ply th a t

th e re a re di s ti nc t u ni t s d eri ve d from th e t w o p a re nts , a nd t h at i n
th e c ou rs e of d e v e l op m e nt th e re i s m ore or l ess s e g re g a ti on o f
th e m — th os e o f th e o n e o ri gi n p re d om i na t i ng s o fa r i n s om e
pla c es as t o gi v e s p eci al lik eness t o one p arent, a nd th os e d e ri v e d
fro m th e oth e r d o i ng t h e lik e i n oth er pl a c es A ll w hi c h i nte r .
re t a ti o n i s i m p o ibl e u nl ess t he h yp oth es i s of c ons ti tu ti o na l

p s s
u ni ts be a d m i tt e d .
7 . I l ength t o th e s p eci al e vid ence referre d to a t t he

c om e a t
o u ts et . I t i s e vi d e nce of the s a m e na tu re as th a t j u s t as s ig ned ,
but c arri e d to a h igh e r s tag e I t i s fu rn i s h e d not by th e s egre

.
gati e n o f t ra i ts d eri v ed from t wo p a rents of th e s a m e v a ri ety ,
but i s fu rn i sh e d by th e s eg reg a t i on o f t ra i ts d eri ve d from p a re nt s

o f di f fe re nt v a ri e ti es I n a rt i cl es on B ud Va ri a ti ons or S p orts
.
( G a r dener s Chr oni cl e 1 8 9 1

’
) D r M as t ers giv es v ari o u s e x a m pl es
,
.
o f th e s ep ara ti o n or u nm i x i ng o f a nc es tra l c o ns ti tut i o ns Mr . .
Nobl e fo rm ed a hyb ri d b e tw een Cl em a ti s J a chm a m a n d C p a l m s

'
. .
O ne of th es e v a ri et i es fl ow e rs i n th e a utu m n on new w oo d w hil e ,
th e o th e r fl o w ers i n t h e s p ri ng o n ol d w o o d ; a nd th e res u l t i s
t h a t fl ow ers o f t w o ki nds q u i t e u nlik e a re p rod u ce d a t di ffere nt
, ,
p a rt s of th e y ear a nd th at by p ru ni ng s o as to c ut a w ay o ne o r
,
o th er s et of s h o ot s th e pl a nt m a y be m a d e t o p ro d u ce e x cl us i v ely
,
for th e t i m e b ei ng one or oth er s ort of flow er .
A nothe r v e ry i nt e e s ti ng c a s e of unm i xi ng or if i t b e pre fe rre d of p a r

r , , ,
t i a l m i x t u re i s a ffo ded by Ne b ert s B e rbe ri s Thi s i s a hyb id b e t wee n

’
r u . r
,
the e v e rgr een p i nn a te l ea v e d Ma h oni a a nd th e dec i du ous s i m p l e l ea v ed

- -
B erb e i s vu l g a ri s a nd i t b ea rs l e a v e s som e of whi ch a re i nterm edi a te i n

r ,
a pp ea ra nce, w hi l e oth e rs a re m u ch l i ke t hos e of one o r ot her o f i t s p a re nt s .
“
A n ot u ncom m on i l l u s t ra t i on of a s i m i l a r ki nd , i s th e p rod uct i on o f a
P ea ch and a Ne ct a ri ne on t h e s am e b ra nch, a nd we ha v e j
u s t l ea rnt from
P H Y S I OL OGI CA L ( OR CONS TIT U T IONA L ) U NIT S .
61 7
Canon E l l a c om be th a t som e of th e Be rl i n Hel l ebor es sh ow e vi dence of th e i r

h yb ri d na t u re b y oc c a s iona l l y p rodu ci ng fol i age [ a nd o we rs fl of t he t wo
p a re nts se p a ra tel y from t he s a m e r oot s tock

-
.
I n a dd i ti on t o the ca s e s gi ve n a b ov e , w e m a y h e re ci te a fe w m ore whi ch

h a v e com e und er ou r n ot i ce, s uc h a s a Ch rys a nth em u m , h a l f t h e ore ts of fl
whi ch a r e of one col our , h a l f of a noth er . A hyb ri d Ca l a nthe , s howi ng a
s i m i l a r p i e ba l d v a ri a t i on, i s sh ow n i n Fi g 1 4
. A v ery curi ou s ca s e w as th a t
.
of t he Na rc i s su s rece i v e d from Mr Wa l k e r, a n d i n wh i ch
. fl
o we rs o f t wo
di s ti nc t v a ri et i e s s p ra ng from th e s a m e bul b Gra p es not uncom m onl y
.
s h o w t h ei r c ros s e d or i i n by p re s enti n
g g a s t ri pe d a pp ea ra nce, one s tri pe
be i ng of one col o ur, o ne of a noth er , a s m ay a l so b e s een i n t h e Orange ,
”
A ppl e , L e m on, a nd Curra nt .
Th u s , h ow e v er th e g erm pl a s m i s c ons t i tu t e d i ts esse nt i a l c om

-
p onente cannot be a l l a lik e B efore th ere c a n be thi s dis s oci ati on

.
o f a n c es tra l c h a r a ct e rs , th e re m u s t be i n th e g e rm pl as m di f fere nt
-
e l e m e nts c a p abl e o f b e i ng di s s oci a te d T hi s s i ngl e fa ct s e em s t o

.
c om p el us to as s um e cons t i tuti onal un i ts .

A P P E ND IX G .
THE I NHE RITA NCE OF FUNCTIONA L L Y-CA USE D

MODIFI CA TIONS .
I N P a rt II , Ch a p t e r X A , I h a v e c onfes s e d th a t t h e p rocess by
w h i ch a s t ru c tu re c ha ng ed by u s e o r di s u s e a ffe c ts th e s p e r m
c e ll s or g erm c ells w h en c e a ri s e d es cen d a nts , i s u ni m a i na bl e :
g
-
w i thou t, h ow e v er, i nferri ng th a t th ere fore s u ch a p ro ce s s d o es

no t e x i s t . W i th oth ers i t s eem s different S om e th ree y ears .
ag o t h e foll o w i ng e x p res s i on o f Opi ni on c a m e t o m e from a
zoo ol gi cal e x p ert

Ma ny z ool ogi s t s — m os t o
f u s h e re a t Ca m b ridge — a re i nten s el y opp os e d
t o th e doct ri n e of the i nh er i ta bi l i ty of a cq ui re d v a ri a ti ons E v e n a ss u m in g
.
th a t th e d ev e l op m e nt a l p ow e r of a g e rm i s dete rm i ne d b y i ts m ol e cul a r
s t ru c tu re (a nd I for o ne w oul d q ue s ti o n t h i s — D ri e s ch a nd hi s s ch ool when
th ey fi nd th a t t h ey ca n s q u ee ze a d ev el opi ng e gg i nto a ll s ort s o f s h a p e s
wi th out a l te ri ng t h e fi na l re su l t , t h a t one b l a s tom er e i n a n e gg wh i ch h a s
di v ided i nt o 8 i s s ti l l a bl e t o rep rod uce a w hol e e m b ry o— q ues ti on i t a l so) ,
we s t i ll fa il t o conc e i v e a ny m ea ns by whi ch , for i ns t a nce , a c h a nge i n t h e
de v el opm ent o f a m u s c l e or n e rv e c an e ffect a corre s pondi ng ch a ng e i n tha t
p a rt of th e g e rm whi ch i s de s ti ne d t o p rodu ce a corr e s pondi ng p a rt i n th e
”
des cenda nt .
H ere i t w ill be Ob ser v ed th a t b eli ef i n t he i nh e ri ta nc e of

s t ru c tu ral ef fe cts w rou ght by u s e a nd di s u s e i s rej e ct e d be cau s e
,
o f i na bili ty
“
t o c o nc e i v e a ny m ea ns by w hi ch th e m odifi ca ti ons
p rod u ced i n a n organ can effect a c orrelat ed m odifi cati on i n th e
e rm o f a d es cend a nt : fa il ure t o c o nc ei v e i s th e t es t T h e i m
g .
p li c a t i o n i s t h a t s o m e a l t e rna t i v e h yp o th e s i s i s a c c e p t e d b e ca u s e
t h e correl a ti ng o f a v a ri a ti on i n a n org a n w i th a c orre s p ondi ng
g erm v a ri a ti o n i s e ffe c t e d by a m ea ns w hi ch i s c once iva bl e T hi s

-
.
i s th e hyp oth es i s of W e i s m a nn . C once rni ng i ts c onc e i v a bili ty l

h a v e i n th e ch a p t er j us t na m ed a l rea dy w ri tte n as follo w s
, ,
If we fol l o w P rof W ei s m a nn w e a re l e d in to a n a s toundi ng s upp os i ti on

. .
He a dm i t s th a t e v ery v a ri a bl e p a rt m u s t h a v e a s p eci al d eterm i na nt , a nd

th a t t h i s r e s ul t s i n t he as s um p ti on of ov er t wo h undred th ous and for t h e
four w i ngs o f a b utte r y fl . L e t u s a sk wh a t m us t h ap pe n i n th e c as e of a
p ea coc k s fe a th er On l ook i ng a t th e e ye ne a r i t s e n d, we s ee t h a t th e
’
.
m i nut e p roces s es on t h e e d e of ea c h l a t e ra l th re a d m us t h a v e b ee n i n s om e
g
j
w a y e x ac t l y a d u s ted, i n c ol ou r a n d
p os i ti on s o a s to fa l l i nt o l i ne w i th t h e
.
j
p roc e s s e s o n a d a cen t th re a ds : oth erw i s e the s ym m e tri cal a rra nge m e nt of
col our ed ri n s w oul d b e i m E a ch o f thes e p roc e ss es , th en, b e i ng a n
g p os s i bl e
.
618
FU NCTI ONA L L Y —CA U S E D MOD I FI CA T IONS .
61 9
i nde p e ndent v a ri a b l e , m us t hav e h a d i ts p a rt i c ul a r det erm in a nt Now th ere .
a re a b out 3 00 t h r ea ds on th e sh a ft O f a l a rge fe a th er a nd ea ch o f t h em ,
b ea rs on t he a v era ge p roce s s es , m aki ng for th e wh ol e fea th e r

O f th e s e p rocess e s For one fe a th e r a l on e t here m u s t ha v e be en
.
dete rm ina nt s , a nd for th e w h ol e ta i l m a ny m i l l i ons A nd t he s e , a l ong w i th th e

.
de te rm i n a nt s for the de ta il ed p a rt s of a l l t he oth e r fe a the rs , a nd for th e

v a ri a bl e c om ponents of a l l organs form ing th e body a t l a rv e, m u s t h a v e been
cont a i n ed i n t h e m i cr o scO pi c h e a d of a s p e rm a t ozo on ! A nd ea ch of th em
m us t , t h rou gh out a l l t h e co m p l e x dev el op m en t a l p r oc es s e s , h a v e p re s e rv e d
t h e a b i l i ty t o fi nd i t s w a v to t he e x a ct p l a ce wh e re i t w a s w a nt e d 1]
If m
y C a m b ridg e c orre s p o nd e nt i s a bl e t o co nce i v e thi s p ro ce s s
i m pli ed by the hyp ot hes i s of W ei s m a nn, I c an o nly s a
y th a t h e
h as a n e n v i a bl e p ow er o f im agi na ti on .
B u t no w c o m esge fa ct th a t a n i m p os s i bili ty of thou ght

th e s tr a n
i m pli e d by W ei s m a nn s hyp oth es i s d o es not ca u s e rej e cti on of i t

’
,
bu t y et i s u rge d a s a r ea s on fo r rej ect i ng a n a lt e rna t i v e h yp o

th es i s w h i ch d oe s n ot i m ply i t On e o bj ec t or c a nnot c once i v e
.
th a t a c h a ng e i n th e d e v e lop m e nt o f a m u s cl e or n er v e ca n
“
ef fe c t a co rres pondi ng ch a ng e i n th a t p a rt o f th e g e rm w h i ch i s
”
d es ti ne d t o p r od uce a corr espond i ng p a r t i n th e d es ce nd a nt ;
a nd a no th e r o bj ect or s a y s i t i s v ery h ard t o b eli e v e th a t a
“ ”
fu nc ti o nally ch a ng e d o rg a n w ill s o a ffect s p erm at ozoa a nd ova

-
th a t one p a r ti cu l a r p a r t of them w ill be s o al tere d th at the

“
o rg a ni s m s w hi ch g ro w u from th em w ill b e a bl e t o p res e nt th e

p ”
s a m e m o dific a t i on o n th e a ppli c a ti o n of a di f ferent s ti m u lus It .
i s t a ci tly as s u m e d by b oth th a t as i n th e h y poth es i s o f W ei s

,
m a nn s o i n th e c o u nt e r h yp oth es i s a p arti cula r p art of the

-
,
germ plas m gi v es origi n to a p a rt i cula r p a rt of th e de vel op ed

-
o rg a ni s m B u t n oth i ng of th e ki nd i s i m pli e d
. T h e n a ture of .
th e c ou nt e r h yp oth es i s ( a t a n y ra t e a s h e ld by m e) i s enti rely

-
m i s a pp reh e nd e d A ny one w h o tu rns b ac k t o th e ch a pt e rs i n th e

.
fi rs t v olum e wh ere th e c oncept i o n of p hy s i ologi ca l u ni t s ( or c on

s ti tu ti o na l u ni t s o r w h o re rea d s th e fo r e g o i ng
) w as s e t fo rt h ,
-
a pp en dix w ill s ee th a t th e re i s a l to g e th er e x c lu d e d a ny id ea o f
,
c orre l a t i on b e tw een c erta i n p a rts o f t h e g e rm a n d c e rt a i n p a rt s
o f th e re s u lt i ng org a ni s m The u n i ts are s u pp os e d t o b e a l l

.
a lik e , a nd d u ri ng th e p rog res s i v e em b ry ol ogi ca l c h a ng es l o c a l
grou p s of th em a re supp os ed t o t ak e on different fo rm s a nd

s tru ctu res u n d e r th e c om bi ne d fo rc e s g e ne ra l a nd l oc a l b roug h t
, ,
t o b ea r on th em T hi s concep ti o n i s n ec e s s i ta t e d by al l th e
.
e v id e nce T he fa ct di s cl os e d by th e e x p eri m e nt s of D ri esch

.
,
Wils on a nd C habry th a t from fra cti ons of an o vum s t ru ctu re s

, ,
m a y be o b t a i ne d li k e th a t obt a i ne d from th e w h ol e o vu m o nly ,
s m a ll er neces s i ta t e s i t
,
Th e fac t t h a t a ny s uffi c i ently l a rg e
.
fragm e nt o f a p ol yp o r pl a na ri a n n o m a tt er fro m w h a t p a rt of ,
th e b o dy t a k e n wi ll d e v el op i nt o a c o m pl et e p olyp o r pl a na ri an
,
nec e s s i ta t es i t Th e fac t th a t fro m a n u ndi ffe re nti a t e d p o rt i on

.
6 20 A P P E ND IX G .
of a pla nt e v en s o s m all a s a s cal e a com pl et e pl a nt m a y a ri s e

, ,
n e cess i ta t e s i t A nd i t i s n ec ess i ta t e d by th e fa ct t ha t a m ong

.
pl a nts roots a re p ro d uc e d by i m be dd e d s ho ots a nd s hoots by

,
ro ot s as well a s b
, y t h e fa ct t h a t l o w a ni m al s s u ch a s hyd ro id s , ,
i f d ep ri v e d of b oth h ea d a nd root w ill d e vel op a h ea d from th e ,
ro o t p a rt a nd a root fro m th e h ea d p a rt i f th ei r r e s p ec ti v e c on ,
di ti o ns a re i nv erte d A l l thi s e vid e nc e s h o w s c on cl u s i v ely t h a t

.
th e com p one nt u ni ts of e a ch s p e c i e s w h e th er e x i s ti ng i n th e g erm

,
o r i n th e d e v el op e d o rg a ni s m a re w h e n n o t , y et d i ffere nti a t ed by
,
lo c al c ondi ti o ns all a lik e a nd th at th e noti o n O f s p ec i a l p a rt s o f

, ,
t h e g e rm pla s m c orre l a t e d w i th s pe c i a l p a rt s o f the res ul t i ng o r

-
g a ni s m i s e nti rely a li e n t o th e hyp ot h es i s

, .
B ut h ow d o th e u n i ts o f a m o di fie d o rg a n a ffe c t th e u ni t s of
th e g er m i n s u ch w i s e th a t th es e p ro d u c e a n i nh e ri te d m o difica
t i o n of th e o rg a n w ill be a s k e d Thi s di ffi c ulty h a s b e en d ea lt
.
w i th i n 9 7d 9 7s wh ere th e a nal ogy b et w ee n t he s oci a l o rg a ni s m

, ,
a nd th e i ndi v id u a l o rg a ni s m h as b ee n b ro u gh t i n a i d : s er v i ng i f ,
no t t o fu rn i sh a c o nc e p t i o n y e t t o fu rni s h a n a d u m b ra t i o n
, Re .
gardi ng ci ti zens as th e u ni ts of a n u nfoldi ng s oci ety s ay a col ony , ,
i t w a s p o i nt e d ou t t h a t th e na tu re th ey i nh e ri t from a m o t h er
s o c i ety giv es th e m a p rocli v i ty t o w a rds a s o ci ety o f lik e s t ru e
t u re th e t ra i t s of w h i ch a re p rog res s i v e ly as s um e d as t h e
,
col o ny g ro w s s u f fi ci e ntly l arg e t o m a k e them p os s ibl e A t th e .
s a m e ti m e i t w a s p o i nt e d o u t t h a t w h il e t h e i nfl u e nc e o f th e
e nt i re a gg reg a t e o n th e i ndi vi d u a l s i s s ee n i n thi s fo r m i ng o f
th em i nto a s oc i ety o f t h e i nh e ri te d typ e th e i nflu enc es of l oca l ,
c i rcu m s ta n ce s a n d o f i ndi v id u al s o n o ne a no th er i n e a ch g ro u p
, , ,
m ak e th e m di f fe re nt i a te i nt o a pp rop ri a te s o ci a l s t ru ctu res t aki ng ,
o n fi t o c cu p a ti ons a n d i nd u s t ri es : t h e i m pli c a t i o n b e i ng th a t i n
vi rtue of th ei r i nh eri t e d natu res th ey a ll h a v e p a rti al cap a ci ti es

for t h e v a ri o us a cti v i t i e s t h ey u nd e rt a k e ; s o th a t a n i m m ig ra nt
cl e rk s ets u p a t a v ern a c o m p os i to r t a k es t o c a rp e nt eri ng a nd a
, ,
u ni ve rs i ty m a n rid es a fter c a ttl e o r i s e m pl oy e d o n a s h eep farm .
E v id e nc e w as gi v en i n t h a t pl a ce a s i n th e a b o v e p a rag ra p h t ha t
, ,
th e c ons ti tu ti o na l u ni ts o f a n org a ni s m s i m il a rly h a v e a l l of th e m

p ot ent i ali ti es for t aki ng o n t hi s or tha t s tru c tu re a nd m od e of
a c t i on w h i ch l o c a l c o ndi t i o ns d et erm i ne I t w a s furth er a rgu e d .
th a t a s c i ti z ens a re c onti nu a lly b ei ng rem ou ld e d by th e i r s o c i e ty

i nto c ongrui ty with i t a nd if ci rcu m s t anc es ch a ng e them t end t o
, , ,
rem ould th e i r s oci e ty ; s o i n th e i ndi v id ua l org a ni s m th ere i s t hi s ,
rec ip ro c a l a c ti o n o f th e w h ol e o n t h e u ni ts a nd o f t h e u ni ts o n
th e w h o l e H e nc e i t w as i nferre d th a t t h e m odifi e d u ni ts i n a ny
.
m o difi e d p a rt t e nd t o di f fu s e m o difi c a ti ons lik e th ei r o w n t hrou gh

t he u ni ts a t l a rg e : b e i ng a id e d by th e c i rcul a ti o n o f p rot opl as m ,
as s u gg es t e d i n 5 4 d a nd 9 7f A nd i t w as u rg e d th a t h o w e v er
.
,
i nconce i v a bly c om ple x s u ch a p roc ess m ay be yet i t s eem s not ,

F U NCTI ONA L L Y—CA U S E D MOD IF I CA TIONS .
621
i ncredible wh en w e recog ni s e th e p roba bili ty th a t a n orga ni s m i s

m or e or l ess p e rm ea bl e t o u n d u l a ti o ns p rop a g a t e d by i ts m ol e
c u l es : ROnt e n r a y s g i v i ng w a rr a nt I f s uch u ni ts t h r ou gh o u t th e
g .
t i ss u es m a y t a k e i n a nd s e n d o ut e th e re a l w a v es w h i c h b ri ng i t
i nt o rh y thm i c a l rela ti ons w i th o th ers of i ts ki nd a nd t e nd to pro
d u c e c ong rui ty i t b e com es if not c onc eiv a ble s ti l l s upp os a ble
, , ,
t h a t th rou g h out th e c i rc ul a t i ng p rot opl a s m th er e g oes on a c on

t i nu a l h a rm o niz a ti on o f i ts c om p o nents — a m ou ld i ng of e a c h by
a l l a nd o f a l l by ea ch S h ould i t be s a id th a t s u ch a p ro cess i s
.
t o o m a r v e ll ou s t o be rea s ona bly as su m e d t h e reply i s th a t i t i s ,
n ot m o r e m a r v ell ou s th a n he re di ty i t self w h i ch w e re i t n ot , ,
fa m ili a r t o u s w ou l d be t h ou ght i nc re di bl e
,
.
B ut as I hav e sa id
i n th e pla c e re fe rr e d t o A t l as t th e n w e
a re o b li ge d to a dm i t tha t th e a ctu a l org ani zi ng p roc es s tra ns cend s
c o nc ep t i on . It i s not e n o ugh t o s a y t h a t w e c a nnot k no w i t ; w e
m u st sa t h at w e c a nnot e v en c o nc e i v e i t c a n only co nc e i v e
y
th e p os s i bili ty o f a gges te d i nterp reta ti o n
su .
H en ce w e h a v e to r ely up on e v id e nces of oth er ki nd s A m ong .
th e s e s o m e w h i c h I th i nk di s p os e a bs olu t e l y o f t h e fa s hi o na bl e
,
h yp oth es i s w h ile t h e y h a rm oni z e wi th th e o pp os e d hyp oth e s i s ,
h a v e n o w t o b e n am e d T h a t th e i r i m pli ca t i o n s h ou ld not h a v e
.
b een g enerally rec og niz e d w ould h a v e s ee m e d t o m e i nco m pre

h e ns i bl e w ere i t not th at I h a v e m y s elf only no w o b s e r v e d th i s
i m pli ca t i o n The fa cts a re thes e
.
Ve r l ot g a rdene r wh o coul d di s ti ng ui sh 1 5 0 k inds of c a m el l i a ,

m ent i on s a
fl
wh en n ot i n ow e r ; a nd i t h a s b ee n p os i t i v el y as s erte d th a t th e fa m ous ol d
fl
D utch ori s t Voorhel m , who k e pt a b ov e v a ri e ti es of th e hya ci nth , w a s
h a rdl y e v er d ec ei v ed i n k nowi ng e a ch v a ri ety by the b ul b a l one Hence .
we m u s t conc l u de t h a t t he b ul bs of the hya ci nt h a nd th e bra n ch e s a nd

l e av e s of t h e ca m el l i a , t h ou gh a pp ea ri ng t o a n unp ra c ti s ed eye a b s ol u t el y
”
undi s ti ng ui s h bl e , ye t
a r e a l l f
f
y di e r
( D a rwin , Va r i a ti on of A n i m als a nd
.
P l a nts , t he , v ol i i , p . .
Mo re re ce ntly t e s t im ony to lik e effect h a s b e en gi v en by D r .
Ma x w ell Ma s t ers a nd h a s a l re a dy b ee n qu ote d by m e i n a not e

,
to 28 6 in ill u strat i on o f a no th e r t ru t h . He sa ys i g
c on c e r n n
s u ch v a ri a ti o ns
To t h e u ntr a i ne d
th e p rim ordi a l di ffe renc e s note d a re Often e ry
e ye , v
v
s l i gh t ; e e n th e b ot a ni s t , unl e s s h i s a tte nti on be s p e ci a l l y d i r e ct ed t o th e
m a t t er, fa i l s t o s ee m i nu t e d i ffe renc es w hi ch a r e p e rce p t i bl e eno ugh t o th e
rai s e r or hi s work m e n . Th e s e a pp a rentl y t ri i n m orph ol o i ca l fl g g
d i fference s a re oft en a s s oci a t e d wi th p hys i ol og i c a l v a ri a ti ons wh i ch rende r
s om e v a ri et i es , s a y of wh ea t , m u ch b et te r e na bl e d t o res i s t m i l d e w a nd di s
e a s e g enera l l y th an ot h e r s Som e , a ga i n, p rov e t o b e b et te r a dap te d for

.
cert a i n s oil s o r for som e c l i m a t e s th a n oth e rs ; s om e a re l es s l i a bl e to in u r

”
y j
from p red a t ory b i rds th a n oth ers , a nd s o on .
I n hi s Veg eta bl e Tera tol ogy, p 4 9 3 , D r Mas te rs nam es a noth e r . .
fact h a v i ng a lik e i m pli ca ti on— the fa ct th a t am ong s e e dli ng

6 22 A P P E ND I X G .
ks w h i ch h a v e not yet flow e red th os e

s to c ,
w hi ch w ill p ro d uce
d ou bl e flow ers a re di s ti ng ui s h a bl e H e s ay s .
Thi s s ep a ra ti on of th e from the dou bl e fl o were d pl a nts , M Ch a ti e

s i ng l e -
.
tel l s u s i s not s o d i ffi c ul t a s m i gh t b e s uppo s e d The s i ng l e s toc ks , h e e x

.
pl a i ns , h a v e de ep gree n l e a v e s ( gl a brou s i n ce rt a i n s p e c ie s ), rounde d a t th e

t op , th e h ea rt b e i ng i n th e form of a sh ut tl ecoc k, a n d t h e p l a nt s to ut a nd
thi ck-s e t i n i ts gene ra l a s p ec t, whil e th e p l a nt s yi e l di ng doubl e fl
owers h a v e
v ery l ong l ea ve s of a l i gh t green col our, ha i ry and curl ed at the edges , th e
he a rt co ns i s t i ng of whi t i s h l e a v e s , c urv ed so th a t they encl os e i t com
”
p l e te l y
.
W hat i s th e ge ne ra l truth i m pli e d ? C l early t ha t t here e x i s ts

no s u c h th i ng as a n i nd ep e nd ent l oc a l v a ri a ti on S om e m a rk e d
.
c h a ng e i n th e fo rm or c ol ou r o f a flo w er o r a fru i t d ra w s a tt en
t i o n ; a nd , b e i ng a ch a ng e wh i c h i nt e re s ts th e fl o ri s t or g ard e ne r,
p ec u ni a rily or othe rw is e , not o nly d raw s a ttenti on but u s u ally
m o nop oliz es a tt e nt i o n : t he na t ura l i m p res s i o n p ro d u ce d b e i ng th a t
thi s v a ri a ti o n s t a nd s t h ere by i ts elf— i s w i th out rel a ti on t o
v a ri a ti o ns el s e w h e re . Bu t no w i t t u rns ou t th a t th e re a re
c o nco m i t a nt v a ri a ti ons a ll o v e r th e pl a nt E ve n i n u nd er
.
ro u n d b ul b s c e rt a i n a pp re ci a bl e di fl ere nce s g o a l o ng w i th c e rt a i n
g
c o ns pi c uo u s differ enc es i n th e fl o w e rs A nd if a long w i th a
.
s triki ng ch a ng e i n a fl ow er w hi ch th e fl ori s t c o ntem pl a te s , th e re
o ch a ng e s a ll o v er th e pl an t no t o b v i o u s t o c a re l es s o b s er vers
g
but v i s ibl e t o h i m , w e m u s t i nf er th a t th ere a re e v e ry w h e re
m i nu t e di ffere nc es w h i c h e v e n t h e fl ori s t c a nn o t p erc e i ve : th e
w h ol e c o ns ti tu ti o n o f t h e pl a nt h as d i v erg e d i n s om e m e as u re
from th e cons ti tut i o ns of ki nd re d pl a nts E v e ry l oc al v a ri a ti o n
.
i m pli es a ch a ng e p erv a di ng th e e nti re org a ni s m , m a nifes ted i n

co nc om i t ant v ari a ti o ns e v e rywh ere el s e .
If s o wh a t b e co m e s o f th e hyp oth es i s o f d e t erm i na nts — th e

hyp ot he s i s th a t t h ere i s a s p e ci a l e l e m e nt i n th e g e rm pl as m -
w h i c h res u lts i n a s p ec i al l o c a l m o difi c a ti on i n t h e a d ul t org a ni s m ?

Th a t t h e re a re n o fa ct s s u pp ort i ng i t h a s be e n a ll a l ong m a ni
fe s t ; but n ow i t i s m a nife s t th a t th e fa c ts di rec tly c o ntra di ct i t .
A t th e s a m e t i m e i t m a y he rem a rk e d t h a t w h il e t he fa cts a re
w holly i nc ong ru ou s w i th th e h yp o th e s i s of d e term i na nts a nd i ts
a c c o m p a nyi ng e l a b ora t e s pec u l a ti o n, t h e y a re no t i nco ngru ou s
w i th th e a lt e rna t i v e hyp oth e s i s I m p oss ibl e though i t m ay be

.
t o i m agi ne t h e na tu re s o f th os e u l ti m a t e u ni ts p ecu li a r t o e ach

s p e c i es , w h i c h h a ve p ro cli vi ti es t ow ard s th e p a rt i cular fo rm
o f o rg a ni z a ti o n c h a ra c t e ri zi ng i t , ye t th a t a c h a ng e of s tru c t u re
a ri s i ng i n o n e p a rt of th e o rg a ni s m i s a cc om p a ni e d by m ul ti tu di
nou s ch a ng es o f s tru c tu re i n o th er p a rt s o f t he o rg a ni s m , i s no t
o nly c o n ru ou s w i th th e b eli e f th a t t h e re e xi s t s uc h c o ns t i tuti ona l
g
u ni ts , bu t yi eld s i t di s t i nct s upp o rt For i f, as a b o ve a rg u e d , a
.
c o ns pi c u ou s l oc a l v a ri a ti o n i s n ot th e res ul t of a ny m o difi c a t i o n o f
u ni ts s p ec i a l to th e l o c ali ty , bu t i s th e res ult o f a m o difi ca t i o n of

F U N CT I ONA L L Y C A U SE D M OD I FICA T I ONS-
.
6 23
th e un i ts at l a rg e
h a pp e n t h a t s u ch m o di fic a ti o n m us t
,
t h en i t m us t
ha v e i ts e ffe c ts o n a ll o th er p a rts of th e orga ni s m ; s o th a t th ere
ca nno t fa il t o re s ult al l th o s e s m a ll c o nc om i t a nt v a ri a t i o ns a b o v e
i ndi ca t e d .
May w e no t a sol les s i ncom p rehe ns ibl e

sa
y th a t i t beco m e s
th a t s tru c t ura l c h a ng e s ca u s e d b
di s u s e a re i nheri te d ?
y u s e a nd
I f as w e s ee a loca l v a ri a ti on sp ont a neou s ly a ri s i ng i s a cc om p a ni e d
, ,
by m u lti tu dinous oth er loca l v a ri a ti ons i m plyi ng a nec es s a ry ,
c o rre l a t i on b e t we e n ea c h l oca l v a ri a t i o n a nd t h e g e ne ra l c o ns ti t u
t i on o f th e org a ni s m ; t h e n i t m a y be a rgue d th a t i f a m a rk e d
c h a ng e of fu n cti o n i n a n o rg a n c a u s es i n creas e o r d ec reas e o f i t
,
t hi s g en e ra l c orrel a t i o n i m pli es th a t th e re m u s t be a r e c ip ro c a l re
a c ti o n b e t w e e n t h e p art a nd t h e w h ol e t endi ng t o r e —es t a bli s h
,
th ei r c o ng ru i ty The c o ns ti tu ti o n a t la rge w ill i n s o fa r he

.
c h a ng e d a nd a l ong w i th i t s c h a n e w ill o c orr e s p o ndi ng ch a ng es

, g g
i n th e s p erm c ell s a nd ge rm cel l s
- -
.
Fi na lly l et m e a dd not a noth er a rg um ent but a noth er fa ct Of

, ,
o b s er v a ti o n o f th e ki n d w h i c h o pp o ne nts d e m a nd
, bu t w hi ch , ,
w h e n t h e y a re fro m t i m e t o ti m e fu rni sh e d a re s e v e ra lly p o oh ,
p ooh e d as no t e nou gh E a ch o f th em i s s p ok en of a s a s oli t a ry

.
fa ct a n d s lighte d as i na d e q u a t e : a nd w h e n by a n d by a noth er i s - -
na m e d th i s i s t re a t e d i n t h e s a m e w a ; s o th a t th e fa ct s wh i c h
, y
if brou gh t tog eth er w ou ld be reco g ni z e d as s u ffi c i e nt a re ne ve r
brou gh t t og e th e r Tha t t o wh i ch I refer i s s et fo rth i n a p a m phl e t
.
by M L e o E rre ra P rofes s or at th e Uni ve rs ity O f B ru ss el s

.
, ,
e nt i tl e d H é rédi t é d u n C a ra ct er e a c qu i s c h e z nu C h a m pig no n
“ ’
pluri c ell ula i re be i ng a n a cc o u nt of e x p e ri m e nt s o f D r H u ng e r .

,
a t th e B ota ni c a l I ns ti tu t e i n B russ el s Fi rst e nu m erat i ng v ari ou s .
i ns t a nc es of a d ap tati ons to cli m at e a s th os e of pla nts w h i ch fitt ed , ,
t o north ern reg i ons p r es e r ve t h e i r co ns ti tu t i o na l ra pidi ty o f

,
ro wt h a nd s e e di ng w h e n brou ght s ou th a nd d o th i s fo r s e v e ra l
g ,
ge ne ra ti o ns h e goes o n t o d eta il th e c ult ure e x p eri m ents of

,
-
M H ung er a nd s u m s u p th e res u lts o f th ese i n t he follow i ng

.
,
w ord s
On d é dui t de l a q u e
° ’
1 L es coni d es d A sp erg il l u s n ig er s ont a da p t é es a l a c oncent ra ti on d u
i
m i l i eu on a v é cu l i ndi vi du q ui l e s p ort e ; c et e fl e t e s t en cor e p l u s m a r q u é
’ °
a p r é s d eu x g é n é ra ti on s p a s s é e s d a n s u n m i l i e u d onn é ( E xp é r I e t I I ) ; .
“ ° '
2 1 1 s a gi t d u ne v eri ta bl e a da p t at i on e t non p a s s i m p l em ent d un
’ ’
a c croi s s em en t d e v i ue u r c h e z l e s coni d i es p r o v en an t d e s l i q ui d e s c onc ent r é s ,

g
ea r ce s m em e s c oni di es g a rm en t m oi n s r a pide m e nt e t donn en t d e s p l a ntes
m o ns v i goureu s e s q ue l e s co n i di e s norm a l e s l o rs q u o n l e s s ei n e d e n o uv e a u
’
i
en s a da p t a nt a ux l i q ui des
’
s ur l e m i l i eu typ e
- concen tr é s , el l e s s e s ont
dé sadap tées da l i q ui de norm al ( E xp é r I I I) ; .
U ne g é n é ra t i on p as s é e s u r l e l i q ui de norm al n e ifa ce p a s l i u fl uence

° ’
3
d une o u de deu x gén é ra t i ons ant é ri eure s pa s s é es s u r u ne l i q ui de p l us c on
’
c e nt r é ( E xp é r IV ) . .
“
Tou s c e s r é s u l ta ts c oncor dent : il s m ontr ent u ne l eg ere, m a i s i ncontes t
”
ab l i i hé édit i e cl e l a da tat o a u m i l i eu
i
’
e t r a n s m ss on r a r n .
S U BJ E CT— IND E X .
( For th i s Ind ex a s i t a p pea r ed i n p re v i ous e d i ti o ns th e A u th or i s i nd e b t e d to F .
H O WA RD C OL L I N S E s q o f E d g b a s ton Bi r m i ng h a m , I t h a s n o w b e e n a d j u s te d
, , .
to s ui t t h e p res ent re vi se d a nd e nl a rg e d e d i t i o n .
)
A CA C I A, fol i a r or g a n s , II ,
4 1 , 2 64 .
g e n e s i s of e rt e b ra t e v s k ul l ,
II ,
A ca l ep h ce en v i ronm e nt , I , 1 05 ; 2 2 7 ; fa l s e oi n t s , , 3 71 j II , 3 72 ; c on
w a t e r i n, I , 1 73 . c e i v a b i l i t y of ri va l h y p oth e s e s , II ,
A ca ri
‘
: sp ecia l c re a t i o n a nd e fi e cts 61 8 a d a p t a t i on to en v i ron m en t

o f, I
4 2 8 ; d i r e c t t ra ns fo rm a t i ons ,
,
i n A s p er g i l l u s , , 6 23 II .
I , 706 ; s e g m e n t a t i o n , , 111 II . A c r o g e ns , t h e t e r m , II , 5 5 —6 .
( S ee
A cor u a ca l a m u s , agam ic p rop a g a A r ch eg oni a teaz ) .
t i o n, I , 642 . A c ti n op h r y s : a p ri m a ry a g g re g a t e ,
Ac q u i re d c h a ra c t e rs , i nh e r i t a n c e II 7 6 , g e ne s i s ,
, II 4 5 2 , .
of : f u n c t i o n a l l y p ro d u c e d m -
od i A c ti noz oa : m u l t i a xi a l d e v e l op m e nt ,
fi c a t i o ns i n p l a nt s a n d a ni m al s, I ,1 6 6 ; w a s t e a n d re p a i r , , 21 3, I
I , 3 07— 1 3 , 3 1 8 , 5 2 6 , 5 4 1 , 5 6 2 , 6 9 2 2 1 9 ; d i ffe r e n t i a t i o n , , 3 9 1 ; p a ra I
5, II
, 6 1 8 2 2 ; c o n c e i a b i l i t y o f,
-
v s it ism , , I
3 9 7 ; i n t e g ra t i o n , , 92 II
on h y p oth e s i s
th e of p h y s i ol og i s y m m e t ry , II , 1 89 , 1 92 ; g row t h
c a l u ni t s , , 3 68 —7 1 I , 69 5 , , 61 8 II a nd g en e s i s , , 44 4 II .
22 ; d i m i nu t i o n of jaw, I , 5 4 1 —2 , v
A c t i i t y : t h e p ri n c i p l e o f, th e es
693 ; c u r r e nt vi ew s o n, I , 5 5 9 - 6 0; s e nt i a l e l e m e nt i n L i fe , I , 1 1 3,
c e s s a t i on of s el e c t i o n , 1, 5 60—3 1 1 4, 1 22 ; n ot i n h e r e nt in l i i ngv
E im er s
’
t h e o ry of or t h og e ne s i s , m a t t e r, I , 1 2 0; n u t r i t i o n g e ne a nd
I , 5 6 0; s p eci es d i lI e r e nt i a t i on, I , si s, é
r s um é II
, ,
4 9 7 —9 ; a n d e vo l u
5 73 ; l o ca t i on of m a m m a lia n t i on, II , 501 —4 .
t es tes, , I
5 73 ; t a c t ua l p e rc e p t i e v A d a p t a t i on : g e ne r a l t ru t h s , 2 27 I ,
ne s s , , 6 02 I
—8 6 3 3 6 6 5 6 6 6 , 6 7 2—3 ,
, , , 3 3 , 2 3 3 —5 ; b ot a ni c a l , , 2 2 7 ; p h y s i o I
6 9 2 ; b l i n d ne s s o f c a e - a n i m v als, 1, l og i c a l , , I
2 2 8 s 3 3 ; p s y c h o l og i c a l .
6 1 2 —3 , 6 4 7 —9 ; c oa d a p t a t i o n Of co I , 2 29 , 2 3 0- 3 ; s t r u c t u r a l , f u n c
op e ra t i ve
p a rt s , , 621 , I 6 6 3 —5 t i ona l , a nd i nt e r d e p e n d e n c e , , I
t r a n s m i s s i on o f d i s e a s e , I , 6 22 —3 ; 2 4 0—1 ,
3 1 8 ; s oc i a l a nd or
h y p ot h e s i s s u p p or t e d b y t e l e g o ny , g a ni c sta bi l ity , ,
2 4 0—2 ; r I é s um é I , ,
1 , 6 2 4 —8 , 6 4 44 3, 6 4 9 —5 0; vi ew s of 2 42 —3 ; t o va ri e d m ed i a , I , 4 7 9 —8 1 ,
D a rw i n a nd n e o - D a r w i ni s t s , I , 489 , 5 5 6 : m u l t i p l i c a t i o n o f e ffe c t s ,
6 30 , 685 , 6 9 0; w h y fa c t s i n s up 1 , 5 1 2 —3 , 5 5 0; d i r e c t e qui l i b ra t i o n,
p or t a re m e a g re , I , 6 3 2 ; d e g ra d a I , 5 22—3 ; n a t u r a l s e l e c t i o n a n d
t i on of l ittl e t oe , I , 6 5 2—3
, 6 73 e q u i l i b r a t i o n, , 5 3 0 5 ; no n a d a p I - -
n e u t e r fo r m s o f s oc i a l i ns e c t s , I , t i ve s p eci fic c h a ra c t e rs , I , 5 65 ;
6 5 8 —9 , 6 6 3 —4 , 6 70, 6 75 ; d e g e n e ra t e d tim e re q u i re d fo r e ffe c t i ng , I ,
i n s t i nc t i n a nt s , , 6 60—2 ; r u d i I 565 6 ; -
an obs ta c l e to re -a d a p t a
m e nt a r y l i m b s o f W h a l e , , 6 69 , I t i o n,II 1 1 ; f s k i n a n d s k e i e
, o
6 9 2 ; i m p or t a n c e of u e s t i o n, ,
q I t o n II 2 1 5 2 1 7 ;
, , ter ti e II , ou ssu , ,
6 7 2 , 6 9 0; m o n s t ro u s d e e l op m e nt v 3 1 2 4 3 8 7 ; s k i n nd m
—
, m m a u co u s e
of h one y -
a nt s ,
“
I , 68 3 —4 ; os t e o l og y b a n e d i ff e nt i a t i on
r II 3 2 1 2 er , ,
—
of P j
un a b i s , I , 68 9 ; s u m m a r y o f 3 8 9 ; v a s c ul y t em II 3 4 3 4 ar s s , ,
-
e v i d e nc e s in s up p o r t , I , 6 9 2 —5 ; os s eo s II 3 5 2 ; m c u l a II
u , , us r, ,
6 26 S UBJ E CT —I ND E X .
3 6 8 —9 ,
3 9 1 ; p e r s i s t e nc e o f fo r c e t i o n, II , 24 4 , 246, 25 1 , 25 2 , 25 6,
a nd p h y s i o l og i c a l , , 39 4 ; II of re 2 72 , 38 5 —6 ; a d a p t a t i on of re p r o
p ro d u c t i e v a cti vit y t o c o n d i t i o ns , d u c t i ve a ct i vi ty t o c o n d i t i on s , II ,
II ,
41 1 -6 ; v e r t e b rae de v e l op m ent , 2 8 9 ; i n t e g ra t i o n, , 29 2 ; i n d e fi II
1 1 , 5 6 3 —6 .
( S ee a lso Co -
ad a p ta ni t e ne s s , , II
2 9 5 ; g e n e s i s a nd d e
t i on .
) v e l op m e nt , ,
463 II .
A fr i c a , c fi e ct of c l i m a t e o n i nh a b i A l i m e n t a ry c a n a l : m e t a b o l i c p ro
t a nt s , I , 30 . ces ses a nd a g e nt s , 1, 68 —9 , 74 ;
Agam og e n e s i s a l t e r n a t i on w i th s t r u c t u ra l t ra i t s , I , 1 9 2 ; p rog r e s
g a m o g e ne s i s ,I 2 6 6 —7 2 72—3 28 4 , , , si ve de v e l o p m e nt , I , 1 9 5 ; re l a t i o n
9 4 3 3 6 5 9 2 II 4 1 5 ; p a a l l l i m i n
, , , , r e s to en v i ro nm e n t I ,1 96 ; fu n c t i on ,
,
k y o k i ne s i s I 2 6 7 —8 ; a p o e
ar , ,
r c ss I 205 ; s e g m
, e n t a t i o n i n a nne l i d s ,
o f d i i n t e g a t i on
s I 2 7 6 —7 ; c nd i r , , o II 1 2 5 ; d i ffe
, r e n t ia t i o n, II , 3 01 ,
t i o n s d e t e r m i n i ng i t s c on t i nu a nc e , 3 02 , 3 2 1 —2 , 3 2 3 —5 3 8 9 ; s p e ci a l i za ,
I 2 8 4—9 4 29 5 —7 3 3 0;
, , , p h y s i o l og i c a l t i o ns i n b i r d s , , 3 25 ; i n ru m i II
u ni t s , I , 35 1 , II , 6 1 3 ; s p o nt a ne na n t s , , II
32 7 —9 ; d i ffe r e n t i a t i o n o f
ous fi s s i on I , , 5 8 2 , 5 8 4 —7 , 5 8 9 —9 2 , l i e r,v , 3 29 II
—3 3
; m us c u l a r i t y , , II
5 9 5 —6 , 5 99 ; re m a r k a b l e e x t e nt o f,
u nd e r fa ou r a b l e v c o n d i t i on s , I , A l l ot r op i s m Of or g a n i c c on s t i t u
5 9 1 —2 , 6 40 1 ; i n A c t i noz oa , ,
-
92 ; II e nt s , I
4 , 9 ; m u s c u l a r a c t i o n, , 5 9
, I .
i n H y d r oz oa , , 1 02 ; i n A nne l i d a , II A l l oy s , m e l t i ng p o i nt o f, , 3 39 I .
II , 1 03 ; i n n u t r i t i on , , 1 79 80 II -
. A l t e r n a t i on of g e ne r a t i on s , m i s l e a d
A g a r i ci nce , II 1 39 , , 25 7 . i ng a p p l i c a t i on of t e rm , II , 84 .
A ga s siz, L . J R . .
, z oo l og i c a l cl a ssi ( S ee A g a m og e n e s i s a nd Ga m og e n
fi c a t i o n, I 38 0
, . esis ) .
A g g r e g a t e s , A ni m al a nd P l a nt ( s ee A m i t os i s , oc c u rr e nc e o f, in m orb i d
M o rp h o l o g y ) . t i s sues, I , 2 64 .
A g i l i ty , v i ta l a t t ri b u t e I 5 78
a , , . Am m on i a :p ro p e r t i e s , I , 7 , 9 ; ne r ve
A g r i m o n y fl or a l s y m m e t ry II
, , , 42 , s t i m u l a t i o n, , 55 I .
1 67 , 1 70 . Am oeba : c e n t ra l de v e l o p m e nt I , , 1 63 ;
A i r, in veg eta l t i s sues , II , 5 6 7—8 , a p r i m a ry a g g r e g a t e , II 8 6 ; , sym
583, 591 , 593 . m e t r y of e n c y s te d , II 1 8 6 ; , sy m
A i r p l a nt s , , 2 08
”
I . b i os i s , , 400 II .
A l b u m e n : p r op e rt i e s , I ,1 2 ; L i eb e r A m p h i bi a : c l a s s i c a t i o n fi o f, I , 3 92 ;
k ii h n
'
s fo r m u l a , I , 1 3 ; d i ffu s l b i l e m b r y o n i c re s p i ra t o r y s y s te m , I ,
i ty , I
1 9 ; i n o rg a ni c t i s s u e s ,
, I , 41 . 45 7 ; s t ru c t u r e a nd m edi a, 483 ; I ,
A l c oh o l s , p rop e r t i e s , , 1 0—1 2 I . l i m b l o co m o t i o n, II , 1 5 ; s eg m e nt a
A l yce : r e p r o d u c t i o n, a nd the dy t i o n, II 22 5 ;
,
1 22 , ou t e r t i s s u e s ,
na m i c e l e m e nt in l i fe , I , 1 1 8 —9 1 1 , 3 1 1 ; r e s p i ra t i o n , II , 3 34 , 3 38
m u l t i c e n t ra l de v e l op m e nt I , , 1 63 , O w e n o n s e l e t o n, k II , 5 5 2, 5 5 7,
1 64 ; axi a l de v e l op m e nt I , ,
1 65
l ocom ot i v e p ow e rs of m i nu t e A m p h i oa u s : s e p a ra t i o n o f s e g m e nt a
‘
fo rm s, I , 1 96 ; u n i fo rm t i s s u e a n d t i on s p h e re s of egg , I , 69 1 ; cm
fu n c t i o n , I , 2 00, 5 8 6 ; g a m o g e ne s i s , b r y o g e ny , II 1 2 1 ; l oc a l s e g m e nt a
,
I , 2 7 1 , 2 7 9 , 2 80, 28 3 , II , 4 4 8 , 4 49 , t i o n, II , 1 2 5 —7 , 605 ; g e n e s i s O f
4 5 0; fe r t i l i t y , I , 582, II , 44 0, 4 4 1 v e r t e b ra t e a x i s II 2 1 3 6 , ,
-
, 21 8,
fi s s i on , I , 5 8 4, 5 85 ; u ni c e l l u l a r 2 22 ; d e v e l o p m e nt II 5 64 , , .
fo r m s, II , 22 ; i n t e g ra t i o n i n C on A m p h i p nous cu c h a v a s c ul a r ’
z ,
a ir
f e r voi d rce a nd C onj u g a tea’, II 2 5 ; , sacs, II , 33 7 .
p s e u d o fol i a r a n d a i a l
-
e x d v e l op A na ba s s ca nd ens , th e clim b i ng fi s h ,
m e nt , , 28 — II
3 3 , 5 7 ; fo l i a r d evel I ,
4 8 0, 48 3 .
o p m e nt , II 7 6 9 1 ; b ra n c h s y m
, , A na ch a r i s ( s e e E l oi d ca ) .
m e t ry , II 1 4 5 : c e l l m t a m
, e or A n ae s t h e t i c s , d i e rs e e fi e c t s v O f, I ,
p h os e s , II 1 7 6 ; t i e d i ff e nt i
, ssu er a s o.
S U BJ E CT —l ND E X .
97
A ng r aecum , a ssim il ati ve fu n c ti on of 4 90 .

( S ee a l so A nne l i d a a nd
roo t , II , 25 5 . A r th r op od a .
)
A ni m al S p i ri t s , v i ta l i s m a nd , I , A nt h r op om or p h i s m , fo r m er p re v a
115 . l en ce o f, I , 41 9 .
A ni m als : nu t r i t i o n a nd m ol e c u l a r A nt s : u t i l i z a t i o n of a p h i d s , I , 6 60
r e a rr a ng e m e n t , 3 6 —7 ; n i t ro g e n I , 1, II 4 03 , 4 05 ; n e s t - m a t e s ,
, II 4 05 ;
,
o u s c h a ra c t e r, , 3 9 — 4 1 ; s e ns i b l e I c a s t e s i n s oc i a l s p e c i e s , I , 6 5 8 —9 ,
m ot i o n , I
5 7 ; m et a b ol i s m ,
, , 62 I ~
6 7 0, 6 7 5 ; l o s s o f s e l f-f e e d i ng i n
7 7 ; m u l t i p l i c a t i o n o f e n e rg i e s , , I s t i n c t i n A m a z on s , , 6 6 0—1 , 6 6 3 I
75 ; c ont ra s t e d t ra i t s of
p l a nt s a n d , 4; m on s t r ou s v e l op m e n t o f H o n
de
I, 1 9 6 ; w ha t is a n i nd i i d ua l ? v,
I e y -a n t s , I 6 8 3 ; b u l k a nd t e c u n
,
2 4 6 — 7 ; s ol a r i n u e n c e, fl I , 5 00, 5 5 6 d ity, II ,
492 . ( See a lso T er m i tes .
)
g e o l o g i c c h a ng e s a fi e c t i ng , I , 5 01 A p hi s : i n d i i d u a l i t y , v I , 2 4 9 , 2 5 0; II ,
4 , 5 49 , 5 5 0, 5 5 6 ; i n t e r d e p e n d e nc e 603 ; p a r t h e nog e ne s i s , I , 2 7 4 —5 ,
w i t h p l a n t s , , 5 04 —6 , 5 1 4 , , 398 I II 289 ; fe r t i l i t y , , 58 2 , I 6 4 0—1 ; II ,
4 01 ; c o m p l e i t y o f i n u e n c e s a f x fl 4 7 6 , 4 9 0; uti l ize d by a nt s , I , 66 0
t e e t i ng , , I
5 06 ; g eog ra p h i c a l i s o ‘
1 ; II , 4 03 , 405 ; o e r - m u l t i p l i c av
l a t i on a nd o r i g i n of s p e c i e s , , I t i on ch ec k e d b y l a d y b i rd , ,
4 06 -
II .
5 68 9 ; -
i ta l v a t t ri b u t e s , I 5 77 9 ,
— A qua t i c a ni m a l s , l a rg e s i z e a tt a i n e d
d i s t r i b u t i on a nd a nt i q i t y fu o by , I , 1 56 .
p l a nt a nd a ni m a l typ es II 2 9 7 ;
, ,
A r a c h ni d a : a v o i d a nc e of d a ng e r , I ,
m utua l d e p e n d e n ce of o r g a ni s m s 92 ; o vi p a r ou s h om og e n e s i s , I , 2 71
a t l a rg e, II 3 9 7 —4 08 ; h y p ot h e t i c a l
, s e g m e n t a t i on , I , 4 69 ; II , 1 1 3, 1 1 4 ;
p l a n t -a n i m al typ e, , 3 9 7 ; p ro II i nt eg ra t i on a nd h om o l og y , II 1 1 1 , ,
g r e s s i v e i nc r e a s e o f s i z e , II 4 01 ;
,
1 2 1 ; b i l a t e ra l s y m m e t ry , II 1 9 8, .
laws of m u l t i p l i c a t i on, II , 4 1 1 —6 A r ce l l a : s y m m e t r y , II , 1 8 6 ; ou t er
rh y th m i n nu m b e rs , II 41 9 ; l a w
, t i s s u e d i if e r e nt i a t i o n, II , 3 09 .
of w e i g h ts a nd d i m e n s i o ns , , II A rch eg oni a teae: m or p h o l og i c a l c om
434 .
p os i t i o n , II , 3 2 —5 ; g r o w t h a n d d e
A ni m als, d om est i ca t e d : v a ri a t i on , v e l op m e n t , II
, 5 04 3 ; t ub u l a r
I , 3 24 , 3 26 , 5 60, 563, 69 3 ; i nt e r s t ru c t u r e , II , 6 2 ; a l t e r n a t i ng
58,
b r e e d i ng , I , 3 4 5 —7 , 3 54 , II , 61 5 ; g e ne ra t i on not d i s t i n c t i e , , 84 ; v II
p ure a nd m ix ed b re e d s , I , 3 54, a s y m m e t ry a nd en v i ro nm e n t II , ,
625 . 1 4 0; i n t e g r a t i o n , 29 3 , 2 96 ; i h II ,
A nne l i d a p h os p h o r e s c e n c e ,
: , 5 0; I d i vi d u a t i on a n d g e ne s i s , , 441 , II
a x v
i a l d e e l o p m e n t , , 1 6 5 , 1 66 ; i n I 45 1 , 463 .
I
t e g r a t i o n , , 3 6 3 ; l a r a l fo r m s a n d v A r c h e n t e r on : p r i m i t i v e e xt e rna l ity,
p h y l og e ny , , 4 4 7, , 1 1 5 ; seg I II II, 3 01 ; fo r m a t i on of c oe l o m , II ,
m enta l fl s s i on, , 5 8 8 —9 ; seg I 3 02 .
m e nt a t i o n , , II
9 8 —1 01 , 1 03 4 , 6 02 -
A r c h i a nne l i d a s e g m e n t a t i o n, II ,
5, ,
II 1 07 —9 ,
1 2 5 —7
; l a t e ra l g e m 1 25 .
m a t i o n, , II
1 05 ; e m b ry og e ny , II , A r eni col a m a r i na : p o l y t ro c h a l lar
1 1 9 ; b i l a t e ra l s y m m e t r y , , 19 II "
v ae , II , 1 09 .
2 00 ; g e n e s i s , , 44 4 , 4 5 3 II . A rm : em b r y o g e ny of h um a n, I , 1 69
A nnu l os a r e g e n e r a t i on , , I
3 61 2 ; -
v i ca ri ou s u s e o f, I , 2 09 .
d i s ti nc ti ve t ra i t s , I, 3 9 2 ; o r i g i n of A r m y , m or p h o l o g i c a l a n a l o g y , II , 6 .
t y p e, 11, 9 8—1 1 0, 602 —6 ; u n i t o f A r t e r i e s ( s ee Va s c u l a r S y s t e m ) .
c om p o s i t i on , , 1 05 ; a p p l i c a t i o n II A r th r op od a u ni a i a l d e e l op m x v e nt ,
of t e r m , 1 1 , 1 1 1 ; e r t e b ra t e s y m v I ,1 6 5 ; p rot op l a s m i c c on t i nu i t y , , I
m e t ry c o m p a r e d , , 2 03 —6 ; s e g II 1 9 0, 6 29 ; e c u r s i e ne s s , , x
481 ; v I
m e nt a l d i ffe re nt i a t i o n , II , 207 —9 ; l i m b l oc o m ot i o n , , 1 5 ; i n t e g ra II
u n i n t e g ra t e d fu n c t i o n i n P l a n a r i a , t i o n a nd h o m o l og y , , 1 1 1 —4 , 1 2 1 II
II , 3 73 ; d e e l o p m v e nt a nd g e ne s i s , b i l a t e ra l s y m m e t ry , II , 1 9 7 —2 00
II , 4 64 ; nu t r i t i o n a nd g e nes i s , II , g e ne s i s , II , 44 5 , 4 5 3 :
698 SU BJ E CT—I ND E X .
A s c i d i a ns : m ul ti a xi a l de v e l op m e n t , B a tr a chi a ( s e e A m p h i bi a ) .
I ,1 6 5 , 1 6 6 ; fu n c t i o n a l d i ff e re n t i a B e a n, va s cu l a r sy s tem ,
II , 5 73 ,
t i o n, , I
2 02 ; c om p os i t e i n d i i d u a l v 591 .
i ty of D ol i o l u m , I , 24 7 ; s e l f- fe r B ea ve r, ta il a nd c o- a d a p t e d s t ru c
t i l i z a t i on , I , 3 42 ; i n t e g r a t i on, II , t ure s , I , 61 6 .
9 4, 9 6, 97 ; s y m m e t ry , II , 1 94 ; B e e s ( s ee n s e c t s ) I .
o r i g i n of v e rt e b ra t e t y p e II , , 1 94, B eg oni a cee : m u l t i p l i c a t i on I , 2 24 ,

5 9 8 , 605 . 3 1 7 , 4 4 2 ; i n d i i d ua l i t y , v I , 25 1 ; d e
A s com y ce tes , r e p ro d u c t i on , II , 450 . v e l op m e n t fr o m s ca l es , I, 28 2
A s s i m i l a t i on : p a red w i t h rea
com s y m m e t ry , II , 1 59 , 1 66 ; de v e l op
s o ni ng , I , 8 1 —7 ; a t r a i t of i t a l i t y , v m e nt , II , 2 71 .
I , 5 77 . Ber ke l ey , M . J .
, i nd e fi ni t e n e s s of
A s ter oi d ea , ra d i a l s y m m e t ry , II ,
m os s e s a n d fe r n s , II , 29 6 .
1 96 . B i l e, a r re s t of x c re t i on I 2 09
e , , .
A s t ro no m y : g ro w t h of cel est i a l B i l i rub i n a nd b i l i v e r d i n e fu nc t i on ,
b od i e s , , 1 35 ; S c h l e i d e n I on i nd i o f, II , 3 3 0, 3 3 3 .
vi d ua l i ty , I , 245 ; e v ol u t i o n ,
I , B i ol og y : fi
d e ni t i o n a nd v
d i i s i on s , I ,
4 32, 4 35 ; cl a s si fi ca t i on of s t a r s , I ,
1 2 4 —5 ; o rg a n i c s t ru ct u ra l p h e no
4 44 ; rh y t h m o f, a nd o rg a n i c m e na , I , 1 2 5 —7 ; a l so fu nc t i on a l , , I
c h a ng e , I , 4 99 5 01 , -
55 7 ; law of 1 2 7 - 9 ; a c t i o n s a nd r e a c t i o n s o f
e q ui l i b r a t i o n, I , 5 1 9 —2 0; c oo p e r a fu n c t i o n a n d s t r u c t u r e , , 1 2 9 —30; I
t i on of s t ru c t u r e a nd fu n c t i o n , g e ne s i s , I , 1 30 1 ; l i m i t e d
-
k no w l
II , 3 . e d g e o f, I , 1 3 1 ; e ol u t i o n, v I , 4 3 2,
A ta i s m v : oc c u r r e n c e o f, I, 3 05 6 -
. 434 ; s oc i ol og i c a l a na l o g i e s .
( s ee
3 1 4 ; di g i t a l a r i a t i on , , v I 3 2 1 —3 . S oc i o l og y ) .
A t om s : u s e of t e rm , , 6 , 3 1 ; e t h e I B i op h o r s , W e i s m a nn s
’
g erm p l a s m
-
r e a l u nd u l a t i on s a n d os c i l l a t i on s , u ni t s ( 8 06 W e i s m a nn) .
I , 3 1 —5 . B i rd s : fl e s h - e a t i ng a nd g ra i n -
ea t
A us t ra l i a : s ett l er s
’
us a g es , I 3 64 ,
i ng c o n t ra s t e d , I
68 ; g r o w t h , a nd
ra t i o o f jaw to s k ul l in na t i v e s I , ,
e x p e nd i t u r e o f for c e , , 1 42 ; I size
5 41 . of egg a nd a d ul t , I , 1 44 ; l i m i t a
A x i l l a ry b ud s , or i g i n a nd de v e l op t i on s on flight I , , 1 55 ; s e l f- m ob i l
m e nt , II , 65 —8 . i ty , I
1 7 5 ; t e m p e ra t u r e ,
, I , 1 76
A xi s : n e u tr a l o f m e c h a ni c s , II ,
f u n c t i o n a l a n d s t r u c t u ra l d i ffe r
2 1 0; g e ne s i s of v e r t e b ra t e , II ,
e n t i a t i on , I , 2 01 ; food of s t a rv
21 2 -
6 , 22 4 —7 . i ng p i g e o n, I, 21 5 ; i i p a rou s vv
ne s s , , I
2 71 ; h e r e d i ty a nd p i g e on
B a c ter i a : fi s s i on I , , 2 70; n o n -n u b r e e d i ng , I , 305 ; a ta v ism in
c l ea te d, II , 2 0; ra t e of i nc re a s e , p i g e on, , I 3 1 4 ; o s s eo u s v a ri a
II , 443 . t i o n i n p i g e o n, I 3 21 ; c l a s s i fi ca
,
B a e r, K . E . v on : e m b ry ol og i c a l for t i on , 1, 3 92 ; m i g ra t i on s a nd
m ula, I , 1 71 , 1 72 , 45 1 , 453, 46 1 , c h a ng e o f h a bi ts , I , 3 99 , 4 02 , 5 00;

4 66 ; z o ol og i c a l cla ssi fi c a t i on ,
I ,
d i s t ri b u t i on i n t i m e, I , 4 1 0; D ar
383 ; on a ni m a l t r a n s i t i o ns , I ,
w in p e t re l s ,
on , 45 5 ; r ud i I ~
480 . m e nt a r y t e e t h , , 45 7 ; e r t e b rae, I v
B a l a nop h or a ’
, i nne r t i s s u e , II 2 7 4 , . I, 47 1 ; , II
5 6 4 ; fe a t h e r d e e l o p v
Bar k : va ri e d de v e l op m e nt II 247 , ,
m e nt , I , 473 ; h ab i ts of w a ter
9; p h y s i o l og i c a l d i ffe r e n t i a t i o n, ouz e l , I , 48 5 ; eg g sh el l s a nd di
II , 2 4 9 —5 0, 2 5 8 , 38 6 . re c t e q u i l i b r a t i on I , , 5 26 ; b one s of
B a s i d i om y cetcs , r e p ro d u c t i on , w a d e rs a nd d i re c t q
e u i l i b r a t i o n,
4 50 . I , 5 27 ; fe r t i l i t y a nd ne r vous de
B a t , i nfe r t i l i t y o f, II , 47 3 . v e l op m e nt , I 5 9 8 ; c el l ul a r con
,
B a t es, H . W .
, p ro t e c t i ve m i m i c ry t i nu i t y , I , 62 9 ; a d a p t a t i on of
of b u t t e r i es, fl I, 3 98 . s t ru c t u r e t o e n i r onm e nt , v II , 12 ;
S U BJ E CT—I ND E X .
6 9
se x ua l s e l e c t i on , II , 2 69 ; w i ng a d a p t a t i on s , I , 6 1 4- 2 1 , 6 74 , 677 ;
s p u rs , II , 313 ; ou t e r t i s s u e d i ffe r r u d i m e n t a ry l im bs of w h a l e, I ,
e n t i a t i o n, II , 3 1 4 —5 , 38 7 ; a li 66 8 , 6 9 2 ; i n h e r i t a nc e o f s e
68 5 ,
m e nt a r y c a na l de v e l o p m e nt II , , q u i re d m o d i c a t i o n s i n P u n a b i s , fi j
3 25 , 3 27 ; m u s cul a r c o l ou r a nd I , 68 9 ; s k ul l de v e l op m e nt II , , 222 ;
a cti v i t y II , , 3 65 —9
; nu t r i t i o n , , II t h eory o f s u p e r n u m e ra r y , II , 223 ;
433 ; c os t of g e ne s i s , , 43 6 ; II C op e o n o ri g i n of v e rt e b ra t e os s e
g r o w t h a n d g e ne s i s , , 4 54 , 4 5 8 II ou s s y s t em , II
2 2 5 7 ; d i ffe r e nt i a
,
-
x
h e a t e p e n d i t u r e a nd g e n e s i s , , II t i o n, II , 3 44—5 6 ; fa l s e oi n t s , , j II
4 68 - 9 , 4 7 4 ; a ct i i t y a nd g e ne s i s , v O w e n s t h o y of v e te
3 7 0—2 ;
’
e r r
II , 47 0—2 , 4 7 4 ; c on t ra s t e d m a m k e l e t on II 5 4 8 66
b ra t e s , ,
— .
m a l i a n fe rt i l i t y , II , 4 70; eg g s of B oo k w orm food

-
f I 77 , o , , .
wi ld a nd ta m 4 7 8 ; fe r t i l i t y
e, II, Bo n G r e x p e i m e n t s o n frog l a r
, .
, r -
of bla c k b i r d a nd l i nn e t co m p a r e d , vae I 3 65 , , .
II 503 ; O w e n
, on s k e l e to n o f, II , B o t n y b i o l og i c l c l
a , i fi ca t i n I a ass o , ,
55 9 , 560, 5 6 1 . 12 4 , 12 5 .
( S ee P l a nt s ) .
B i s c h ofi , em b ry og e ny of h um an B oth ri oc ep h a l us , de v e l op m e nt , II ,
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B i s o n, m od i fi c a t i on s e nt a i l ed by i h B otr y l l/i d w: d e v e l op m e n t I 1 66 ; l n , ,
c re a s e d w ei gh t of h ead, I , 512 . d e p e nd e n c e of c o m p on e nt s , , 24 7 ; I
B l a c bi rd ,k c ont ra s t e d w it h l i nn e t a g a m og e n e s i s , I , 64 1 .
in de v e l o p m e nt II 503 , , . B o w e r, P rof .
, o n a l t e r n a t i o n of g e n
B l a i n v i l l e d e d e fi n i ti o n o f
, , l i fe , I , e ra t i o n s , II , 84 .
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, ,
B l a s t u l a , d e ni t i on fi of l i fe a nd for 55 .
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B l oo d : sim i l a ri t y of i r on p e ro i d e , x e v o l u t i on I , 5 5 3 ; a n a l y s i s o f s ub
,
I , 17; m et a b ol i c p roc e s s e s , , 69 ; I s t a nce , I , 5 96 ; w eigh t l n h ig her

s e g r e g a t i o n of a b no r m a l c o ns t i t u a ni m a l s , I , 5 9 8 —9 ; s i z e i n c i i l i z e d v
e nt s , I , 1 8 0; p rot o z o o n l i fe o f c o r a nd u nc i v i l i z e d II , , 53 0 .
p uscl e s , ,I1 8 6 - 7 ; m or b i d c h a n g e s , B ra nch e s ( s ee M or p h o l og y ) .
I, 22 1 , 7 01 ; assim i l a ti ve p ow e r B r a nchi az ( s ee R e s p i ra t o r y S y s t e m ) .
a nd o r g a n i c r ep a i r , I , 22 1 - 2 ; re s B ra s s , e ffe c t o f a n t i m o n y o n , I , 1 21 .
p i ra t ory t i s s u e d i ff e r e nt i a t i o n, II , B rea d, d ia m a g ne t i s m , I , 3 70 .
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( S ee a lso Va s c u l a r Sy st em ) . a nd - i h , I , 344—7 , 35 3 ; II , 61 5 ;
B l o w -fl y , Weis m a nn on n ut r i t i o n p u r e a nd m i e d , x I , 3 54 , 6 2 5 .
a nd g ene s i s i n, I , 6 78 - 9 . k
B ri c s , c h a ng e d e q u i l i b ri u m s h ow n
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ra t e
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B ro w n-S e qu a rd , on i nh e ri t e d ep i
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m am m a lian cer vica l ve r t e b rae, I , B r y op h y l l u m , p e c u l i a r p rol i fe ra t i on,

394 ; e v o l u t i on a nd v e r t e b ra l c o l II , 29 5 .
u m n, I , 4 7 0—1 ; p a rt i a l d e v e l op B r y op h y ta , l a rg e s i ze a t t a i ne d by
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fl u e nc i ng , I , 5 1 2 , 5 3 6 - 9 ; d i re c t B r y oz oa , g e m m a t i on, , 58 8 I .
e q u i l i b ra t i o n a nd s t re ng t h , , I B u d d i ng ( s ee G e m m a t i on) .
5 27 ; na t u ra l s e l e c t i on a nd co B ud s : d e v e l op m e nt I , , 1 6 7—8 ; t h e
630 S U BJE CT—I ND E X .
c ri e s of h e red i ty a nd c a u l i n e, I , t i on a nd re p ro d u c t i ve fun c t i on,

3 58 —9 , 3 60 ; a x i l l a ry II ,
65 9 ;
,
-
ef 11 , 4 60 .
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fl
B u t t e r y : p ro t e c t i e v m i m i c ry , I , C a s t r a t i o n, e ffec t of, on g ro w t h , II ,
3 9 8 ; i n s t a nc e of t a m e, I , 68 4 .
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CA BB A G E v a r i e t i e s ,
o f, I , 3 02 . II, 4 9 3 -
6 .
C a c ta cece : fo l i a r a n d a a l d e e l op xi v Ca t a l y s i s, a nd vital m e t a m or p h os i s ,
m e nt, , II
4 7 9 ; d i ffe r e n t i a t i o n i n ,
-
I , 39 , 4 3 .
II ,
25 8 , 2 76 , 28 2 ; a s c ul a r s y s t em , v C a t t e l l , M c K e e n, on t a ct u a l p e rc e p
II 28 2 ; d y e p e r m
,
ea b i l i t y a nd ci r t i v e ne s s , I , 66 6 .
c u l a t i o n, , 5 71 , II 5 72 ; w oo d fo r C a ul er p a , s i m u l a t i on of h ig h er
m a ti on II 5 7 5, , 5 7 7 , 5 78 , 5 8 0 .
p l a n t -
fo r m s, II , 22 .
C a l l us , b u d d i ng fr o m , I , 3 58 , Ca v e a ni m a l s
-
, d e g e ne ra t i on of e y e s,
3 59 . I , 309 , 6 12 -3 , 6 1 4 , 64 7 9 , 69 3 -
.
Cam el, na t u r a l s e l e c t i o n a nd h um p C el l , the : i nc o m p r e h e n s i b i l i t y o f

o f, I , 5 34 . fo r c e s a t w or i n, , k
1 1 8 ; p ro t o I
C a n a d i a n s , F r e n c h , fe r t i l i t y o f, II , p i a s t s a nd t h e i r t ra i t s , I 1 81
,
t h e c e l l -t h e o r y , , 1 84 , 2 5 2 , I II 1 7 ,
C a nc e r , t h e d e fi ni t i o n of l i fe , I
, 1 1 1 ; 21 , 8 5 ; d i iT e r e n t i a t i on , , 1 88 —9
, I
( e s op h a g e a l , II , a nd v a s cul a r 1 94 ; t h e c o n t i n u i t y of p ro t op l a s m ,
sy s te m , II , 3 43 . I , 1 9 0 2,
-
1 94 , 6 28 - 3 0, II , 2 1 ; i ts
C a o u t c h o u c , l e a f s t r u c t u re , , 58 9 II . s t r uc t u r e , I , 2 5 3 —5 ; f u n c t i o n of
C a p i l l a r i e s ( s ec Va s c u l a r S y s t e m ) . c e n t ro s o m e , I , 25 4- 5 , 25 7 ; s t ru e
C a p i l l a r i t y , a nd e g e ta l a s cu l a r v v t ur e fu n c t i o n o f n u c l e u s ,
a nd I ,
s y s tem , , II
2 7 9- 8 0, 2 8 6 , 5 68 , 5 7 0, 25 5 - 6 , 258 —9 ; a ry o i n e s i s , , 25 7 k k I
5 8 5 , 5 8 7, 5 9 2 6 -
. 8 ; fu nc t i o n o f c h ro m a t i n , , 25 9 I
C a rb o h y d r a t e s : i ns t a b i l i t y , , 1 0; I 6 5 ; f e r t i l i z a t i o n a n d fu n c t i o n of
the t erm h y d ro -
c a r b o n, ih p ol a r b o d i e s , , 2 6 6 8 ; t h e o ri e s I -
of
m o l e c u l a r c h a ng e s i n , I , 42 - 3 ; or h e re d i t y b a s e d on t h e o ry , I , 356 ;
i Ve i s m d i ffe r e n t i a t i o n i nt o
’
g a ni e t ra ns fo r m a t i o n, I, 43 , 48 ; a nn s
'
m e t a b ol i c p roc e s s e s , I , 63 - 7 7 , r e p ro d u c t i ve a nd s om a t i c, I , 622 ,
2 6 2 —3 , , 3 62 II . 628 —3 0, 63 3 —4 4 ; n u c l e u s a b s e nt or
C a r b on : p rop e r t i e s , I 3 —5 , 2 0; c o m
,
d i s p e rs e d , 20, 8 5 ; II , m o rp h o l o g i
p o u nd s , I , 6 , 7, 9 , 1 0 1 2 , 1 3 24 —5
-
, . c a l d i ffe r e nt i a t i o n, II , 1 75 - 7 ; a ni
C a rb o n i c a cid ( c a rb o n d i o i d e) x : m al m o rp h o l o g y , II , 22 8 —3 0; m or
p ro p e r t i e s , , 6, 7, 9 ; I in a ni m a l p h o l og i c a l s u m m a ry , , 2 33 ; II
a nd p l a n t func t i o n s , I , 62, 2 1 4, ve g e t a l t i s s u e d i iI e re n t i a t i on , 1 1 ,
II , 3 98 : d i ffu s i b i l i t y , , 33 1 II . 2 49 - 5 0, 38 6 ; a s cul a r v de v e l op
x
C a r b o n i c o i d e , p rop e rt i e s , , 6 I . m e nt , II , 2 7 9 - 8 4 , 38 9 .
v
C a r ni o r e s : ni t rog e n o u s food , I , 4 7, C e nti p ed e , b i l a t e ra l s y m m e t ry , II ,
68 ; k a t a b o l i c p ro c e s s I 7 1 ; re 1 9 8 2 00 -
.
, ,
s tri ct e d e n v i o n m e nt I 3 96 ; r , ,
Ocp h a l op od a : b i l a t e ra l s y m m e t ry ,
t h e i r b e ne fi c i a l e ffec t s o n a n i m a l II 2 03 ;
,
v
a s cul a r s y s t e m , , 3 41 II .
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C a rp e nt e r, W . B . : on fu nc t i o na l C cr e us , t i s s u e d i iI e r e n t i a t i o n, II ,
s p e c i a l i z a t i on, I , 2 08 ; re p r o d u c 2 7 6 , 28 3 .
t i on of s e a -w e e d , I
, 582 ; eg e t a l v C e s a l p i no , I , 3 77 .
ce l l m u l t i p l i c a t i o n, I , 585 ; s t ru e C es tod a ( s e e E ntozoa ) .
t u re a nd m u l t i p l i ca t i o n o f c om C haatop od a , s e g m e nt a t i on, II , 98 ,
p o u nd o rg a n i s m s , I , 5 8 6 —9 ; on 1 03 , 6 05 .
f und a m e nt a l t ra i t s o f s e x , 595 ; I , C h a j a , w i ng s p u rs , , 31 3 II .
nu t ri t i ve s y s te m v
o f i n e rt e b ra t e s , C h a ng e , a nd fi
d e ni t i on of l i fe , I ,
I , 5 9 5 ; ll . a c r ocy s t zs ,
’
II , 4 50; nut ri 8 1 9 0, 1 1 3
-
.
S U B E CT I N D E X J -
.
31
C h a rl e s , R . H .
, on i nh e r i t a n c e of o l og i c a l , I , 38 0— 9 ; i nc om p l e t e
ac q ui re d m od i fi c a t i on s in l eg e q u i v a l e nc e of g rou p s , , 38 9 , I
b o ne s of P un a bi s , j I , 68 9 . 4 4 5 —6 , 44 8 , 555,5 7 2 ; g rou p at
Chatie , o n s i ng l e a n d d ou b l e s t oc ks , t ri b u t e s , I , 3 90 3 ; t h e t r u t h s
-
ln
II , 6 22 . t e rp re t e d , I , 3 9 3 —4 ; e t h no l og i c
C h e m i s t ry : p ro p e r t i e s of o rg a n i c a nd l i ng u i s t i c v
e o l u t i o n, , 44 1 - 6 I '
e l em e nt s , I , 3 —5
, 20, 22 ; o f d i a org a n i c e v o l ut i o n , I 44 3 4 4 7
, , ,
t om i c com p ou nd s , , I 7 1 0; t r i a
-
555 ; di fl
’
e r e nc e s in k i nd a n d d e
t om i c , , I
1 0—1 2 ; p o l y a t o m i c , , I g re e , I , 44 4 —6
; a nt e c e d e nt s t r u e
1 2 1 3,-
2 5 ; t r a i t s of e o l u t i o n, , v I t u ra l s i m i l a ri t y , , 44 7 , 4 4 8 —9 ; I
23 4 ; - e t h e rea l u nd u l a t i o n s a nd Vo n B a er
’
s fo r m ul a , I , 451 —4
,
a t om i c os c i l l a t i on, I , 31- 6 ; ch em i 555 ; o r g a ni c, n ot uni s e ri a l , II ,
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x C l a s s i fi ca ti on o f t he S c i ences , T h e,
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o rg a n i c d e v e l op m e nt a nd d i ffe r e n Co -
a d a p t a t i on o f c oo p e ra ti v e p a rt s
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ca l u ni t s , I , 22 5 , II
6 12 ; , p ri m i 51 1-3, 5 1 4—5 ; s l ow op e r a t i o n of
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e v o l u t i o n o f o rg a n i c c o m p ou n d s , a n a l og y , , I
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C h e s t nu t , l eaf s y m m e t ry , II , 1 49 , a n a l og y co n t i nu e d , 17
) the ca s e
1 53 . of b i s on
’
s h ea d, I , 5 1 2 ; n a t u ra l se
C h i ton : s i m u l a t i o n o f s e g m e n t a t i o n, l e c ti on an i na d e qua t e e x p l a na
II 1 1 6 , 1 1 8 ; s y m m e t ry ,
,
II , 2 02 . t i on , I , 5 35 , 6 1 4—2 1 , 69 2 ; R om a ne s
C h l o r o p h y l l : fu n c t i o n , I , 65 , II , on
“
c e s s a t i on of s e l e c t i on as
2 63 ; n u t ri t i o n a n d a b s e nc e o f, II , e ffe c t i n g , I , 5 60, 5 6 1 —2 ; Weis

7 4 ; c o n s t i t u t i o n, II
,
2 2; sy m bi m a nn s
’
t h eo ri e s , I , 5 6 0- 3 , 6 6 3 —5 ,
ot i c p re s e n c e in a nim a l s, II , 4 00 . 6 7 0, 6 7 4 5 ; na t u r a l s e l e c t i o n a n d
-
C h ond r a c a n th us g i bbos us , e no r m o u s e c o n o m y o f g ro w t h , 5 62 ; p h y I ,
de v e l op m e n t of r e p ro d u c t i ve sys s i o l og i c a l p roc e s s e s i n o l e d , v v ,
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’
C h o r d a ta , a fii n i t i e s , I , 466 . a rt i c i a l fi s e l e c t i o n, , 61 5 ; w hat I
C h ro m a ti n ( s ee C e l l ) . a re c oo p e ra t i ve p a rt s
? I
61 6 7 ;,
-
C i rc l e , th e, a nd e v ol ut i on h y p ot h
“
i n t ra - s e l e c t i o n e x a m i ne d , ,
I
es i s, I ,
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-
a nd fe r t i l i t y , II ,
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4 9 4- 5 . C oc c o s p h e r e s v i t a l p rob l em p re
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g e nes i s , II , 5 29 - 3 1 . 1 1 9 ; i m b ri c a t e d p l a t e s , I , 1 82 .
C l a d op h or a : i nt e g ra t i on , II , 25 ; k
C oc r o a c h , ou s t i n g of E u ro p e a n
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J
S U B E C T —IN D E X .
C ael enter a ta r u d i m e n t a ry c o n t ra e t i on, II

2 6 7 9 ; s e u a l s e l ec t i o n ,
,
-
x
tile o rg a n s , 58 ; I
i t a l c h a ng e s
, v ’
II , 2 69 ; a c t i i t y a n d m u s c u l a r , v ,
II
i n p ol y p , I, 9 5 ; a i a l d e e l o p m e nt ,x v 365 -
9; p h y s i ol o g i c a l u ni t s a nd
I , 1 65 , 1 66 ; e n i ro n m e n t a nd v m i x t u re o f, in o ffs p r i ng , II , 61 6,
s t ru c t u r e , I , 1 7 3 ; s e l f- m o b i l i t y , , I 61 7 .
1 75 ; , II
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I , 2 7 9 , 28 0; II , 2 5 , 28 ,
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( S e e A l g ae ) .
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, 28 1 ; d i ffe r e n I C on j u g a t i on i n A l y ce ,
, , 2 79 ; i n P r o I
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’
t i on, II , 9 1 ; t e r t i a r y a g g re g a t i on , fi c a t i o n, I , 1 89 .
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1 9 2 - 3 ; s e g m e nt a l d i ffe r e n t i a t i o n , s i o n, I , 3 07 .
II 2 07 ; p h y s i o l og i c a l d i ff e r e n t i a
, C o -o r d i n a t i o n of a c t i on s ( s ee L i fe ) .
t i o n i n H y d ra a n d a n a l og y , ,
II Cop e, E . D .
, on o ri g i n o f v e r t e b ra t e
3 00; c i l i a t i on o f b l a s tu l a , II , 3 01 s t ruc t ur e, II , 22 5 —7 .
t i s s ue r e d u p l i c a t i on, II , 3 01 —2 , C or m op h y ta : s l igh t i nt e rn a l dit

f er
3 8 9 ; o u t e r t i s s u e d i ff e r e n t i a t i o n, e nt i a t i o n , II , 273 ; v a s c ul a r sys
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m e nt g e ne s i s ,
a nd II 4 6 2 ; , nu t ri C ow w h a t p ro m p t s h e r t o m
: um bl e
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-
.
3 02 3 -
. C ra b ( s e e C r us ta cea ) .
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d i fi us i bi l i ty , I,
1 8 21 ; -
o rg a n i c , I ,
1 54 , 29 2 .
21 , 25 , 26 ; p l i a bi l i ty a nd elas C roo kes , Sir W h y p ot h et i ca l

t i ci ty , I , 27 ; c a p i l l a ry a fii n i t y , I , ch e m i ca l u ni t p ro t y l e , I , 22 , 23 .
28 ; i s om e ri s m , I , 5 9 ; i ns t a b i l i t y , C r uci fer az, fl o ra l s y m m e t r y , II , 1 64 ,

I , 3 5 0: m ol e c u l a r m o b i l i t y a nd d i f 1 71 .
fu s i b i l i t y , II 3 3 1 ; i ns t a b i l i t y o f,
, C r us ta cea : l oc o m ot i on of l ob s t e r , , I
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6 1 ; a nd m u s c u l a r t i s s ue, II 3 6 1 —4
, . 3 60 , 5 89, II . 76 : h om og e n e s i s , I ,
C ol o ni e s , a u t og e no u s d e v e l o p m e nt 2 71 ; g e ne s i s a nd n u t r i t i on in
a nd p a ra l lel i n h e red i ty , I , 3 66 D a p h n i d az, , I 29 0—1 ; g ro w t h a n d
8; , II
62 0 .
g e ne s i s , I , 29 2 ; d e g e ne ra t i on o f
C o l ou r : s e ns a t i on o f, I , 54 ; p h e no e ye i n ca v e - i n h a b i t i ng , I , 3 09 , 6 1 4 ,
g a m i c, II , 75 , 2 65 6 ; l i g h t a n d -
64 8 ; h e rm i t c ra b p a ra s i t e , I , 39 7 ;
Ve g e ta l , II , 26 1 - 2 ; ora l fe rt i l i z a fl c h a ng es of m e d i a , ,
4 01 , I 48 1 - 2 ;
S UBJ E CT—I ND E X .
633
r e t ro g ra d e de v e l op m e nt in c i r ri c r os s i n g I 3 4 7 , 6 1 1 , 66 9 ; h i s t h e
,
p edes , I , 458 ; s e g m e nt a t i o n , I , o ry of p a ng e n e s i s e a m i n e d , , x I
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h o m o l og y , II , 1 1 1 —4 , 1 2 1 , 603 ; h i t i on a nd n a t u ra l b a r ri e r s , I ,
4 02 ,
l a t e ra l sym m e t ry , , II
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h e rm i t c ra b , , 3 2 2 , 3 8 7 ; fe r II t ri b u t i o n in tim e a nd s p a ce, I ,
t i l ity , II
4 5 3 ; n u t r i t i o n a nd g e ne
,
4 1 0; l i ng u i s t i c c la ssi fi ca t i on . I ,
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-
I , 1 8 —21 ; or m e nt of C i r r h i p ed i a , I 45 8 ; j a w s
,
g a ni e , , 2 1 2 , 2 6 I -
. a nd l eg s of C r us ta cea 4 71 ;
, I ,
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s t or m g l a ss , , 96 ; g r o w t h , , I I t i on s of sp e ci e s i n G a l a p a g os
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22 1 , 22 3 ; e q u i l i b ra t i on , I , 33 7 D a rw i n a nd L a m a rc k I 491 ; , , th e
p h y s i ol og i ca l p o l a ri t y , ,
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e a n d fo r m a t i o n , , 77 II . 53 0; i n d i re c t e q ui l i b ra t i on by
C ten od r i l us , s e g m e nt a l i nd i v i d u al na t ura l s e l e c t i o n, , 5 3 0-
5 ; in I
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,
t e rs , I , 5 65 ; foo d o f b l o w - fl y l a r t erm n a t u ra l s e l e c t i o n , 6 09 , , I
v aa, I , 6 7 8 ; a r t h r op o d s e g m e n t a 69 5 ; c a s t e g ra d a t i o n s a n d a w s j of
t i o n, II , 114; e g g - p r od u c t i o n of v
d ri e r a n t s , , 65 8 ; a t t a c h m e n t I of
C o ng e r , II ,
425 . cl im b i ng p l a nt s , II ,
2 7 6 —7 ; ve g et a l
Ca ttl e fi s h , -
i nd i i d u a l i t y v of H ec t o fr u c t i fi ca t i o n , II , 29 4 ; e a r th
co ty l u s , I , 25 0 . w o rm , II , 402 ; a ni m a l s t e ri l i ty
v
C u i e r, z oo l o g i c a l cl a s si fi c a t i on I , , a nd d om e s t i ca t i on , II , 4 8 0, 483 :
38 1 . v a ri a t i o n i n h y a c i n th a nd c a m el
C y a no g e n, p ro p e r t i e s , , 7, 9 I . l ia , II , 621 .
C y c l i ch th y s , d e r m a l s t r u c t u r e , II , D a rw i n , D r . E .
, m od i fi a b i l i t y of or
3 06 .
g a ni s m s , I , 4 90, 4 9 2 7 -
.
D ea t h : a n a r re s t of v ita l c orr e
D A L YE LL , Sir J .
, r e g e ne r a t i on in s p o nd e nc e , 1 02 ; o n l y I , lim it to
D a s y eh o ne, I 3 6 1 ; p ro p a g a t i o n
,
of v e g e t a l g ro w t h , , 153 ; I c e s s a t i on
H y d ra , II , 4 76 . o f c oo r d i n a t i o n o f a c t i on s , I , 5 78 ,
D a p h m d az,
‘
h e t e r og e n e s i s a nd nu t r i 5 79 ; W ei s m h y p ot h es i s ,
a nn s,
’
I
t i o n, I , 290 1 -
. 63 6 - 8 ; p h y s i o l og i c a l i nt e g r a t i o n ,
D a rw i n, C . : O ri gi n of S p ec i es , I , II , 3 7 4 , 39 2 ; c a u s e of n a t u r a l , , II
1 29 , II
5 28 ; na t u r a l s e l e c t i on a n d
,
4 1 3 ; re l a t i o n t o b i rt h s , , 41 7 II .
fu n c t i o n , , 3 08 - 9 69 3 : a t a i s m ,I , v D e fi n i t e ne s s : of v i t a l c h a ng e I , , 87
I 3 1 4 ; os s eo u s a r i a t i on s i n p i g v 90, 1 06 , 1 09 ; d e v e l op m e n t a l I
“
, , ,
eo ns , ,
I
32 1 ; p l a n t a ri a t i o n a nd v 1 78 ; fu nc t i o n a l , I , 21 2 ; s e g re g a
d om e s t i ca t i on, , 32 5 ; s p o nt a n e I _
t i on of e v o l u t i on , I , 5 1 4- 6 .
ous v a ri a t i on , I , 32 8 , 69 7 ; fl ora l De fi ni t i on , d i fii c u l t i e s of, I , 78 ; II ,
f e r t i l i z a t i on , I , 340 , II , 1 68 , 2 6 7 , 17 .
4 07 , 6 08 ; i nt e rc ros s i ng a n d s el f D e g e ne r a c y , m orp h o l og i c a l ob s c u ra
fe rt i l i z a t i o n , , 3 4 4 , 3 4 5 ; i nt e r I t i o ns d u e t o , II , 1 2, 1 3 .
6 34 S U BJ E CT —I ND E X .
D end r obi u m ( s ee O r c h i d s ) . D i ffu s i o n , of c ol l o i d s a nd c ry s t a l
D es m i di a ceaz: u n i c e l l u l a r, II , 21 ; l oi d s , I , 1 8 - 2 0; II , 331 .
l i ne a r a n d c e n t ra l a g g r e g a t i o n , II , D i g e s t i on a c t i on of ni t ro g e n ou s
23 ; n a t u ra l s e l e c t i on a nd sym c om p ou n d s , , I 69 ; obes i ty , II ,
m et ry , II
1 3 4 , 1 3 5 ; m o r p h o l og i c a l
,
4 8 0—4 ; fe r t i l i t y , II , 514 .
d i fi e r e n t i a t i o n, , 1 77 ; t i s s u e , , II II D im o rp h i s m : fl o ra l I , 5 34 ; s e u a l ,
, x
2 44 ; g e n e s i s , II , 4 4 0, 4 4 9 . i n p a ra s i t e s , I , 3 1 5 ; s oc i a l i ns e c t s
D e t e r m i na n t s , Weism a nn s
’
g e rm ( s ee In s e ct s ) .
p l a s m uni t s ( s ee G e r m p l a s m ) -
. D i nos a u r s , s i z e o f, I , 1 39 .
v
D e e l op m e n t : an i n c re a s e of s t ruc D i p h y es : i nd i i d u a l i t y , v I , 2 46 ; sym
t u re , I , 1 62 , II
4 6 1 ; p ri m a ri l y
,
m e t ry , , 1 92 II .
c e nt r a l , I , 1 6 2 , 1 6 6 ; u n i a nd m u l D i s ea se : s e g r e g a t i on of b l oo d c on
t i - c e n t ra l , I , 1 63 —4
, 1 6 6 —7 ; a i a l , x s t i t u e nt s , I , 1 79 ; c h a ng e s in
I , 1 64 , 1 6 7 ; u ni a nd m ul t i - a xi a l , b l o o d fro m , I , 2 2 1 , 7 01 ; h e r e d i t y ,
I , 1 6 5 - 6 ; a c h a ng e t o c oh e r e n t I , 3 06 —7 , 31 2 3,
6 22 4 ; b e l i e f i n -
de fi n i t e h e t e rog e n e i ty , I , 1 6 7 — 7 0, s u p e r n a t u ra l o r i g i n, , 4 1 9 ; p a ra I
1 7 9 ; Vo n B a e r s fo r m u l a , , 1 7 1 —2 ;
’
I s i ti s m a nd s p ec a l i c re a t i o n, I ,
i ndi i d ua l vd i ffe re n t i a t i on f ro m 42 7 ; m orb i d p ro d u c t s a s s p e c ific

en v i r on m e nt I , , 1 7 2 -
8 ; c e l l fo r m a c h a r a c t e rs , I , 5 67 ; t e l e g o ny , I ,
t i o n, I2 25 ;
, d i s c o n t i nu o u s , a n d 64 6 ; d e r m al s t r u c t u re , 1 1 , 3 06 ; l n
agam og e n e s i s , I
2 75 ; P ro f
, . H ux t e s t i na l m u s c u l a r h y p e r t ro p h y , , II
l ey
’
s cl a ss i fi c a t i o n, , 276 ; I s oci o 3 25 ; i nd i g e s t i o n a nd a l i m en ta ry
l og i c a l p a r a l l e l t o a u t o g e n ou s , 1 , c a na l de v e l op m e nt II , , 3 28 ; a u n j
3 64—8 , II , 6 2 0; r e t r og ra d e , , 45 7 I d i ce a nd b i l i ru b i n, II , 3 3 0; l oc a l i
8 : i ne q u a l i ti e s a m o n g c o -o p e r a t i ve z a t i o n of e x cr e t i o n II , , 331 ; m em
p a rt s , , 61 7 ; I
h e t e ro c h r o ny , , I b ra n e s in i nfl a m m a t o ry , II , 3 43
6 5 ; c o n t i n u o u s a nd d i s c o n t i nu o u s
5
o s s e ou s d i ffe r e n t i a t i o n i n ri c kets ,
v eg eta l ,
II , 52 ; s u m m a ry of II , 3 52 ; fa t t y d e g e n e ra t i o n , II ,
p h y s i o l og i c a l , II , 38 4 9 0; -
n u t ri 482 .
t ion a nd g e n e s i s , r es u m e, , 49 7 II D i s i n t e g ra t i on, p h y s i ol og i ca l ( s ee
9; e v o l u t i on , , 50 1 -
II
5 ; c om m e n c e P h y s i o l og y ) .
m e nt g e ne s i s ,
of II , 5 06 ; of ve r D i s tom a : m e t a g e n e s i s , 1 , 2 7 3 - 4 ; d i s
t e b ra t e l i m b s , II , 553 .
( S ee a ls o i n t e g ra t i o n o f g e ne s i s , , 2 76 ; I
M u l t i p l i ca t i on ) . c y cl e of
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D evel op m en t H y p oth cs l s , T h e, I , 41 7 . D i s t r i b ut i on : p h y s i c a l l i m i t s , , 3 9 6 I
D i a l e c t s ( s ec L a n g u a g e ) . o rg a ni c en v i ro n m e nt , I , 39 6 8 -
D i a l y s i s , a nd d i iT u s i b i l i ty , I , 1 9 , 20 .
p a ra s i t i c c o nd i t i o ns , I , 3 9 7- 8 ; s im
D i a sta se, d e c o m p os i t i o n o f, I , 38 , u l t a ne i t y of ag e nc i e s a ffe c t i ng ,
40 .
I ,398 ; m u t ua l e n c roa c h m e nt s of
D i a tom a ceaa: t i s s ue, II , 2 4 4 ; g e ne s i s , s p ecie s, , 3 9 8I-

4 01 , 4 7 7 , 4 8 9 ; fa c t s
II , 4 40, 448 . d i s p ro v i ng p re a d a p t a t i o n t o h a b i
-
D i a t om i c c om p ou nd s ( s ee Ch em ta ts, I , 40 1 —3
, 4 1 1 —2
: o f a ni m a l s
i s t ry ) . a nd p l a nt s i n t i m e, I , 4 04 - 1 1 , 4 1 2
D i c o t y l e d o ns : g ro w t h , , 1 3 9 , 1 43 , I o u s t i ng o f na t i ve s p ec ies i n Ne w
II , 6 3 —4 , 6 9 —7 2 , 78 , 8 2 - 3 ; u n i a i a l x Z e a l a nd I 4 7 7 ; , , l oca l fl ue nc e s in ,
de v e l op m e nt , I , 1 65 ; s tem a nd I 4 77 9 48 9 ;
,
-
, t h ro u g h v a r i ed
l e a f fu nc t i o n s , II , 2 5 7 ; m e c h a ni m edi a I 4 7 9 —8 5
, , , 48 9 , 556 ; past
c a l s t re s s a nd w oo d for m a t i o n, II , a nd p r e s e n t o rg a ni c fo r m s , , I
2 7 7 ; g ro w t h a n d g e n e s i s , , 45 1 II . 4 8 5 - 9 , 5 5 6 ; com p l e o rg a n i z a t i o n x
D i ffe re nt i a t i o n ( s ee M o r p h o l og y a nd a nd , II , 29 6—7 .
P h y s i ol o g y ) .
v
D i i s i on of l a b our, p h y s i ol og i c a l
D i fl l ug i a : p r i m a r y a g g re g a t e , II , 86 ( s ee L a b ou r) .
7 ; s y m m e t ry , II 1 8 6 ; o ut e r
, t i s s ue D og : c o n t ra s t e d li ves of t or t o i s e
d i f e r e nt i a t i o n , II 3 09 ,
. a nd , I , 1 03 , 1 04 ; i nh e r i t e d h a b i t s ,
S UBJ E CT -
IND E X .
63 5
I , 3 09 , 5 73 ; a b nor m a l d ig i ts, I , E g g s ( s ee E m b r y o l o g y ) .
3 24 ; i n t e r b r e e d i ng o f di ve rg e n t E i m e r, T , t h e o r y o f . or t h o g e n e s i s ,
v a ri e t i e s I , 5 6 5 ; d e c re a s e o f j a w ,
, I , 5 63-4 .
I , 61 5, 69 3 ; t e l e g o ny , , 6 45 ; c on I E l a sm obr a nch i i : p rotop l a s m i c c on

d i t i o ns a fl e c t l n g fe r t i l i t y ,
‘
, 4 74 , II t i nu i t y , I , 6 2 9 ; s e g m e nt a t i o n , , II
4 79 . 1 26 .
D o h rn , t h eo ry of v e rt e b ra t e s t ru e E l e c t ri c i t y : g e n e s i s i n o r g a n i c m a t
t u re , II , 606 . t e r, , I 5 0—2 , 60; m u s c u l a r s e
D ol i ol um , c om b i na t i o n of i nd i i d u v t i o n, I , 59 ; i nc o m p r e h e n s i b i l i t y ,
a l i ties, I , 24 7 . I , 1 21 .
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D ou b l e d a y , E o n n ut r i t i o n o f g e ne 4 5 9 , 5 06 ; c e r e b ro- s p i n a l s y s t e m , I ,
sis, II , 5 1 0- 2 . 5 9 8, 599 .
D ri e s c h , s e p a ra t i o n of s e g m e n t a t i o n E l k, I ri s h , h or n s a nd c o r re l a te d
s p h e re s of E ch m us o vum I , , 69 1 ; p a rt s , I , 5 3 7, 6 74 .
II , 618 . E l o i d ca c a na d ens i s : v
i nd i i du a l i ty , I ,
D r op s y ( s ee D i s e a s e ) . 2 48 ; e no r m ou s aga m ic m u l t i p l i ca
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l i t e ro ns g r o w t h , II , 75 . E l o n g a t i on , a nd l oc o m ot i on in a ni
D u B o i s R e y m ond , -
E . H .
, el ec m al s, II , 15 .
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D um a s , a nt i t h e s i s of a ni m a l s a nd i ni t i a l a nd fi na l o r g a n i c b u l s , , k I
p l a nt s , , 62 I . 1 4 3 , 1 5 8 , 1 6 1 ; foet a l fl es h c on s t i t u
D w a rfs , H i nd u fa m i l y o f, I , 316 . e nt s , , I
1 54 ; h u m a n a rm de v e l op
m e nt , I , 1 69 ; Vo n B a e r s
'
fo r
E AR , de v e l op m e n t of v ert e b ra t e II , , m ul a, 1, 1 7 0—2 , 4 5 1 —4 4 66 : . em
3 1 8 , 32 0 . b r y on i e h e a t , I , 1 77 ; s p h erica l or
E a rt h , c l i m a t i c r h y t h m a nd o rg a n i c g a ni c fo r m , I, 1 77 ; u ni t - l i fe in
c h a ng e , I , 4 9 9 —5 01 , 557 . m u l t i c e l l u l a r org a ni s m s , I , 1 8 5- 6 ;
E a rt h - w o r m : b i l a t e ra l s y m m e t ry , fu nc t i o n a l d i ff e r e n t i a t i on , I , 2 03
II , 1 99 , 2 00; m ou l d p ro d u c t i o n, 11 , i nd i vi dua l i ty I , , 2 46- 7 : u ns p e c i a l
402 . i zed r e p r od u c t i ve t i ss ue, I , 279

E ch i nococc us ( s e e E nt ozoa ) . 83, 3 1 7 ; c h a n g e s fo l l o w i ng i m
E ch i nod e r m a ta i n d e p e n d e nc e of p r e g na t i o n , , 2 8 3- 4 : nu t r i t i on a n d I
b l a s t o s p h e re c e l l s , , 1 8 5 ; p ro t o I v eg e t a l g ro w t h I , 28 5 - 8 , 29 5 - 7 ;
plasm ic c on t i n u i t y in em b ry os , I , a nd a ni m a l g ro w t h , I , 28 9 9 4 , -
1 90; s e p a ra t i o n of s e g m e nt a t i o n 29 5 —7 ; p h y s i ol og i ca l uni t s a nd
s p h e re s of o vum I , 69 1 ;
, II , 618 ; h e re d i t y , , 31 7 9 ; I -
v a ri a t i o n a nd
s y m m e t ry , II , 191, 1 9 5 -6 .
p a r e nt a l f u n c t i o n a l c o n d i t i o n , I ,
E c o no m y : of g ro w t h i n n a t u r a l s e 3 2 4 ; u t e r i n e e n i r o nm e n t , , 3 2 7 —8 v I
l e c t i o n, I , 5 3 6 , 5 6 2 ; a t ra i t of o r p h y s i o l og i c a l u n i t s a n d a r i a t i o n, v
g a n i e e o l u t i o n, , v
501 , 5 04 II . I , 33 0—4 , 4 58 ; fe r t i l i z e d a n d u n
E c t od e r m : fu n c t i o n a l d i ffe r e nt i a fe r t i l i z e d o v a , , 3 4 0- 1 ; h e rm a p h I
t i o n, I
2 02 , 2 03 ; fu n c t i o na l
, v i c a ri ro d i s m , I , 3 41 2 , -
3 44 ; s o c i o l og i
o u s ne s s , , 2 09 ; r e p ro d u c t i I ve fu nc ca l p a ra l l e l , I , 3 6 6 —8 : e vo l u t i on
t i on , I , 28 1 . h y p oth e s i s , I , 434, 43 6, 45 3 , 454 ,
E if e c t s , M u l t i p l i c a t i on of : v a ri a 5 5 5 ; p e t r e l d e e l op m e n t , , 4 55 v I
t i on, 1 , 3 29 -3 0, 3 3 3 ; o rg a n i c e vo s u b s t i t ut i on a nd s u p p re s s i o n o f
l a t i on , , I
5 1 1 —4 , 5 1 5 , 5 1 7 , 549 , o r g a ns , , 4 5 6 -
8 , 4 I
66 , 4 7 2 -
3 ; s t ru e
55 7 , , 4 0 5 -6
; II m o rp h o l o g i c a l de t u ra l p r oc l i i t i e s o f p h y s i ol og i c a lv
v e l op m e nt , , II .
7 —9 , 234 ; ph y sio u ni t s , I , 4 58 ; a bri d g m e nt of
l o gi c a l d i ffe r e n t i a t i on, II , 39 0- 1 , s t a g e s, I , 4 58 —9 , 4 64 : d i s a p p ea r
39 2 . a nc e of i n t e rm edi a te fo rm s , , I
63 6 S U BJ E CT -
IND E X .
4 5 9 - 60, 463 ; p re -
a d a p t a t i on , , I 5 84 ; nu t ri t i on a nd g e ne s i s , 64 1 I ,
461 - 3 ; d i s c r i m i na t i o n o f s p e c i e s II , 4 88 ; d i re c t t ra n s fo r m a t i o n , ,
I
i n e a rl y 4 6 1 ; a n o m a l ou s
s t a g es , I , 7 06 ; i n t e g r a t i o n, II , 1 02 ; s eg
p e r s i s t e n c e of a n c e s t r a l t r a i t s , , I m e n t a t i on, II , 1 07 ,
1 08 ; i n t e r d e
4 6 3 5 ; p h y l og e ny , , 4 66 ; e g g - s h e l l
-
I p e nd e n c e a n d o rg a ni c i nt e g r a t i on,
I
f u n c t i o n, , 5 2 7 ; g e ne s i s o f g r a d e s II , 4 04 .
in s oc i a l i ns e c t s , I , 6 5 4- 6 , 65 8 9 , -
E nv i ro n m e n t : d e g re e of l i fe a nd
6 7 9 - 8 0; d e t e rm i na t i o n of s e x, I , c om ple xi ty o f, 1, 1 04 —7 ; r e l a t i on
657 ; o rd e r of de v e l op m e nt q u a l i to o rg a ni c s t ru c t u r e a nd fu n c
fi e d by ne ed s, , 6 7 9 ; o s t e o l og y o f I t i on , I
1 72 —8 ;
, , 1 2 - 5 ; a d a p t a t i on II
P un a bi s, j
, 68 9 ; d i r e c t t ra n s for I to v
a ri e d m e d i a a n e i d e nc e o f v
m a t i o ns a n d p h y s i o l o g i c a l u n i t s , v
e o l u t i on , , I
4 79 —8 1 , 5 5 6 ; i n u e nc e fl
1, 706 : t ra n s fo r m a t i on of bia s of s ol a r s y s t em , I , 5 00, 556 ; ln
t em a, II , 2 0; a r re s t o f g ro w t h a n d h e ri t e d a d a p t a t i on t o, II , 6 23 .
i nnu t r i t i on, II , 73 : d e e l op m e n t of v E ol l s , b r a nc h i ae, II 1 1 8 , .
s e g m e nt e d a n i m a l s , II , 1 00- 2 , 6 02 ; E p i d erm i s ( s ee S k i n) .
a da p ti v e v e r t e b ra t e s e g m e nt a t i o n, E p i lep sy : d e n i t i on fi of l i fe a nd
II, 1 1 8 2 3 , 1 2 4 , 2 2 3 —4 , 605 - 6 ; a n i
-
m o v e m e nt s i n, I , 1 1 2 ; h e re d i t y , I ,
m a l c e l l m o r p h o l og y , , 2 28 ; p r i II 312 .
m a r y d i ffe re n t i a t i o n s o f g e r m i n a l E p i t h e l i um : ci l i a t ed, I , 1 8 7 ; H ert

l a y e rs , II 3 00 2 ; l ng d e v e l op ,
-
u w i g s c l a s s i c a t i o n,
’
fi I , 1 89 ; r e p ro
m e nt II 3 3 3 —4 ; m a m m a l i a n o a
, , v d u c t i ve fu n c t i o n, I , 2 80; pa ve
m at a t i on
ur II 34 2 3 : m o v m nt s , ,
— e e m e nt a nd c y l i n d e r, II , 2 29 .
of o II 3 5 6 3 6 3 ; m d i fi c a t i ons
va , , , o E p i zoa : d i s t r i b u t i o n , I , 3 98 ; s p ec ia l
i n m o l e II 3 9 1 ; g n i s a nd n
, , e es u c r e a t i o n a nd e f fects o f, , 4 28 : i n I
t i t i o n II 4 24 4 2 5 ; fi s h o a
r , II
, , v , , t e r d e p e nd e n c e a nd o rg a n i c i nt e
4 35 4 5 4 ; c
, t of g e n i II 4 35 os es s , , g ra t i o n, II 404 ; n t i t i , u r on a nd
6 ; n m b e of b i d
u g g s II 4 54
r r s
’
e , , g e ne s i s , II 48 7 , .
6 4 7 8 ; h e a t a n d g n e s i s II 468
, e , , , E q u i l i b ra t i n : v a i a t i on a nd
o r law o f,
4 74 : a c t i v i t y nd g e n e i s i n b i d s a s r ,
I , 3 26, 334 ; m ol e c u l a r a r ra ng e
II 4 7 0—2 4 7 4 ; v t e b a t e l i m b
, , er r m e nt , I , 3 3 7- 4 5 ; of o r g a ni c
d v e l op m nt II 5 5 3 ; os i fi c t i o n
e e , , s a c h a ng e , I , 3 4 8 , 54 7 , 557 ; d i re c t
i n ve t b te s II 5 5 6 ; O w e n s a nd i nd i re c t , I 5 1 9 - 22 , 5 73 a d ap
’
r e ra , , , :
v e t e b t e t h y II 5 6 3 ; d v l
r ra e or , , e e t a t i on by d i re c t , I , 5 22 - 3 , 55 1 ,
op m e nt o f v t b II 564 ( S er e rae , , . ee 5 57 : nu t r i t i o n , d e fe nc e , a nd fe r
a l so M u l t i p l i c a t i on ) . t i l i z a t i on of
p l a nt s , I , 5 2 3-5 ; d i
E m b r y o l o g y o f c o n c e p t i o ns , 45 1 , I . re c t of a ni m a l s , I , 5 25 - 8 , 5 5 1 ,
E m i g ra n t s , typ e of o rg a n i z a t i o n 5 57 ; n a t u ra l s e l e ct i o n a nd i nd i
a m o ng , 3 64 , ,6 20 I II , . re c t, , I
5 3 0 4 , 5 5 2 , 5 5 7 ; o f na t u ra l
-
E nd o d e r m : fu n c t i o n a l d i ffe re n t i a s e l e c t i on , 54 ,3 —7
, 5 5 2I—3 , 557 ;
t i on, I
2 02 , 2 03 ; fu nc t i ona l
, v i c a ri i nc r e a s i ng i m p o r t a n c e of d i re c t , I ,
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E n d og e n, a p p l i c a t i on of t e rm , II ,
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( S ee M onoc ot y l e d o ns ) . no t a cc o u nt e d for b y , I , 5 73 ; t i s
E ne r g y : e v ol ut i on o f, in a ni m a l s , I ,
s ue d i ffe r e nt i a t i o n , 245 ; g e ne II ,
7 1 —7 ; o rg a ni c
g ro w t h a nd e x p e n s i s o f n er o u s s y s t e m , v , 3 07 - 8 II
I
d i t u r e , , 1 41 ; f u n c t i o na l t r a n s fe r. fu nc t i o na l , , II
39 1 - 4 ; l a w s o f m u l
I, 2 01 —6 ; c h ro m a t i n a s t h e s o u rc e t i p l i c a t i o n, , 4 1 1 -6
II
; in hum a n
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. a nd s oc i a l ev l o u t i o n, II , 53 7 .
( S ee
( S ee a ls o F or c e ) . a lso Ac q u i re d c h a ra c t e r s a nd
E ntoz oa : m e t a g e ne s i s , I , 2 7 3 , 64 1 N a t ura l s e l e c t i on .
)
s e l f- fe rt i l i z a t i o n, 3 42 ; d i s t r ib u I ,
E rre r a , Li nh e r i t e d a d a p t a t i on
.
, on
t i o n, I , 3 9 8 ; a n d s p e c i a l c re a t i on. to v
e n i ronm e nt i n A s p erg i l l us , , II
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S U BJ E CT - I ND E X .
637
E t h no l og y : h e re d i t y , I , 3 03 4 , 3 1 0
-
n e i t y a nd d e fl n i t e ne s s o f, I , 5 1 4 —8 ,
p l a s t i c i t y of m i xed ra c e s , I , 3 54 5 5 0; n a t u r a l s e l e c t i o n a nd g e n e ra l
v
p ri m i t i e i d e a s , I ,
41 7 ; e v o l u t i on d o c t r i ne o f, 1 , 5 4 3 - 8 , 5 5 7 ; fa c t o r s
a nd c l a ss i fi c a t i on I , , 44 1 3 , -
4 46 ; t a b ul a t ed , I , 5 5 1 ; i nd uc t i e e v i v
n a t u ra l s e l e c t i o n , I , 553 . d e nc e s s u m m a ri z e d , 1 , 555 6 ; -
s ur
E u p h or bi a cece : fo l i a r a nd a xi a l de v i v i ng d i s b e l i e f i n F ra n c e , 5 59 ; I ,
ve l o p m e nt , II
4 7 —8 ; p h y s i o l o g i c a l
, c u r r e nt t h e o ri e s of o rg a n i c , 1,
d i ffe r e nt i a t i o n , , 2 58 ; d y e p e r II 5 5 9 —7 4 ; E lm er s
’
t h eo r y of o rt h o
m e a b i l i t y a nd c i r c u l a t i o n , II , 5 71 ; g e ne s i s , , 5 63 —
4 ; I
Gul ic k on
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58 9 —9 2 , 5 96 . or i e s , , 5 7 3 I
-
4 ; i no r g a n i c a n d t h e
E v a p ora t i on : org a ni c c h a ng e I , , 28 ; S y s tem of P h i l os op h y , I , 69 6 ;
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E v o l u t i on : c h e m i c a l e l e m e n t s a nd 7 01 , 7 02 ; d i s s ol u t i o n a nd p rob l e m s
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, or d i a l o f m o rp h o l o g y , , 4—6 ; m or p h ol II
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og y a nd fo r m ula , II , 7 9, -
23 1 - 5
1 81 ; II
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; , , fi I d i fli cul t i e s of fi
d e ni t i o n, II , 17
1 07 - 1 0; g ro w t h t h e p r i m a r y t ra i t c e l l - d oc t r i ne , II , 1 7 - 21 , 8 5 ; u ni c e l
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r s um é of p l a n t m or p h o l og y , , II
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de , ,
-
, 7 8 - 8 0; o ri g i n a n d d i ffe r e n t i a t i o n
1 7 8 —9 ; p rog e s s i v e s t r u c t u ra l d i f
r of p h ae nog a m i c t yp e, II 83 ; ,
fe r e n t i a t i o n, I , 1 81 -
4 , 1 9 2—6 , 2 1 1 p h y s i ol o g i c a l p ro b l e m s , II , 23 9 4 3 -
2 ; l i fe b e for e o r g a n i z a t i o n, I , 2 1 0; t i s sue d i ffe re n t i a t i o n , II , 2 44—6,

h e t e rog e ne i t y of fu n c t i o n , I , 21 1 38 5 ; i nt e g ra t i o n o f o rg a n i c w or l d ,
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v
5 1 8 ; i nd i i d u a l i t y , , 2 47 ; c e l l o r I -
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w i th s p e c i a l - c r e a t i on h y p ot h e s i s , e nc i ng h um a n, II 5 2 5 8 ; f t e
,
-
u ur
I , 41 5, 4 3 1 —40 ; d e ri v a t i on of hy of p o p u l a t i o n, II 5 3 2 7 ; s e l f s uf
,
— -
p ot h e s i s , , I
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, r r ,
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, 43 9 ; ex I II , 563 6 -
.
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, 4 3 3 5 , 4 3 9 ; d i r e ct e i d e n c e ,
-
v 3 03 ; l oc a l i z a t i on of, , 33 1 3 II -
.
1 , 4 3 5 - 7 , 4 39 ; m a l e v o l e nc e not i m x
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p l i ed by , , 43 7 - 9 ; I e v i d e n c e from ( S ee D i c ot y l e d on s ) .
c l a ss i fi c a t i on , I , 4 43 , 44 4 , 44 9 , E x p e nd i t u r e ( s ec M u l t i p l i c a t i on) .
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v
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,
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D a rw i n on
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-
a ri a
’
v m o l og y o f p e t a l s , II , 4 3 —6 ; sym
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.
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,
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w or k I 2 1 5 —6 ; g ro w t h a nd l n
, ,
it recu r ? I , 29 4 —7 , 3 3 6 - 4 0; e fi ec t
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d e p e n d e nc e o f s o c i a l a n d o rg a n i c , t i o n, II , 39 ; a l t e r n a t i ng g e ne ra
I , 2 3 7—9 , 2 4 0- 2 ; s t ru c t u r e a nd t i on i n l i ve r w o rt s , II , 8 0—4 ; m o l
h e r ed i t y , 3 1 8 9 ( s ee A c I , 3 06 —1 3 ,
-
l u s c a n h om og e n e s i s II
, , 1 1 6, 1 1 7
q u i re d C h a ra c t e rs ) ; a i d s na t u ra l 8; v e rt e b ra t e II , 1 1 8 ; g ro w t h ,
, , II
s e l e c t i o n, , 3 08 ; o r g a n i c i nt e r d eI 2 66 .
( S ee a lso F e r t i l i z a t i on, a nd
p e n d e nc e , I , 3 1 8 9 ; p a r e n ta l c o n
-
M ul t i p l i c a tion ) .
d it ion a nd v a ri a t i o n , , 3 24 , 32 6 I G a s ter op od a ( s e e M ol l us ca ) .
va ri a t i on a nd a l t e re d , I , 3 25 —6 , Geddes a nd
T h o m p s on , o n t h e de
3 33 -4 ; as ca u s i ng v a ri a t i on I , , t e r m i na t i on o f s e x , , 65 7 I .
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c a t i on of ef
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s t ru c t u r a l c oo p e ra t i o n , II , I ,
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3 , 21 7 v i c a ri o u s v e g et a l , II , 2 70 G e n e s i s ( s ee M u l t i p l i c a t i o n) .
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a rr a ng e m e n t , II ,
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-
G e n u s : i nd e fi ni t e v a l u e I , , 38 9 , 44 6 ;
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i ns t a b i l i t y of h om og e n e o u s a nd G o ri l l a , c a l l os i t i e s , II , 31 2 .
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, B i r ds o f A us tr a l i a , II ,
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’
x e nt s w i th
4 04 - 1 1 , 412 ; s p eci a l c r e a t i o n, I , fro g l a r ae , , 3 65 v I .
41 9, 426 ; e vol ut i on I , ,
432, 43 7 ; Gra h a m , T p r op e r t i e s o f w a t e r ,
.
, ,
I
re c ord c o n g r u ou s w i th e vo l uti on , 9, II , 3 5 9 ; c o l l oi d s a n d c r y s t a l
1 , 4 8 5 - 9 , 5 5 6 ; o rg a n i c i n u e n c e o f fl l oi d s , I , 1 5-8 , II , 3 56 ; t h ei r d if
c h a ng e s , , I
501 3 , 5 4 9 , 5 5 0, 5 5 7 ; -
fu s i b i l i t y , I , 1 8 - 2 0; sa pid a nd ia
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v
e ol u t i on, I 5 65 - 6 ; r i s e
, of i ns e c t 2 63 ; ora l fl s y m m e t ry , II , 1 65 ;
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m a n e o l u t i on v a nd c h a ng e s , II , 25 7 .
534 . G r a m i n i v o re s , food c o nt r a s t e d with

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1 , 4 3 3- 4 . G ra s s i , on foo d h a b i t s of T er m i tes ,
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G e rm -
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, v eg e t a l c i rc ul a
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G erm -
plasm , W eis m a nn s
’
t h e o ry o f, G ra v i ty .
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of l os t l i m b s , 3 62 ; I , v a r i a t i ons in s y m m e t ry , ,
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, 1 3 5 - 7 ; s i m u l a t i on o f, I , 1 3 6 ; l i m i ts
of v
a r i a t i o n s i n n e u t e r i ns e c t s , , I t o, , I
1 3 7 , 1 5 5 —7 ; s t r u c t u ra l c om
65 9 , 6 63 - 5 ,6 7 5 ; c or r e l a t e d 671 , p l e x i t y , 1 , 1 38 —4 0, 1 4 5 — 7 , 1 6 1 ; nu
v a r i a t i on s i n s t a g , , 6 7 7 ; i ns u p e r I t r i t i o n, , 1 4 0, 1 4 7 —9
, I 1 6 1 ; e xp en
a b l e d i f cu l ti e s, , fi
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6 0, 1 61 ; na l fi a rr e s t o f, 1,
va ted p l a nts , II , 621 - 2 . 1 49 —5 5 , 63 9 ; w h e re u nc e a s i n g , I ,
G h os t t h e o r y , -
Vi t a l i s m a nd , I , 114 . 1 54 ; r s u m é
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G i r a ffe , c o - a d a p t a t i o n o f s t ru c t u re s , I , 1 6 1 ; d e fi n e d , , 1 62 ; , 461 I II
I , 61 5 . i nc re a s e d fu n c t i o n , , 228 - 3 3 , 2 3 4 I
G i z z a rd , de v e l op m e n t of bi rd s , II , 5 ; fu n c t i o na l i n t e r d e p e nd e n c e , , I
3 20 . 2 35 —9 , 24 0; n u t r i t i o n a nd e g e ta l , v
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-
G l y c og e n , in a ni m a l m e t a b ol i s m , I , g e n e s i s a n d na t u ra l s e l e c t i on, I ,
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II , 4 3 —4 , 54 3 , 5 44 ; a rc h e t y p a l hy e n d o g e nou s , II
6 0—2 , 7 8 ; e o g e n , x
p ot h e s i s , , 1 22 ; II
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c a t i o n a nd n u t r i t i o n , , 1 8 0; t h e II t i o n II , , 1 29 - 1 3 1 ; t i s s u e d i ffe re n
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2 23 ; on the s k u l l II , , 561 . a da pti ve b o ne -

s t r u c t u re s , II , 3 70
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S U BJ E CT I ND E X -
.
64 1
w i th v o l ut i on I I 401 2 ; v e g e t a l
e , ,
-
, 60; g rowt h a nd org a ni c , , 1 5 2 -
3 I
a n d a s e x u a l g e ne s i s II 4 39 4 2 , ,
-
a ni m a l , v
e g e t a l , a n d e n i ro n m e nt , v
a ni m a l a n d a s e x u a l g e ne s i s
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, , i n t e rre s t ri a l , I , 4 98 , 557 ; e ff e c t
ua l g e ne s i s , II , 44 8 5 1 ; -
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a nd s e x ua l
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n u t r i t i on a nd g ene si s , r é b ra t e e x p e nd i t u r e
g enes i s, , a nd II
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e v ol u t i on a nd , 4 68 9 , 4 74 ; i n s e c t g e ne s i s ,
-
, 4 76 ; II
II , 501 —5 ; c o m m e n c e m e nt of g en s e a so n a l v
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, a nd II 4 84 5 ; i n g e r m i n a t i o n ,
,
-
, 61 5 II .
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,
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-
, 3 01 - 4 I
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5 ; , 6 12
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I , 5 1 2, 5 3 7 . c i o l og i c a l p a r a l l e l , I , 3 6 6- 8 ; n a t u
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-
p l y i ng a g e n c i e s , , 75 I . 557 ; e t h no l o g y a nd na t u r a l s e lec
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, 1 0 —1 2 2 3 2 4
, , ; I m ate p ro c e s s i n c om p re h e ns i b l e , , I
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r e s p i ra
87
642 S U BJ E CT—IND E X .
t ory II 3 1 1 ; os s e ous d i f
s y stem , ,
t r i b u t i on of p l a nt s a nd a ni m a l s ,
fe e n t i t i on
r II 35 1 ; m u c u l a
a , , s r II , 29 7 ; b e a n v a s cu l a r s y st em ,
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’
6 77 .
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4 09 ; fe r t yp es , I ,
t e rs , , I
6 9 1 ; m e a ni ng o f fe r t i l i z a t i l i ty , , I
5 9 8 ; w e i g h t o f b ra i n , , I
t i o n, ,
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, h e re d i t a r y
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. . I
t i on , I
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“
d i s c o n t i n uou s
”
d e e l op v
84 90; o f d e e l o p m e n t ,
-
v I
1 70, 1 78 ; , m e nt, I , 1 64 ; cl a s s i fi c a t ion of de
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a nd i ns t a b i l i t y o f h om o g e n eo u s , I , 38 3 ; o n
“
r i
P e s s t e t T y p es ,
n
”
I ,
509 —1 1 , 5 1 7 , 549 , 55 7 : s e g r e g a t i on 4 08 —9 ; a n c e s t ra l e q u i ne ty p es , to
a c c om p a ny i ng , I , 5 1 4—6, 5 1 7 —8 , s e g m e nt a t i on of a rt i c u l a t es , I ,
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1 1 3 ; a ga m i c m u l t i p l i ca
,
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33 6 ; a ni m a l n u t r i t i o n, 1 , 1 ; II
, 4 76 ; ce l l d o c t r i
n e, , 21 ;
-
II v er
28 9 —01 , 2 9 5 - 7 ; n a t u ra l s e l e c t i on, , I t e b r a t e e m b r y o, , 1 1 9 , 1 2 0; II m ol
2 9 5 —8 ; h e r e d i t y , , 3 01 I . l us ca n s y m m e t ry , II
2 02 ; t eg u
,
H i nd u s : food , , 68 ; d w a r f fa m i l y , I m e nt a r y o rg a ns , II , 3 1 4 , 3 1 5 ; ve r
I
, 31 0 . t e b ra t e s e n s o ry II
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’
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v a ri a t i on I ,3 3 0, 33 4 , 342 ; e
, v ol a H y b ri d s , s e p a ra t i o n of a n c e s t ra l
t i o n, I , 5 09 —1 1 , 5 1 7 , 54 9 , 55 7 ; m or t r a i t s i n, II , 6 1 6 —7 .
p h o l og i c a l d e e l op m v e nt , II , 7 —9 , H y d ro -
c a r b on s : p ro p e r t i e s , I , 6 —9
2 3 4 ; d i re c t i o n of veg eta l g ro w t h , t h e t e rm c a r b o - h y d ra t e s (q . I ,
II 1 8 1 ; ra d i a l s y m m e t ry ,
, , 1 90; II 10 .
p h y s i ol o g i c a l d i ffe r e n t i a t i on, , II H y d roc h l or i c a cid, i n g a s t ri c j ui ce ,
38 4, 3 9 2 . I , 69 .
H om og e ne s i s ( s ee G a m og e n e s i s ) . H y d rog e n : p hy sica l ch em i ca l a nd
H om o l og y , s i m u l a ti on o f, by a na l p rop e r t i e s , , 3 5 ; c om p ou nd s , , I -
I
og y , II , 14 . 6 , 8 , 9 , 1 0—1 2 ; 1 2—1 3 .
H oo k er , Sir J
I i x ; E u op e n . D .
, ,
r a H y d r ozoa ( s e e Oazl enter a ta ) .
p la nt s i n N e w Z e a l nd I 4 7 7 ; a , , H ym en op ter a ( s ee ns e c t s ) I .
p la nt d i t ri b u t i o n I 4 7 9 ; a d a p t a
s , , H y p e r t rop h y ( s ee D i s e a s e) .
t io n o f p l a nt t va i d m d i I s o r e e a, , H y p os p a d i a s , t e l e g on i c t r a ns m i s
4 8 4 ; p l a n t g r o w t h , 1 1 , 5 6 ; B a l a no s i o n, I , 64 6 .
p h on e a n d R a mes i a c eaz, , 274 ; II H y p os t a s i s of a r e l a t i o n, exe m pli

s t ru c t u ra l c om p le x i ty , II , 29 5 , fi ed in e x p l a na t i o n s of fe r t i l i za
29 7 ; re l a t i ve a nt i q u i ty a nd d is t i o n, II , 61 3 .
S UBJ E C T—IND E X .
6 43
I D E AS ( s ee P s y c h o l og y ) . h
p y s i o l o g i ca l , i n p l a nt s , , 29 2 —5
, II
I nd i vi d u a l i ty : t h e b o t a ni c a l , I , 2 44 29 5 —8 , 39 0; o f o rg a n i c w o r l d , , II
6 ; th e z oo l og i c a l , , 2
I
4 6 —7 ; t h e t e r 3 9 6 4 08 ; g e ne s i s ,
-
, 4 2 4
, II
ti l ize d g e rm p rod u c t , , 2 4 8 —9 ; I I n t e l l i g e nc e, a i t a l a t t ri b u t e , v I ,
de fi n i t i o n o f l i fe , I , 250 . 5 79 .
I nd i v i d u a t i on : a nd g e ne s i s , , 5 83 I I nt e r no d e s : va ri ed de v e l op m e nt II , ,
4; II , 4 28 —3 0, 49 9 ; t ot a l c o s t , II , 4 5 ; n u t r i t i on a n d l e n g t h , , 1 7 8 —9 II .
4 35 —7
; g e ne s i s a nd e v o l u t i on , II ,
I nt e s t i n e ( s ee A l i m e n t a r y C a n a l ) .
501 —5
, 5 29 , 5 3 0 . Int ra s e l e c t i on,
-
R ou xs’
t h e ory o f, I ,
I nfu s or i a : fu n c t i on a l s p e c i a l i z a t i on, 5 6 2 , 6 7 6—8 .
I , 3 9 1 ; p ri m a ry a g g re g a t e , II , 87 ; I r i s h , n u t r i t i on a nd g e ne s i s , , 51 0 II .
a s y m m e t ry , II , 1 8 7, 1 88 ; d i fife r I ro n : c o l l o i d a l fo r m o f p e r o x i d e , , I
e n t i a t i o n, II , 29 9 , 38 5 ; g e n e s i s , II , 1 7 , 2 0; m ol e c u l a r r e a r r a ng e m e nt ,
44 2 , 4 4 6 , 4 5 2 . I 3 3 7 , 7 04 ;
, ve g et a l a b s o rp t i o n , II ,
I nj u ri e s , rep a i r of a ni m a l , I , 21 9 , 5 73 .
22 2 —4 , 3 1 6 , II , 1 02 , 6 1 1 . I ro n i nd u s t r y , i n t e r d e p e nd e n c e of
In s a ni t y , i nh e ri t e d , I , 31 4 . s oc i a l f u n c t i on , I , 23 7 —4 1 .
Ins e ct s : t e m p e r a t u re , I , 4 7, 1 74 I s ol a t i on , a nd s p e c i es d i ffe r e n t i a
p h os p h o r e s c e n c e , I , 49 ; s e l f- m o t i o n, I , 56 8 —9 .
bi l ity , I 1 7 5 ; p a r t h e no g e n e s i s ,
, , I I s om e r i s m : o f o r g a n i c c o ns t i t u e n t s ,
2 74 5 , -
2 7 7 , 2 9 4 , 5 9 2 , 64 0; g r o w t h I 4 , , 25 ; 9, t ri a nd l
p y a t om
o -
ic
a nd r e p rod u c t i on, I , 29 2 ; sp e ci es c om 1 1 , 1 3 , 25 ; m u s c u l a r
p ound s , I ,
d i s t r i b u t i on d e t e r m i ne d by p res a c t i o n, , I
5 9 ; o r g a n i c e o l u t i o n, , v I
e nc e o f, I , 3 96 —7 ; ey es of c a v e in-
7 00, 7 03 ; d i ffe r e n t i a t i o n o f n e r e v
h a b i t i ng , I , 309 , 6 1 2 — 3 , 6 1 4 , 6 4 7 - 9 , t i s s ue , ,
5
II
3 6 —6 0, 3 6 1 ; o f m u s c u
69 3 ; p e r s i s t e n t t y p e s , I , 4 08 ; re l a r t i s s ue, II , 3 6 1 —4 .
t ro g ra d e d e e l o p m e nt , v I , 458 ;
s e g m e nt a t i on , I , 4 68 9; -
II , 114; J A C K S ON J , . H u g h l i ng s , on i nh e ri t
a borte d I
o rg a n s , , 4 74 ; E a st In a nc e of ne r o us v p e c u l i a ri t i e s , I ,
dia n d is t i b t i on I 4 7 8 ; fl o
r u , ,
ra l fe r 3 1 3 , 69 4 .
ti l i z a tio n I 5 25 ; II 1 68 —9
, , , , 406 7 , J a u n d i c e ( s ee D i s e a s e ) .
6 08 ; a p p l i a n c e s fo r c l ea ni ng an J a w s of u nc i v i l i e d a nd
, z ci vi l i zed ,
t e n n ae , , 65 1 ; I eyes, I , 65 8 , II , I , 5 41 2 , 612 , 693 .
3 1 8 ; i n t e g ra t i o n a n d h om o l o g y , II , J o h n s on , G . L i nd s e y , on i nh e r i t e d
1 1 1 —3 , 1 2 1 ; b i l a t e r a l s y m m e t r y , II , m y op i a , I , 694 .
1 98 ; se x ua l s e l e c t i o n, II , 269 ; J on e s , T . Ry m e r, on fi s s i on I , , 58 5 ,
ey es, II
3 1 8 ; e n i ron m ent ,
, ,
v II 5 90 .
4 3 3 ; c os t o f g e n e s i s , , 4 3 6 , 43 7 ; II Jul i n , C .
, on c a s t ra t i on p a ra s i
v
d e e l o p m e nt a n d g e n e s i s , ,
461 II t a i re i n C ru s t a c e a ns , II , 4 9 3 —6 .
n u t r i t i on a nd g e ne s i s , II , 4 76 , J u ng cr m a nn i a ce w: m o r p h o l og y , II ,
33 4 ; -
re l a t i o ns of high a nd l ow
Ins ec ts, S oc i a l , ori g i n of ca s t e typ es, 35 , 55 ; II , c ont i nu ou s a nd
g ra d a t i ons i n, I , 6 5 4—6 5 , 6 70, 6 7 4 , d i s c on t i n u o u s de v e l o p m e nt , II ,
6 75 , 6 7 8 8 4 , 68 6 - 8 -
. 52 —
5 , 92 : tubul a r s t ru ct u re , II , 58 ,
In sta bi l ity of t h e h o m o g e n eo u s ( s ee 6 2 ; p r o l i fe r o u s g r o wt h , II , 67, 9 1 ;
H om og e ne o u s ) . c ol ou r , , II
7 5 , 26 5 : s y m m e t ry , II ,
In s ti nc t : o rg a ni c e v ol u t i o n a nd co 1 4 0; fe r t i l i t y a nd g row t h , II 44 1
, .
o rd i na t i on o f, in m a s o n- w a s p , I , J us s i eu, A . d e, p l a nt cl a s s i fi c a t i on ,
5 74 ; a v
i t a l a t t ri bu t e , , 5 7 8 ; l os s I 1 , 3 78 .
o f s e l f fe e d i n g , i n A m a z on a nt s ,
-
,
I
66 0 1 , 6 6 3 —4
-
. K AR Y O K I N E S IS, I , 257, 25 9 , 2 6 3 —5 .
I nt g a t i n
e r o : in ch e m i cal e v ol u t i on , Ke n A r e r, .
, o n c a u l i ne buds, I , 3 58
I 23 ; m o
, r p h o l og i c a l c om p o s i t i o n, p l a n t -
c l a s s i fi c a t i o n i n N a tu r a l H i s
II 4 6 ; a
,
— r t h ro p o d , II , 1 1 1 4, 1 21 -
t o ry of P l a n ts , I , 3 78 —9 .
64 4 S U BJ E CT- IND E X .
Ki dd , B en j h is a c c e p t a nc e of 25 6 —9 , 2 70, 38 6 ; d i s t ri b u t i o n of
W eism a nn i s m , I 6 90 , . s tom a t a , II
2 60—1 ; w a x d ep os i t
,
K i t to , D r .
, his vis a l m em u o ry a nd o n, II , 2 60, 2 6 1 ; l i g h t a n d c ol o u r ,
d e a fn e s s , I , 23 0 . II 2 6 1 —2 ; s u p e r c i a l d i if e r e nt i a
, fi
Kl e b s , on H y d r od i c ty on, I , 2 8 8 ; Va u t i o n, , 2 6 3 — II
5 , 2 7 0, 3 8 7 ; a b o rt i e v
c h cr i a ,
f
II , 84 . i n p a r a s i t i c p l a nt s , II , 2 74 ; s ub
K l ei n , E .
,
m u l t i p l i c a t i on of Bac m e rg e d , l n a ua t i c q p l a nt s , ,
II
ter i a , II , 44 3 . 2 7 4 5 ; i nn e r t i s s u e
-
d i iTe re n t i a t i o n,
K o rs c h e l t , E .
, a n nu l os e s e g m e nt a II ,
2 7 8 , 38 8 ; a s cu l a r t i s s u e d i f v
t i o n, II 1 03 601 , ,
-
3 , 605 ; A r em
‘
col a fe re n t i a t i on, , 28 6 , 2 8 8 , 38 8 ; d y e II
lar vae II 1 09 , , . a b s o rp t i on a nd c i rc u l a t i o n, , II
5 7 0—4 , 5 77 ; va s cu l a r sy stem ,
II ,
L AB OU R , p h y s i o l og i c a l d i i s i onv o f, 5 88 9 2 , 5 96 ;
-
a r r a ng e m e nt , II , 6 08
I , 2 04 , 2 07 , 5 9 1 , II , 3 73 ; i t s m e a n 11 .
i ng a nd W ei s m a nn
'
s fa l l a c i o u s l n c i d op ter a ( s ee I ns e c t s ) .
t e rp r e t a t i o n , I , 6 3 4—5 .
c i d os i ren os s i fi c a t i n II 2 1 8 ; o , ,
L a c a z e - B u t h i e rs , o n o r i g i n o f a nnu r e s p i r a t i o n, II 3 3 8 ; s k e l e t on II
, , ,
l os e t y p e , II , 110 . 5 53 , 555, 560 .
L a m a rc k: z oo l og i c a l cl a s s i fi c a t i on , c i d os teu s a r m ou r, I , 5 26 ; air
I , 3 82 : op i n i o n s of E . D a rw i n b l a d d e r, II , 3 34 .
a nd , I , 49 1 , 4 9 3 —7 ; ne o- D a r w i ni s t s L e ro y - B ea u l i e u , P i e rr e , on A u s t ra
a nd , I , 63 0 1 -
. l i a n m i ne rs
'
u sa g es, I , 3 64 .
L a m i na r i a cea p s e u d o-fo l i a r a nd L es s oni a : H oo ker on g ro w t h , II , 56 ;

a xi a l de v e l o p m e nt , , 3 0; t i s s u e , II b r a nc h s y m m e try , II , 1 46 .
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t i p , , 607 I .
p l i c a t i o n, II , 27 ; H oo e r k on
L an kes t er , Si r E . B ay, a b s e nc e of g ro w t h , II
5 6 ; t u b u l a r s t r u c t u re ,
,
n uc l e u s 1 83 ; di
i n A r che r i na , I , II, 5 7 ; i n t e g r a t i o n, , 2 9 3 ; d ua l II
v e r s i t y o f P r otoz oa , i h ; z o o l og i c a l . na t u r e , , II
3 99 ; re p ro d u c t i o n, , II
c l a s s i c a t i o n, fi
, 38 7 ; b l i nd n e s s o f I 450 .
ca ve -
a ni m a l s , I , 64 7 —8 , 6 4 9 . L i eb i g , B a ro n , ni t rog e no u s food
L a ug h , d e fi n i t i o n o f l i fe a n d I , , 1 12 . s t u ffs , I , 47 8 -
.
L a u re l , l e a v e s o f II 1 49 2 4 9 , , , . L i fe c o -o r d i n a t i o n of a c t i o ns , I ,
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v e l op m e nt a nd a g g re g a t i o n , , II i ng , , I
7 8 , 1 7 8 ; It l c l i e ra u d , , 79 ; I ’
3 7 —4 2 , 7 6 ; s t e m l i e s t a l s , k k II I
'
, 41 -
D e B l a i nv i l l e , 79 , 9 3 ; L e w e s , r,
,
h o m o l og i e s , ,
42 , 7 5 —7 , 8 3 ; nu II 80; d e fi ni t i o n y i e l d e d b y c on t ra s t
t r i t i on a nd c om p ou nd , , 42 ; II i ng k e k i nd s I 8 1 —8
m os t u nl i , ,
fo l i a r a n d a i a l d e e l o p m e n t , , x v II c h a ng e s s h o w i n g I 9 1 ; v i ta l a c , ,
— — II t i o n s a n d e nv i ro nm e n t 1 92 3
"
46 50, 5 4 1 7 ; a d na t e , , 58 ; , ,
-
p ro l i fe rou s g ro w t h , II , 6 7 , 9 1 ; nu re s u l t i ng a d d i t i on to c o n c e p t i on ,
t r i t i on a n d d e e l op m v e nt , , 76 8 II I , 93, 3 26 ; C o rnt e
’
s fi
d e n i t i on , I ,
s y m m e t ry , a nd of b ra n c h e s , II , 93 ; c o r re s p o n d e n c e of e x te rna l
1 4 8 50, 1 5 1 ;-
s i ze a nd d i s t r i b u t i on a nd i nt e r n a l re l a t i o ns , I , 936 ,
of l ea ets , fl II
1 5 2—5 ; t ra n s i t i on
, 1 00; II
5 2 3 ; c on t i nu o u s a d u s t
,
j
from c om p ou n d t o s i m p l e , , 1 55 II m e nt of s u c h re l a t i o ns , , 9 9 ; c om I
8 ; u n s y m m e t r i c a l fo r m , 1 58 II , p l e t e ne s s p ro p o r t i on a t e t o c o rre
9 ; na t u ra l s e l e c t i o n a n d d i s t r i b u s p on d e n c e , , 1 01 —4 , 1 09 , I 3 49 ;
II
t i on , 1 7 9 ; m orp h o l og i c a l s u m
, l e ng t h a nd c om p l e i ty, x I , 1 03
m a y II 2 3 4 —5 ;
r , , t i s s u e d i ffe re n t i a c om ple xity of v
e n i ro n m e nt a nd
t i on II 2 4 7 ; d i, , s t r i b u t i on , II , 2 49 d e g re e o f, I , 1 04—6 ; d e n i t i o ns fi of
o ut e r t i s s ue s of s te m a nd , II , e v ol u t i on a nd , I , 1 07—1 0; d e fi c i e n
S U BJ E CT—I ND E X .
645
c i e s of fo r m ul a , I , 1 1 2 —3 ; a ct i vit y L o g w oo d , v eg et a l s t a i n i ng , II , 56 9
t h e e s s e n t i a l e l e m e nt , , 1 1 3 ; hy I 7 4 , 5 7 7—8 1 , 58 4 .
p o th e s i s of i nd e p e n d e nt i ta l v L o ng e vi ty , a nd c om ple xi ty of l i fe ,
p ri nci p l e e x a m i ne d I , , 1 1 4 —
7 ; dif I , 1 02 —3 .
fi cul t i e s p h y s i co of ~
ch em i ca l th e L ub boc k , Sir J . : on g r o w th a n d
o ry , , I
1 1 7 - 2 0; u l t i m a t e i nco m p r e g ene s i s in i ns e ct s a nd c ru s t a
h e n s i b i l i ty , , 12 0- 3 , 3 7 3 : a l i d ityI v c e a ns , I , 29 2 ; a qua t i c fi l e s I , , 4 00 .
of c o nc l u s i o ns re a c h e d , I , 1 23 ; i s L u ng s ( s ee R e s p i ra t o r y S y s t e m ) .
o r g a n i z a t i on p r od uc e d by ? I , 1 97 ; L y m p h a ti c s y s t e m : a m a ab o i d c el l s,
p re ce d e s o r g a n i z a t i o n, I ,
2 1 0; I
, 1 8 7 ; s t ru c t u ra l t ra i t s , I , 1 92 ,
fi
d e ni t i o n s o f i nd i i d u a l i t y v a nd , I , 1 93 .
2 5 0; f e ct
ef of i nc i d e n t for c e s o n,
I , 3 4 8 —9 , 3 5 5 : l e ng t h i n i n d i v i d ua l s MA CB RI D E , E . W .
, I , vi , II , vi ;
a nd sp ecies, 1 , 42 2 ; e q u i l i b ra t i on z oo l o g i ca
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3 0—6 , II
4 33 ; ni t r og e no u s p l a n t s ,
,
i nc om p re h e n s i b i l i t y , , 1 21 I .
I , 40; o rg a ni c p h o s p h o re s c e n c e , , I M a i l l e t , B d e , m od i fi a b i l i ty
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m i ca l v a r i a t i on o f, a nd org a ni c g r ow t h , I ,1 4 1 ; e p e nd i t u r e o f x
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e nc e s , II , 1 3 0, 1 3 1 , 1 47 , 1 4 9 , 1 5 8 e nt s , I
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, , 1 74, I
in fl u e nc e on fl o w e rs , II , 1 6 7 —8 , 1 7 7 ; s e l f- m ob i l i t y , , 1 7 5 ; fu n c I
608 —1 1 ; veg et a l t i s s u e -d i ffere n t i a t i o na l a n d s t r u c t u r a l d i ff e r e nt i a
t i o n, II , 25 3 5 , 2 5 8 , 2 5 9 ; a c ti on on
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t i o n, I
2 01 ; h e a r t - fu n c t i o n ,
, I , 206
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m e nt o f s e ns o r y o rg a n s , II , 3 20 . m e nt , I , 3 27 ; cl a s s i fi ca t i on I , , 3 92 ;
L i l i a ceaa, fi o ra l s y m m e t ry , II , 1 70 . cer v i c a l v e rt e b r ae , I , 394 , II , 5 64
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L i nd l ey,
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J .
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d e e l o p m e nt , ,
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; s i m ul a te d I
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,
7 0; v i t a l i t y o f e x c i s e d I 1 1 1 ; , , de t i on a nd i na c t i e p a r t s , , 5 34 v I
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Li ver fl uk e -
( s e e D i s tom a ) . o r g a ns , I
5 63 ; l o c a t i on of t e s t e s
,
L i v e w o rt s
r ( s e e H ep a ti ca ) . a n d c u r re n t t h e o r i e s , , 5 73 ; fe r I
L i za rd, re g e n e ra t i o n of l os t t a l l , I , t i lity a nd v e l o p m e nt I 5 8 3 II
de , , , ,
3 60 . 4 6 5 ; fe r t i l i t y a n d n e v o u s d e v e l op r
L oc o m o t i o n ( s ee M ot i on) . m ent , I
5 9 8 —9 ; , l o co m ot i on a nd
L og ic, r e a s o n i ng a nd d e ni t i on fi of e l on g a t e d fo r m , 1 5 ; s y m m et ry , II ,
l i fe , I , 8 1 —6 . t e g u m e n t a r y s t r u c t u re , , II
6 46 S U BJ E CT I ND E X -
.
314; c i r c u l a t i o n, II , 34 0; v a s c ul a r ~
6 2 1 ; s e p a ra t i on o f a n c e s t r a l c on
i ty a n d o v a - m a t u r a t i o n, II , 3 42 —3 ; s t i t ut i o n s i n p l a n t h y b ri d s , II ,
a c ti vi ty a nd m u s cu l a r c ol o u r, II , 61 6 ; s i ng l e a n d d ou b l e s t oc ks ,
II ,
3 6 5 9 ; fu n c t i on a l i n t e g ra t i o n ,
-
, II 62 2 .
3 7 5 ; o u t e r t i s s u e d i ff e r e n t i a t i o n , M at ter l nc o m p r e h e n s i b i l i t y
, of ln
II 3 8 7 ; g row t h a n d g e ne s i s ,
, , II t e ra c t i o n s , , 12 1 —2 I .
4 56, 4 5 9 ; c o m p a r a t i e fe r t i l i t y , v M e c h a ni c s : t r a nv e r s e s t ra i n s , II ,
II , 4 6 5 , 4 70; h e a t e p e n d i t u r e x a nd 2 09 —1 2 ; g e n e s i s o f e r t e b r a t e v a xi s,
g e ne s i s
, II
4 6 7 —9 ; a c t i i t y , v a nd II , 2 12 —6 , 2 1 6 —8 , 22 4 , 2 2 5 —7 ; os s e
fe rt i l i t y , II 4 72 ; , n u t ri t i on a nd ou s II 34 5 5 1 ; d i s
d i ffe r e n t i a t i o n , ,
-
g e ne s i s , , II 4 79 —8 0 . i n t e g a t e d m ot i on II 3 7 5 ; a n a l
r , ,
M a n : e ffe c t o f c l i m ate on v i g ou r I , , og y f om l o c o m ot i v e
r II 5 1 7 9 ; , ,
—
3 0; fle s h a nd g ra i n e a t e rs c o m f t u ur e h u m a n e v o l t i on II 5 24 ; u , ,
p a re d , , I 68 ; l o ng e i t y a n d l i fe , v I , st i n a n d v e g e t a l s t u c t u e II
ra r r , ,
1 03 ; c o m p l e x en v i ro nm e nt I , , 1 05 5 7 4 —88 , 5 9 2 —6 .
em b r y og e n y of a rm , I1 69 ; fe r
, M ca naw : c o n t ra c t i l e fu nc t i o n s , I ,
t i l i ty a nd c on d i t i o n s a ffe c t i n g i t , 58 ; , II
3 7 4 ; i nd i i d u a l i t y , , 248 v I
I , 3 00, 5 7 0, 5 83 , , II 506 —2 148 4,
; h e t e r og e ne s i s , , 2 7 3 ; fe r t i l i t y , ,
I I
i nh e r i t a n c e of f u n c t i on a l l y p r o 5 82 ; s t r ob i l i z a t i o n , I , 592 ; sy m
d uce d c h a ng e s , I , 3 1 0 3 5 4 1 , 605 -
, , m e t ry , II , 1 88 —9 1 .
6 08 , 612 , 65 2 , 67 3 , 68 9 , 69 3 —4 ; M e h n e rt , E .
, on fe e t of p e nt a d a c
h e re d i t y a nd s ex , I 3 1 5 —6 ; f nc
, u ty l e v e r t e b ra t e s I , , 4 61 .
t i on of b i l i r ub i n, I 3 3 0; co i n
, us M e ns e l
'
s s a l t, t e m p e ra t u r e a nd iso
m a r ri a g e s , I , 34 6 , II 6 1 5 ; p ri m i
, m e ri s m , I , 77 .
ti ve no t i o n s , I , 4 1 7 9 ; i nu t i l i t y o f
-
M e ta b ol i s m a nt i t h e s i s b e tw e e n
A p p end i x verm i fo r m i s , , 474 ; d i I p l a n t s a nd a n i m a l s , , 6 2 —3 ; c v o I
m i nu t i o n o f jaw , I , 54 1 , 6 1 2 , 6 9 3 l u t i o n h y p o t h e s i s a n d p r i m o rd i a l ,
-
c o - o r d i n a t i on of a ct i o ns g re a t e s t I , 6 3 —4 ; i n p l a n t s , , 04 —7
; a ni I
i n, , I
5 7 9 ; f u nd a m e n t a l t ra i t s o f m als, I , 6 7 —7 7 ; n e r vo m u s c ul a r
-
ac
s e x, , 5 9 4 —7 I
; ob e s i t y , , 5 94 ; s u b I t i vi ti e s, 1 , 71 - 7 ; s u m m a ry , I , 77 ;
s t a nc e a nd w e i g h t o f b ra i n , , I c el l p ro c e s s e s , I , 2 61 .
596, 5 99 ; d i s t r i b u t i o n o f t a c t u a l M et a l s : re m a r ka bl e i n t e ra c t i o n s of
p e r c e p t i e n e s s , , v
60 2 8 , 6 6 5 —6
, I -
s om e , I1 21 ; m e l t i n g o f a l l o y s ,
, I ,
6 72 —3 , 6 9 2 ; t e l e g o n y , , 6 2 5 , 6 4 4— 5; I 3 39 ; a t om i c t e -a r r a ng e m e nt , I ,
d e g r a d a t i on o f l i t t l e t oe , I , 652, 352 .
6 7 3 ; t r a n s m i tt e d o s t e o l og i c a l p e M e ta m e r i s m ( s ee S eg m e n t a t i o n ) .
c u l i a r i ti e s of P un abi s, , 68 9 ; j I M e ta z oa : c e l l u l a r s t r u c t u r e I , , 1 8 4,
t ra i t s of t w i n -b e a r i ng w o m e n, II ,
1 94 , II , 21 ; s ub ord i n a t i o n o f un i t s ,
457 ; c om v
p a ra t i e m a m m a l i a n fe r I ,1 8 5 - 7 ; g e n e r a l c h a ra c t e r s o f t i s
t i l i ty , II , 4 65 ; f u t u r e e o l u t i o n , v s ue s ,, I
1 88 —9 ; p r o t o p l a s m i c c o n
II , 5 2 2- 3 7
(8 . 66 a l so L a ng u a g e t i n u i t y , 1 , 1 90—2 , 1 9 4 , 62 8 ; g e n e s i s
a nd S o c i o l o g y ) . o f fOO d c a i t y a n d v
i s u a l o rg a n , v
M a n a t e e na i l l e s s p a d d l e s
, , I , 473 . I , 1 95 ; W ei sm a nn s
’
d l iI e re n t i a t i o n
M a nx c a t s I 3 03 , , . t h e o ry , I , 63 7 —4 3 .
M a r c h a n ti a c ea : s y m m e t r y ’
, II , 1 40 M e t e o ro l o g y : no n - v i ta l c h a ng e s
ou t e r t i s s ue d i ffe r e n t i a t i on, II , sh ow n i n, I , 82, 84 ; c ry s t a l l i za

t i on of s t or m g l a ss, I , 96 ; spe
M a rm ot, h y b e r na t i o n a nd w a s te , I , cia l c re a t i o n, I , 41 9 ; rh y t h m i n,
2 1 4—5 . a nd o rg a n i c c h a ng e , I , 4 9 9 - 5 01 ,
M a r r i a g e ( a ce M u l t i p l i c a t i o n) . 557 ; v
a r i a t i o ns due to g e o l og i c
M a rs h 0 C o n t e l e g o n y I 6 4 4
, . .
, , , . c h a ng e , 5 03 I , .
M a s t e r s M T o n fo l i a r h o m o l o g y
, . .
, , M i c r os tom i d a , s e g m e nt a l r e p r od u c
II , 4 6 —7 ; s e l e c t i o n o f i n c o nS p i c u t i o n, II , 1 02 .
o us v
a r i a t i o n s i n p l a nt s , , 2 98 , II M i g ra t i on of a ni m a l s p eci es, I ,
S U B E C T—IND E X J .
647
39 6 —401 , 4 1 1 ; s o l a r i n u e nc e s , , fl I 2, 22 5 ; r u d i m e nt a ry o r g a ns , I ,
5 00; p a r t p l a y e d b y , i n o r g a ni c 4 7 2 —5 , 5 5 6 ; s t r u c t u r a l a nd fu n c
e v o l u t i on I , , 5 68 ; ca us e s o f, II , t i o na l c o - op e r a t i o n , II
3 , 23 9 ; i n
,
5 33 —4 . t e g r a t i o n, II , 4- 6 , 1 8 1 — 9 6 ; c h a ng e
Mi l k , h ea t a n d s u p p l y o f, II , 4 68 . of s h a p e , II , 6 ; fo r m u l a o f e v ol a
M i l ne -
E d w a rd s , H .
, p h y s i o l og i c a l t i o n, II , a s i n t e r p re t e d by
v
d i i s i on of l a b o u r,
”
I, 2 04 ; W e i s p h y l og e ny , II , 1 0—6 ; e v o l u t i on a nd
m a nn s
’
e r ro ne o u s a p p l i ca t i on of cel l d o c t ri ne , II ,
1 7 —2 1 .
i t, I , 63 4 ; o n oc u l a r s t r u c t u r e , II , M o rp h o l og y , A ni m a l : e v o l u t i on a nd
318 . s e g m e n t a t i o n of A r ti c ul a ta , I ,
468
M i n d ( s ee P s y c h o l og y ) . 9; v erteb ra l col um n de v e l op m e nt ,
M i t os i s ( s ee K a r y o k i ne s i s ) . I , 4 70; si m ul a t ed h om o l og i e s , II ,
M o b i l i t y m o l a r a nd m o l e c ul a r
, , I , 1 4—5 ; p r i m a r y a g g r e g a t e s , , 85 II
14; en v i ron m e nt a nd s el f m ob i l 8 , 12 3 —4 ; s e c o n d a ry , , 88 9 1 , 1 24 II -
ity, I , 1 77 . t e r t i a ry ,
, 9 1 —3 II
; i nt e g r a t i o n a n d
M oh l , on p h aenog a m i c g ro w t h , II , i nd e p e nd e nc e o f i n d i i d u a l i t y , , v II
82 . 9 3 —9 , 12 4 ; a n n u l os e s e g m e n t a t i o n,
Mol e ,
m od i fi c a t i ons due t o h abi ts, II , 9 8 —1 01 , 1 06 —1 0, 1 2 5 —7 , 6 02 7 ; -
II , 391 .
p r og r e s s i e a n nu l o s e v i nt e g r a t i o n ,
M ol e c u l e s : m e c h a ni c a l l y c o n s i d e r e d , II , 1 00—5 , 1 1 1 —5 , 12 1 , 12 4 , 22 3 ; un
I, I 3 3 7 4 0; n v
1 4 ; s t a b i l i ty , ,
-
er e i nt e g ra t e d m ol l u s ca n fo r m , II ,
’
II 35 5 —6 1 3 7 9 8 2
d i tfe r e n t i a t i o n , , ,
— . 1 1 5 —7 ; e rt e b r a t ev s e g m e n t a t i on
M ol l c x i a l d v l p m nt I 1 6 5
us a : a e e o e , , a nd i n t e g ra t i o n , , 1 1 7 —2 3
, 1 2 4—7 , II
g e ne i I 27 1 II 444 ; h
s s, , , ,
er 2 2 3 —4 602, 6 06 —7 ; m ot i o n
, a nd
m a p h od i s m I 34 1 ; c l a i fi ca t o y
r , , ss r s y m m e t ry , 1 8 3 —5 ; sy m m eII ,
t its
ra I 3 9 2 ; d i t i b t i on i n
, , s r u t ry o f p ri m a r y a n d s e c o n d a r y a g
tim e I 4 05 408 4 1 0 44 6 —7 :
, , , , , g r e g a t e s , , 1 8 6 , 1 8 7 —9 1 II
; of
t r oc h o p h o r e a nd its re l a t i on s h i p s , c om p o u n d C ozl en tc r a ta , , 1 9 2—4 ; II

I , 447, II , 1 08 , 1 09 , 1 1 5 ; de el v s i m u l a t i on p l a nt s h a p e s , of , II
o p m e nt , I , 4 60; a m p h i b i ou s a n d 1 92 ; s y m m e t ry of P o l yz oa a n d
t e r re s t ri a l , I , 48 1 ; i nd i re c t e qui T uni ca ta , , 1 94 ; of II
P l a ty h e i
l i b ra t i o n, I , 5 34 ; s eco n d a r y a g m i n th es a n d E c h i no d e r m s , II , 1 95
g r e g a t i on , II , 1 1 5 —7 ; s y m m e t r y , 7; of A nnu l os a , II
1 9 7 —201 ; o f ,
II , 2 01 — 3 ; out e r t i s s ue , II , 3 1 0, m ol l us c s , II ,
2 01 —3
; of e rte v
3 87 ; a l im e n ta r y s y s tem , II , 32 5 ; b ra t e s , II , 2 03 —6 , 2 08 ; s i m i l a r i t y
va s cu l a r s y s te m , II , 3 4 0—1 . of a n i m a l a nd p l a nt , II , 2 06 ; c el l
M ol l us co i d a , II , 598 .
( S ee P ol y z oa s h a p es , II , 2 2 8 —3 0; e v l o u ti on a nd
a nd T u n i c a ta .
) g e n e r a l i z a t i on s s u m m a ri z e d, II ,
M o no c o t y l e d o n s : g row t h , I , 1 38 , 2 3 1 —5 .
( S ee a l s o S t r u c t u re ) .
1 3 9 , 1 4 3 ; u ni a e nt , ,
x i a l d e e l op m v I M or p h o l og y , Ve g e t a l s i m ul a ted
1 65 ; c o t y l e d o n o u s g e r m i n a t i o n a n d h om ol og i e s , II , u ni c e l l u l a r
e n d og e n o u s g r o w t h , , 5 9 —6 2 , 69 II p l a nt s , , II
2 1 ; a g g re g a t i on a n d l n
7 2 , 82 —3 , 1 8 1 —2 ; a b s e n c e o f h e l i c a l t e g ra t i on, , 2 2 —6
, 7 8 —
II
9 ; p s eu d o ?
p h y l l ot a xy i n R a vena l a , II , 1 82 ; fo l i a r d e e l op m e nt , v , 2 6 —8 II
s u r fa c e 2 5 7 ; ou t e r
c on t r a s t s , II , p s eu d
-
a xial , II 28 ,
—9
; p s eud o
l ea f t i s sue, , 2 63 ; w o od fo r m a II fo l i a r a nd a x i l IIa , , 3 0 2 ; com
t i on, , II
2 7 8 ; g ro w t h a n d g e n e s i s , p o s i t i o n o f A r c h e g on i a t e s , , 33 II
II , 45 1 . v
5 ; l e a f d e e l op m e n t a n d a g g r e g a
M o n s t r os i t i e s i n p l a nt s ,
, II , 7 8 , 54 1 , t i on , II , 3 7 —4 2 , 7 5 —8 ; fo l i a r h o m o
5 46 ; v e r t e b ra t e , , 118 II . l og i e s , II , 4 2 —6 , 7 5 8 ; fo l i a r a n d -
M or g a n , T . H .
, on re g e n e r a t i on of a xi a l dev II 4 6 —5 0 5 4 1
e l o p m e nt , , ,
P l a na r i a , II , 1 02 , 6 1 1 -
. 7 ; g ow t h r nd d e v e l o p m e nt f a o
M or p h o l o g y : p ri s e d i n , ,
fa c t s c om I A h e g n i a t s II 5 0 6 ; o f t
rc o e e , ,
—
1 2 5 —6 ; m orp h o l o g i c a l u n i t s , , 1 90 I n og a m II 5 6 —64 7 8 8 0; a x i l
s, , ,
- s
6 48 S U BJ E CT -
I ND E X .
l a ry b ud de v e l op m e nt II , , 6
5 —9 M u c ou s m e m b ra n e , d i flfe re nt i a t i on,
p h ae nog a m i c m o d e s o f g r o w t h , , II II , 32 1 —2 , 3 8 9 .
6 9 —7 2 ; h o m o l og i e s , , 7 3 —5 , 8 0—4 ; II M u l t i p l i ca t i o n : d e c l i n e of fer t i l i t y
v
d e e l op m e n t o f fo l i a r i n t o a i a l x wi th e v o l u t i on , I, 1 03 , II , 43 1 ;
o r g a ns , II
7 5 —8 ; r s u m , ,, 78 é é II v i ta l i s m I , 1 1 6 ; p h e no m e n a
, cl a ss i
8 0; c ri t i c i s m s a n d re p l i e s , , 80 II fi ed , I , 1 3 0; t h e t e r m “ g e n e s i s ,
4 ; c a n p l a n t -s h a p e s b e fo r m u I , 26 9 p r oc e s s e s c l a s s i
: fied I , , 2 70
l a t ed ? II , 1 2 8 ; g ro w t h a nd d i ffe r 6, 336 , 5 83 ; a p roc e s s of d i s i nt e
e n t i a t i o n, II
, 1 29 —3 1 ; k i nd s of g ra t i on , , I 2 76 ; r e p ro d u c t i ve tis
s y m m e t ry , II , 1 3 1 —3 ; s y m m e t ry o f s ue l n g a m og e n e s i s , I , 2 78 —84 ; n u
p ri m a ry a g g re g a t e s , II , 1 3 4- 7 ; of t r i t i on a nd g ro w t h , I , 2 8 5 —94
,
s e c o n d a ry , II , 1 3 7—4 0; t e rt i a ry , II , 2 9 5 —7 , 29 9 ; ,
n a t u ra l s e l e c t i o n, I
1 40—3 ; s y m m e t ry a nd en v i ro n i n g 29 5 —8 ; h e r m a p h ro d i s m , , 34 0—4 I
in fl u e nc es II , , 1 43 -4 ; s y m m et ry o f l n-a n d l n b r e e d i ng ,
-
, 344—7 ; p h y I ~
b r a n c h es , II , l ea f a nd s i o l og i c a l u ni t s , I , 35 0- 5 ; l a w of
b ra n c h s y m m e t ry , II , 1 4 8 —5 0; ra c e -m a i n t e n a nc e , I , 581 ; , 42 0 II
p h ae n og a m i c u n i t h om o l og y , , II 3, 4 30; et
f ec t of m e n ta l a p p l i ca
1 5 1 ; s i z e a nd d i s t ri b u t i o n o f l e a f t i o n, I , 5 9 7, II , 5 1 1 —4 , 5 1 6 —9 , 5 3 0;
l ets, II
1 5 2—5 ; t ra n s i t i o n f ro m
,
i nd i v i d u a t i on a n t a g o ni s t i c t o, I ,
c om p ound t o s i m p l e l e a e s , , v II 5 9 8 —600, II , 42 8 —3 0, 4 35 —7 , 4 99 ,
1 5 5 —8 ; u ns y m m e t r i c a l l ea f d e ve l 501 —5 ; ch ec ks p ut by c a rn i v o re s
o p m e nt , II
1 5 8 —9 ; d i ffe re n t i a t i o n
, o n, II , 4 05 ; fou r fa c t or s i n ra t e
of h om o l og o u s u n i t s , , 1 5 9 —60; II o f, II , 4 1 6 , 4 3 5 ; d e s t ru c t i e v a nd
fl ora l s y m m e t ry , II , 1 6 1 - 74 ; c e l l p re s e r va tiv e forc e s , II , 4 1 7—2 0;

d i ffe r e nt i a t i on a n d m e t a m o rp h o rh y th m of s p e c i e s ,
4 1 9 ; n u t ri II ,
s i s, , 1 7 II
5 —7 ; n u t r i t i o n a nd d i ff e r t i o n a n d d i s i n t e g r a t i o n o f, , 424 , II
e n t i a t i on , 1 7 8 ; a nd i nfl o r e sII ,
42 5 , 4 3 0; i n t e g ra t i o n a nd g e ne s i s ,
c e nc e, , II
1 7 9 ; h e l i c a l g ro w t h o f II, 42 6 —8
; i n u e nc e fl of en v i ro n
p h aen o g a m s , , 1 8 0— 1 ; s u m m a ry II m e nt , , II
43 2 —3 ; a nd v a r i a t i o ns of
o f s y m m e t ry , , 2 34 ; s t r e s s a n d II e x p e nd i t u r e II 43 3 5 ; g ro w t h , ,
-
s t r u c t u re , II ,
2 7 5 —9 , 3 8 8 . ( S ee a ls o a nd a s e x ua l g e ne s i s II 4 3 9 —4 6 ; , ,
S t r u c t u re ) . a s e x u a l a nd s e x u a l d i s t i ng u i s h e d ,
M o rt o n ,
L o rd , qua g g a -
m ar ked foa l , II 44 8 ; s e x ua l g e ne s i s a n d
,
I , 624 . g ro w t h , II ,
4 4 8 —5 8 , 4 9 5 ; a nd de
M os e r i m p re s s i o ns
, p rod u c e d by ve l o p m e nt, II , 4 6 1 4 3 ; p l a nt ex
l i g h t o n m e ta l s , , 352 I .
p e n d i t u re , II 4 6 7 ;
, a ni m a l expe n
M os s e s : v a ri e d de v e l op m e nt II , , 50 d i t u re, II 4 68 , nu t r i t i o n in
1 , 52 ; h om o l og i e s , II , 8 0, 8 1 ; l n p l a nt s , II 4 75 1 1 ;
, , 5 in a ni m a l s , II ,
d e fi ni t e n e s s , II , 29 6 ; m u l ti p l i c a 4 7 6 —8 4 , 5 1 1 ; s e a s o na l v a r i a t i o ns ,
t i on , II , 4 41 .
II , 4 8 4 - 5 : nu t r i t i o n , r és m né , II ,
M oth , c l ot h e s , fo o d of la r va , I ,
48 6 , 49 7 —9 ; n u t r i t i on a nd p a ra
77 .
s i t i e, II , 4 8 6 —90; re v e rs i o n to
M o t i on : o r g a ni c , a nd en v i ro n m e nt ,
a g a m o g e ne s i s , II , 490- 2 ; h um an
I, 7 5 —7 , 1 7 5 8 , 1 9 6 ; of a n i m a l s a nd fe r t i l i t y , II , 5 06 —1 0; D ou b l e d a y o n,
w a ste, ,
I
2 1 4 , 2 2 0; s i m u l a t i o n o f II 5 1 0—
, 2 ; c i i l ized v a nd u nc i vil
l oc om o t i ve s t r u c t u re s , II , 15 . i zed, , II
5 1 4—6 ; h u m an e vol u t ion
M ot o r o rg a n s , d i ffe re n t i a t i o n o f, I ,
a nd d e c l i ne i n,
, 5 29 —3 1
; the II
2 62 .
fu t u re of o
p p u l a t i o n, , 5 3 2 —7: II
M ou n t a i ns : cli m atic e ffe c t s , I 504 ; e q u i l i b r a t i on a nd e v o l u t i on , II ,
g ro w t h of t re e s on , II , 1 4" 5 37 .
M ou s e : fe r t i l i t y o f, II ,
42 1 , 473 Mus c l e : e l e c t ri c a l c o nt ra s t s , I , 50
t a p e w orm p a ra s i t i s m , II , 4 90 a c t i o n o f, I
5 9 ; m e t a b o l i s m , , 70,
,
I
c om p a re d w i th ra t, II , 5 03 —4 . 7 1 —4 : de fi n i t i o n o f l i fe a n d a c t i o n s
M ucor , II , 2 2, 1 2 3 . o f, 1, 1 12 3 -
: g ro w th a nd func t i o n,
S U B E CT—l ND E X J .
64 9
I , 1 51
1 55 ; d e , v e l op m e nt I 1 7 0; , , 5 5 2 ; h um a n ra c e s , I , 553 ; c u r r e nt
fi c a t i o n of t i s s u e s views
'
H ert w ig s c l a s s i 5 5 9 —60;
'
, , I, p a nm i x i a a nd
I , fu n c t i o n a l d i ffe r e n t i a t i o n, c e s s a t i on of s e l e c t i o n, , 5 6 0-3 : I
I, 2 03 —4
; w a s t e a n d re p a i r , 2 1 5 4 i n t r a —s e l e c t i o n, I , 5 62 , 6 7 6 —8
m od i fi a b i l i t y a nd a d a p t a b i l i t y , , I E im er s
’
t h e o ry o f o r t h og e n e s i s , I ,
2 2 8 —9 , 2 3 0, 2 3 2 ; c or r e l a t e d v a ri a 5 64 ; M r C u nn i n g h a m s c r i t i c i s m ,
.
’
t i o ns , , I
5 3 6 —9 , 6 1 4 —2 1 , 6 76 , 69 3 ; I , 5 65 —6 ; l oc a t i o n o f m a m m a l i a n
r e s i s t a nc e to s t r a i ns , I , 6 39 ; ac t es tes , , I
5 7 3 ; c o o rd i n a t e d i n -
t i o n o n b on e s i n P u n j a bis I , , 68 9 s t i nc t s of m a s o n- w a s p , I , 5 74 ;
d i if e re nt i a t i o n , II 3 6 1 —9 ; a c t i i t y
, v t a c tua l p e rce p t i e ne s s , , v I 6 03 —8 ,
a nd c o l o u r, II , 3 6 5 —9 : i nt e g r a t i on , 633, 64 6 , 6 6 5 , 6 7 1 , 6 7 2 , 6 9 2 ; e r ro n e
II , 3 76 , 3 82 : q u i l i b r a t i on i n a c
e ou s l y i d e nt i fied w i th a rt i fici a l se
t i o n, II , 39 3 ; a c t i v i t y a nd fe r t i l i t y l e c t i on, I , 609 , 6 9 5 ; re e rs e d s e v
II 4 70—2 ; fu t e h u m n
i n b i rd s , , ur a l e c t i on , I , 6 1 1 ; b l i n d n e s s of c a e v
e v o l t i on II 5 2 3 ; i g i n of e
u , , or v r a ni m a l s , I , 6 1 3 , 6 1 4 , 64 7 —8 , 69 3 ; c o
t b e t e t y p e II 5 9 8 —6 00
ra , , . a da p t a t ion of c o -o p e r a t i ve p a rts ,
M i c l i m i t e d a d p t a b i l i t y of v o i c e
us : a I , 6 1 4, 62 1 , 66 3 5 , -
6 70, 6 74 , 6 7 5 ,
nd
a e I 2 3 1 ; i nh e i t a nc e o f
a r, , r 68 9 , 6 9 2 ; W h e re op e ra t i 6 32 ; ve I , ,
fa c l t y I 3 1 1 2 69 4
u , ,
—
, . W e i s m a nn on c on c e i a b i l i t y of v
M t i l t i on s t h q e t i o n of t h e i r
u a , e u s p ro c e s s , I , 65 1 ; d e g e n e r a t i o n of
i nh it n e I 631
er a c , , . l i t t l e t oe , I , 65 2 —3 , 6 7 3 ; g e n e s i s of
M y cetozoa , g ro w t h a n d r e p r od u c t i o n , ca s t e g ra d a t i on s i n s o c i a l i n s e c t s ,
I , 2 98 —9 . 1 , 6 5 4 — 60, 6 6 3 , 6 7 0, 6 7 5 , 68 4 ; s e l f
M y oc o m m a ta (m y ot o m es ) , a nd ver f e e d i ng i n s t i n c t i n a n t s , , 6 6 0 —2
, I
t e b ra t e s k e l e t o n II , , 21 6 , 21 7- 8 , 6 70; ru di m e n t a r y o rg a ns , I , 6 6 7 —9 ,
2 22 .
'
6 71 , 6 9 2 ; h o r ns o f s t a g , , I 6 76 —8 ,
M y op i a i nh e ri t a n c e o f I 3 06 6 9 4
, , , , . 6 9 2 ; m u s i c a l fa c u l t y , , 6 94 ; th e I
M y r i a m d a fa s c i a ta I 3 6 1 ; II 4 4 5
’
, , , . n e o- D a r w i n i a n p o s i t i o n r e i e w e d , v
M y r i a p od a : g e m m a t i o n I 5 8 9 ; s e g , , I 6 94 —5 ;
, v
e g et a l n u t r i t i o n , II , 51
m e n t a t i on, I , 5 9 0, II , 1 1 3, 1 1 4, 2 ; up r i g h t e g e t a l g row t h , v II , 56
6 01 ; d e g e ne r a t e d eyes of ca ve l n -
7 ; e nd og e n ou s g r o w t h , II , 5 7- 8 ;
h a b i t i ng , , 64 9 ; I i nt e g ra t i o n a nd e x og e nou s II , , 64 ; N a vi c ul a sy m
h om o l og y , II , 1 1 1 —4 ; g ene s i s , II , m e try , II , 1 35 ; fol i a r, II , 1 58 ;

4 45 . fo l i a r d i s t r i b u t i on , II , 1 67, 1 79 ;
M ywoth a l l op h y ta , I , 3 78 . fl ora l fe r t i l i z a t i o n a nd s y m m e t ry ,
II , 1 68 - 70 608 —1 1 , ; h e l i c a l p h aen o
NAI L S , m a m m a l i a n, I , 4 73 .
g a m i c g ro w t h , II , 1 8 1 ; E ch i n oder
Na l s : re g e n e r a t i o n of d et a ch e d m a ta a n d b i l a t e r a l s y m m e t ry , II ,
p a rt s , I , 21 9, 361 . 1 97 ; ve rteb ra t e s t ruc t ur e , II , 21 4

N a rc i s s u s , s e p a r a t i on of a n c e s t ra l 2 0, 22 7 ; p h aenog a m i c t i s s u e d i ffe r
t ra i t s i n h y b r i d s , , 61 7 II . e nt i a t i o n , , II
2 4 8 : p h y s i o l og i c a l
N a t u ra l s e l e c t i o n : s t r u c t u r a l m od i fi d i ffe r e n t i a t i o n , 2 5 2 , 2 5 6 ; roo t II ,
c a t i o n, I
2 1 1 ; i n c e l l p r oc e s s e s , ,
, I l ets of ivy, II , 2 54 ; s tom a ta a nd
2 63 —4 ; m u l t i p l i c a t i on , , 2 9 5 —8 ; I fo l i a r s u r fa c e s , II , 2 6 1 , 2 62 ; fl ora l
a i d e d b y f u n c t i o n, , 3 08 —1 0; s p e I fe r t i l i z a t i o n , II , 2 6 8 —9 ; s e xua l se
c i a l c re a t i o n, , 42 6 7 : t h e t e r m I -
l e c t i o n, II , 269 ; v e g e ta l ti s sue d if
“
s ur v i va l of th e fi tt e s t , I , 5 3 0; f e r e nt i a t i o n , II , 2 7 9 ; w o o d fo r m a
i nd i r e c t e q u i l i b ra t i on ,
I , 5 3 0-5 , t i o n, II , 2 8 7- 8 , 2 9 0; a n i m a l t i s s u e
5 5 2 —3 , 557, 5 7 1 ; c h a ng e s u ne x d i ffe re nt i a t i o n, ~II , 3 04—8 ; e vol u
p l a i ne d b y , 1 , 5 3 5 —42 , 5 7 1 , II , 3 71 t i on of n e r v ou s sy s tem , II , 3 07 - 8
t e nd e n c y t o e c o no m y, I , 5 3 6, 5 62 r e s p i ra t o r y syst em , II , 31 1 ; der
d e c re a s e of ja w , I 54 1, , 69 3 ; g en m al c a l l os i t i e s , , II
3 1 2-4
; s e ns or y
e ra l d o ct r i n e of e v ol t i u o n, , 5 43 I o rg a n com p l e x i ti es, 32 1 ; s i n II . k
8, 557 ; u n c e a s i ng o p e r a t i o n, I , a nd m u c ou s m e m b ra ne d i ffe r e n
650 S U BJ E CT —I ND E X .
t i a t i o n, II 3 22 ; l oc a l i a t i on of e , z x fa c t o r s i n e v ol u t i on of, II , 307—8
c e t i on
r II 33 3 ; re p i t y o
, , s ra or r d i ffe r e n t i a t i on fr o m m us c l e, II ,
g n
a s o f fi s h II 3 3 5 —8 ; h t es , ,
ea r 3 63 .
( S ee a lso P s y c h o l og y .
)
a nd va l y s te m II 34 1 3 4 4 ;
s cu a r s , , , Ner v o u s ne s s , h e r e d i t a ry t r a ns m i s
o ss
e s dif ou e nt i t i o n II 3 5 5 ; er a , , s i o n, I , 3 07 .
al so m u la II 3 63 3 68 —9 ;
scu r, , ,
N e u ri n e , I , 5 94 , 5 9 7 .
f l s e j o i nt s
a II 3 7 1 ; i n c t n , , se u N e u t e r i ns e c t s ( s ee ns e c t s)
-
I .
t i t i n nd g n
r o i II 4 9 9 ; e c o no
a e es s, , Ne w Z
e a l a nd : i n a s i o n o f a l i e n v spe
m i s of e v ol t i n
c II 5 01 —5 ; a u o , , u ci e s , I , 4 77 ; k i ns h i p of pa st a nd
n n c i a t i o n of s u r v i v a l o f p r e s e n t fo r m s , , 48 9 I
’
th s
or e u .
t h e fi t t e t i n 1 8 5 2 II 5 2 8 —9 ; e v i l s
s , , N i t r og e n : p rop e r t i e s , , 3 — 5 , 2 0, 2 4 I
of i nte f nc e w i t h r i n m n II
ere , a , , c o m p o u nd s a n d t h e i r p r op e r t i e s ,
5 3 2 3 ; v eg e t l t i
— e fo m t i on a ssu r a , I , 6 , 8 , 9 , 1 2 —1 4 2 5 - 6
, , 39 , 4 1 , 4 2 —
3,
II 5 8 2 5 9 4—6 ; o i g i n o f v e t e
, , r r II , 2 5 0; o rg a n i c
i m p o r t a nc e , , I
b rat e t y p e II 5 9 9 , , . 4 2 —3 ; e v o l u t i on o f h e a t a n d o i d a x
N a t re u m e ,c m p l ex tha n s p
or o u t i o n, I , 47 ; v i o l e nt o r g a n i c e ffe c t s
p os e d I 25 2 4 5 0 of c o m p ou nd s , I 5 4 5 ; f un c t i o n i n
-
, , , .
,
N a i c l a s y m m e t y II 1 3 4—5
v u , r , . m e t a b ol i s m , I , 6 3—4 , 6 6, 6 8 76 ; -
N eb l a H y p t h s i
u r I 23 o e s, , .
p re s e n c e i n p rot op l a s m , , 66 ; a c I
N e g a t i on i n c on c e i v b i l i t y o f
, th e a , t i o n i n d i g e s t i o n, , 6 9 ; fa t a c c u I
u l tim te t t af t t h I 6 75 es o ru , , . m u l a t i o n a nd fe r t i l i t y , II , 483 .
N e g ro e s , t e l e g o n y i n U n i t e d S t a t e s , N i t ro g l y c e r i ne,
-
v i o l e nt e ffe c t s o f,
I , 64 4—5 . I , 5 5 , 1 22 .
N em er ti d w c o n t i nu i ng v i ta l i ty of N ot o c h o r d : s e g m e n t a t i on , II , 1 25 ,
p i li di um , I , 2 5 0; bi l a t e ra l sy m 2 1 8- 2 2 ; for m a t i o n, II ,
2 1 7 —8 , 6 00;
m e t ry , II , 1 95 . in P e rm ian v e r t e b ra t e s II , , 225 .
N e o D a rw i ni s t s ,
-
a nd L a m a rc k I , , N ou m e no n, l i fe no t m a n i fe s t e d a s,
6 3 0; t h ei r p os i t i o n re v i e wed I , , I , 580 .
6 9 4—5 . N u c l e i n, II , 21 .
Ner ves : e l e c t r i ca l p h e n o m e n a , I , 51 Nucl eus c e n t ra l de v e l o p m e nt , I ,
g e n e ra t i on o f n e r ve -
fo r c e , I , 5 2 —6 , 1 63 ; i n sim p l e or g a n i s m s , I , 1 83
6 0; cor p u s c u l a 7 5 ; H e rt
ta c tu s , I , p h e no m e na x
e h i bi te d by, I , 25 5
w ig
’
s cla ss i fi c a t i o n o f t i s s ue s , , I 8; c u r re n t h y p ot h e s e s of fu n c t i o n ,
1 89 : s t r u c t u ra l t r a i ts , I , 1 92, 1 9 3 I,
25 8 —9
; p rop e r t i e s a n d f u n c t i o n
en v i ro n m e n t a nd s t ru ct u re , I , o f c h ro m a t i n , , 2 5 9 6 5 ; fu s i o n i n I
1 9 6 ; d i ff e r e n t i a t i o n , , 2 03 ; , I II fe r t i l i z a t i o n, I
2 8 3 4 ; fu n c t i o n i n
,
-
3 5 5—6 1 ; a s o m ot or s y s t e m , v , 2 06 ; I u ni c e l l u l a r r e p ro d u c t i o n , , 5 9 5 —6
; I
v i c a r i o u s fu n c t i o n , , 2 09 ; a c t i i t y I v a b s e nc e o f, II , 2 0—1 ; d i tf u s e d
a nd w a s te , I , 21 6 ; a d a p t a b i l i ty , I ,
for m , II , 85 ; m a c ro a nd m i c ro
2 29 , 23 2 , 2 3 6 ; p a r a l l e l i s m i n c e l l nu c l e u s l n c o n u g a t i on , j II , 452 .
p ro c e s s e s , , 260 2 ; h e r e d i t y , , I -
I N ut ri t i on : o rg a n i c m ol e c u l a r r e -a r
’
3 1 3 ; e fi e c t s o f s e e ra n c e , , 3 49 ; v I r a ng e m e nt , I , 3 6 ; n i t ro g e no u s a nd
re l a t i ve de v e l op m e nt in m e n a nd n o n- n i t ro g e no u s , I ,
4 7 —8 , 68 , 7 1 —4 ,
w om e n, I , 5 9 4 ; a na l y s i s of b ra i n 77, II , 3 6 2 ; foo d a s si m i l a t i o n a nd
s ub s t a nc e , ,I 5 96 ; i n d i v i d ua t ion r e a s o ni ng , I 8 1 ; n e e d fu l for
,
i ta l v
a nd de v e l op m e n t o f, I , 5 98 , 599, c h a ng e, I , 9 4 ; re l a t i o n t o g ro w t h ,
6 00; d i s t ri b u t i on of t a ctua l p er I , 1 4 0, 1 43 , 1 44 , 1 47 9 , -
1 5 0, 1 5 7,
c e p t i ve ne s s , I , 603 8 , 63 3 , 64 6 , 1 61 ; e x p e n d i t u re o f e n e g y I r , ,
6 6 5 —6 , 6 7 1 , al l eg ed cos t 1 5 7, 39 1 ; fl u i d I 2 08 ; v e g e t a l , ,
l i ne s s 66 2 ;
of t i s sue , I , i ns t i nc t f ru c t i fi c a t i o n , I , 267 ; II , 2 66 ;
d e g e ne r a t i o n i n a n t s , l b s e ns a veg et a l g ro w t h a n d g e ne s i s , , I
t i on a re a s ,
"
I 6 66 ; , s e g m e n t a t i on 29 3 , 2 9 4 —7 , 3 3 6 : a n i m a l g r o w t h
in A nn e l i d s , II 1 2 5 ; , e cto d e rm a l a n d g e ne s i s , , 2 89 —9 3 , 2 9 5 —7 , 3 3 6 ; I
d e ri v a t i on , II 3 03—4 ; , c oo p e ra t i ng c o nd i t i o ns ua l i fy i ng a n ta g o n i s mq
S UBJ E CT—I ND E X .
651
of g ro w t h a nd g ene s i s , , 29 9 ; I O rch i d s : p o l l e n p r op u l s i o n , , 57 ; I
c o m p e t i t i on am o ng p a r t s o f a n o r l e a f for m a t i o n i n D end r obi um , , II
g a n i s m fo r, , I
5 6 2, 5 6 6 , 6 76 ; s e x 60—1 ; a er i a l r oo t s a n d p h y s i o l og i
d i ff e r e n t i a t i o n , , 594 —5
; ce ll m ul I -
ca l d i fi e re nt i a t i o n , , 25 5 , 25 7 ; II
t i p l i c a t i o n, I
6 38 ; d i if e r e n t i a t i o n
, fo l i a r s u r fa c e , II , 26 4 .
of ne u t e r i ns e c t s , , 6 5 5 — 6 0, 6 7 0, I O rg a n i c p ro p e r t i e s o f e l e
m a t ter :
6 74 , 6 8 6 8 ; m o ns t r o u s
-
a nt fo r m s, m e nt s , , 3 — I
5 , 2 2 ; of c o m p ou n d s ,
I , 6 8 3 —4 : l e a f d e e l o p m -
v e nt , II , 39, I, 5 —1 3
, 2 5 ; m ol a r a n d m o l e c u l a r
4 2 , 7 3 —8 ; v
e g e t a l d e e l o p m e nt , , v II m ob i l i t y , , 1 2 —1 4
; I
co l l oi d a nd
5 1 — 2 , 1 7 8 , 2 76 ; a i l l a r y b u d s , , x II c r y s t a l l oi d 1 5 —8 , 25 ;
fo r m , I ,
65 —9 , 7 3 —4 ; e ffe c t o n a n i m a l a g g re t h e i r d i ffu s i b i l i t y , , 1 8 —2 1 , 2 6 ; e x I
g a t i o n, II
9 3 ; i n t e r no d e s
, a nd in t r e m e c om p l e i t y , , 2 1 ; l a w s of x I
fl o re s c e n c e , , 1 7 8—8 0; II h el i ca l v
e o l u t i o n a n d g e n e s i s o f, , 2 2 —4 ; I
p h aen og a m i c g r o w t h , II , 1 81 ; ac m od i fi a b i l i ty ,I , 27, 4 4 ; c a p i l l a ri t y
t i on of b i l e , , 3 30 ; II o s s e ou s de a n d os m os i s I 2 8 ; e fife t s f h at , , e o e ,
v e l op m e n t , II 3 49 , , 3 5
3 ; g e ne s i s , I 2 9 ; f l i g h t I 3 0 4 ; n i t og n
, o , ,
— r e
II , 4 1 9 , 42 2 , 42 7 , 4 3 5 —7 , 4 5 2 ; p a r o us, I 3 9 —4 3 ; o x i d a t i on nd e vo
, a
enta l l os s in fe e d i n g y ou n g , II , l t i on o f h
u t I 4 6 6 0; g e n i s ea , , , es
4 24 , 429 ; di v ers e s ou r c e s , II , of l c t i c i t y I 5 0 2 6 0;
e e r n ib le , ,
-
, se s
433 ; C a rp e n t e r on r ep ro d u c t i on m ot i n in I 59 ; t
o s n f m a t i ns
, , ra s or o
a nd , II , 4 6 0; a ni m a l de v e l o p m e nt a n d p e s i t e n c e o f fo r e s I 61 rc , ,
a nd g e ne s i s , II , 465 ; e x p e nd i t u e r m eta b l i s m I 6 2 7 7 ;
o ti fi ci l p o, ,
— ar a r
a nd g e ne s i s , II , 46 8 ; v a ri a t i ons of d c t i o n o f c om p o n d
u I 64 ; c n u s, , o
g e n es i s ; ,
II 4 75 —8 0,5 1 1 ; obe s i t y t ted w i th i n
ra s g ni c m a t t I or a e r, ,
a nd g e ne s i s, II, 4 8 0—4 , 5 1 1 ; g e n 1 07 8 ; i n c m p e h e n b i l i t y o f v i t a l
— o r sx
e ra l d oc t r i n e of g e ne s i s , , 48 6 ; II c h ng a i n I 1 22 ; i n t b i l i t y I
es , , s a , ,
g ene s i s a nd veg et a l p a ra s i t i s m , 1 49 , 5 08 ; p h os p h o r u s i n c e l l -o r
II , 4 8 6 ; a l s o a ni m a l , II
, 4 8 7 —9 0, g a n i z a t i o n, , 2 6 0—1 ; h e t e rog e I
49 5 ; i ns e c t g ene s i s , , II 4 9 0—2 nei t y , I , 35 0—5 ; s p ont a n e ou s g en
g e n e s i s ,
r s um , , é 49 7 —9
; a nd e voé II e ra t i o n a nd e v o l u t i on o f, I , 696
l u t i o n, , 5 0 1 II
—4
; of b l a c bi rd k 7 01 ; c e l l - d oc t r i ne a nd e v o l u ti on
a nd II 5 03 ; g e ne i i n
l i nn e t , ,
s s o f, II , 1 7 —2 1 .
h m n
u ca II 5 08 1 0 5 1 4—6 ;
ra e, ,
-
, O rg a n i z a t i o n ( 8 6 6 S t ru c t u r e ) .
D b l d y on
ou e II 5 1 0—2 ; f t e
a , , u ur O r m e ro d , Dr .
, o n s e x a nd nu t r i t i o n
h m n v l t i on II 5 2 6 5 3 1 ; fl o
u a e o u , , , in wasp s, I , 656 .
l m on s t o i t i s II 5 4 2 54 6 54 7 I
’
ra r s e , , , , . O rt h og e n e s i s , E l m er s t h eor y o f, ,
5 63 —4 .
O BESIT Y , nu t r i t i o n a n d g e n e s i s , II , Os m os i s : o rg a n i c e ffe c t s , I , 2 8 , 29
4 80—4 , 5 1 1 . in a ni m a l s , I , 58 ; in v a s cu la r sys
Od o r i fe r o u s g l a nd s , na tu r a l sel ec tem , II 339 ; in , eg e t a l t i s s u e, v II ,
ti on a nd, I , 5 34 . 5 68 , 5 75 , 5 77 , 5 8 5 , 5 9 2 —6 .
O do u rs : fl ora l fe r t i l i z a t i o n , II , 2 68 O s t e ol og y ( s ee B o n e ) .
9 ; a ni m al p ro t e c t i o n , II , 434 . O vu m ( s ee E g g a nd F e r t i l i z a t i o n) .
O ff s p r i ng : p a r e nt a l l os s il e d by
e nta O w e n, S i r R m e t a g e ne s i s a n d p a r
n u rt u r e , II 42 4 , ,
4 2 9 ; i n u e nc e of fl t h e no g e n e s i s , 273 4 ; fo s s i l I ,
-
age on , II 5 07 , . m am m a l s, ,
I
4 1 0; h u m a n p a r a
Ok e n, L a r c h et y p a l h y p ot h e s i s , II , s i tes , I ,
42 7 ; c o n t i n u o u s o p e r a t i o n
1 22 ; t h e or y of s u p e r n u m e ra r y of c rea t i p ow e r, ve
, 492 ; I fi s s i on
b o n es , II , 223 ; on the s k u l l II , , in I nfu s or i a , , 5 8 4, 5 8 5 , I 595 6 -
5 61 .
p a r t h e no g e n e s i s , , 5 9 2 ; th e o r y o f I
O liv e r, F . W cl as si fi c a t i on of v e r t e b r a t e s e l e t on, , 1 23 , 5 48 k II
p l a nt s , , 3 7 8 —9 I . 66 t h e or y of s u p e r n u m e ra r y
O p h ry ot r och a p unr i Z/i s , c i l i a t i on of b on e s , II , 223 ; E s ch ri ch t on
s e g m e nt s , II , 1 09 . A s ca r i s , II , 4 88 .
652 S UBJ E CT—I ND E X .
Owa l i s : ra d i a l s y m m e t ry , II , 1 52 ; II , 12 ; a p h i s a nd a nt , I , 66 0 1 , -
II ,
fo l i a r s u r fa c e , II , 2 64 . 4 03 , 405 ; as an i nt e g ra t i ng
O xen com p a ri s o n w i th sh eep , I , a g e nc y , II 402 —4 ; i t c m p a a t i v e , s o r
1 5 8 , 1 6 0; c e r e b ro - s p i na l s y s tem , I ,
re c e nc y II 404 ; n t i t i on a n d
, , u r
5 98 .
g e n e s i s i n v e g t a l II 4 8 6 ; i n a ni e , ,
O x i d a t i on ( s ee x
O y g e n) . m al II 4 8 7 90 4 9 3
, , a s t a ti n —
, c r o
Oxyg n : p e ro p e r t i e s , I 3— 5 , 2 0, 22 ;
, p a a s i t a i re
r in c t c ns II r us a ea , ,
c om p o u nd s , I , 6 —7 , 1 0—1 3 , 2 2 , 2 4 —5 '
49 3 —6 .
a c ry s t a l l oi d , I ,
21 ; c om b i n i ng P a r a s ol A nt s , ori g i n of cl a s s es, I ,
p ow er a nd a t om i c w e i g h t , 33 ; I , 68 7 —8 .
o rg a n i c c h a ng e fro m , I , 3 7 ; h eat P a r t h e nog e n e s i s o c c u r re n c e , I ,
g e n e ra t i o n , , 4 6I—9
; p h o s p h o re s 2 74 —5 ; a l t e r na t i ng wi th gam o
c e nc e , I , 4 9 ; n e r e for c e d e p e nd v g e ne s i s , I , 28 9 —9 1 ; w e n o n, O I ,
e nt o n, I , 5 3 ; a ni m a l m e t a b o l i s m , 5 92 ; l a w s o f m u l t i p l i c a t i o n, II ,
I , 7 2 , 7 3 ; n e c e s s a r y t o a ni m a l l i f e , 4 1 5 ; i n a rt i c u l a t e a ni m a l s , II , 44 5 .
I , 94 —5 , 5 7 7 ; a c t i i t y a n d a m o u nt v P a s t e u r, L .
, sil k -
w or m d i sea s es, I ,
i nh a l e d , I , 21 4 . 6 22 3 .
P e a c oc k : t h eo r i e s of h e re d i ty a nd
C
P A K AR D , A . S .
, on ey es of ca ve / of
s t ru c t u r e t a i l fe a t h e r, I , 3 72 3 ,
a ni m a l s , I , 648 —9 , 69 3 . 695 , II , 6 1 8 —9 .
P a g et, Sir J b l o od c h a ng e s in P e a r, fo r e r i g h t s h oo t s , I , 28 7 .
s m a l l - p ox a nd s c a r l a t i na , I , 221 , P e l ori a : i n g l ox i ni a , II , 1 66 ; p h aeno

7 01 .
gam s, II , 1 80 .
P a l ae o n t o l o g y : d i s t r i b u t i on i n t i m e, P e n g u i n, d e rm a l s t ru c t u r e , II , 314 .
I ,4 04—1 1 , 4 1 2 ; s p e c i a l c r e a t i o n, I , P e p s i n, I , 69 .
4 2 5 ; c o ng r u i t y w i t h e vo l u t i on hy P e ri c y c l i c fi b re s o f m o noc o t y l e d o n s ,
p o th e s i s , I ,
4 85 —9 , 5 5 6 ; r e l a t i on s II , 2 78 .
of p r e s e nt t o e x t i nc t s p e c i e s , , II P er i p a t us ca p ens i s , p r ot o p l a s m i c
1 0—1 1 ; s c a rc i t y of r e m a i ns , II , c on t i n u i t y , I , 6 29 .
34 —5 ; s e c on d a r y th i c k e ni ng in P e ri - v i s c e ra l s a c, fu nc t i on a nd dif
p l a nt s , , II
5 6 ; C op e o n o s t e o l og y fe re n t i a t i o n, I , 39 1 .
o f P e r m i a n Ve rt e b r a t e s , II , 2 25 —6 . P er k in , W
'
. H I , vi .
P a ng e n e s i s , D a r w i n
’
s t h e ory o f, I , P e rs i s t e n c e of fo r c e , c o ro l l a r i e s
3 5 6 , 3 5 7 , 3 5 9 , 3 6 0, 3 62 , 3 7 2 . fr om : p r op e r t i e s o f c o m p o u n d s ,
P a nm i i a , x W ei s m a nn s
’
h y p ot h e s i s I , 3; o r g a ni c t ra ns fo r m a t i o n, , I
of : its re l a t i on t o R om a nes
‘
c es 6 0; g ro w t h , I , 1 5 0: o rg a ni c eu
s a t i on of s e l e c t i o n,
"
I , 5 60; ai e rg y , ,
2 2 0; I v a ri a t i o n I 335 ;
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, ,
l eg ed s e l e cti ve p ro c e s s d e n i e d , I , g e ne s i s , , h e re d i t y ,a n d v a r i a t i on
5 6 1 —3 , 6 6 7, 68 5 ; d i s t r i b u t i on of I , 3 5 4 —5
; m o rp h o l og i c a l s u m m a ry ,
t a c tua l p e rce p t i v e ne s s , I , 608 ; II , 23 5 ; v
e g e t a l t i s s u e d i ffe r e nt i a
r u d i m en t a ry eyes of ca va -
fa u n a , t i o n, , II
2 4 5 ; p h y s i ol og i c a l d e e l v
I , 6 1 2 —3 , 64 7 ; R om p ro a ne s on o p m e nt , II , 394 .
ce s s , I , 64 9 , 6 6 7 ; d e g e n e ra t i o n o f P eta l s : fo l i a r h om ol og y , II ,
43 6
s e l f- fe e d i n g i ns t i n c t i n A m a z o n a d na t e , II , 58 .
a nts , I , 6 6 0—2 , 6 7 0; r u d i m e nt a r y P e t re l s , D a rw i n on , I , 45 5 .
l i m bs of w h a l e,
6 85 ; a I , 66 8 —9 , P h ae n og a m s : p ro d u c t i on o f s p e r m a
p u r e s p e c u l a t i o n , , 6 7 1 ; m a r i ng s I k t ozo i d s , I , 1 86 ; m or p h o l o g i c a l
o n l e g b o n es o f P u n a b i s ,
-
, 68 9 j I . com p o s i t i o n, , II
3 7 —7 9
; l e a f t ra n
P a r a m aec i u m :p a r a s i t e i n fe s t i ng , I , s i t i ons , II , 3 7 - 4 2 ; fo l i a r h o m o l o
4 2 7 ; re p r o d u c t i o n , , 44 3 , 45 2 II .
g i es , , II
42 —9
; o ri g i n o f t y p e , , II
P a r a s i t e s : s e u a l d i m o rp h i s m , x I , 4 9 —8 4 ; v
e r t i c a l g ro w t h , , 5 6 —64 ; II
3 1 5 ; l i m i t s t o d i s t ri b u t i on, , 39 7 ; I a x i l l a ry b u d s , , 6 6 ; c ot y l e d on II
s p e c i a l -c r e a t i o n a n d , 1 , 4 2 7 —9 , 4 3 8 ou s g e r m i na t i o n a nd e n d o g e n o u s
r e t rog ra d e de v e l op m e n t I , , 4 5 7, g ro w t h , , II
69 —7 2 ; a i a l h o m o l o x
S U BJ E CT I ND E X -
.
653
g ies, II 7 3 5 ; i re g l a r d e v e l op
,
— r u m e nt , II , 76 ; fa l s e- j oi nt s ,
"
II ,
m nt e II 7 5 —8 ; d g e of c o m
, , e re 3 7 1 —2 ; d i s s oc i a t i o n of a n c e s t ra l
p i t i n II 7 8 ;
os o ep d c ti v e h o
, , r ro u t ra i t s in h y b ri d s , II
6 1 6- 7 ; i n
,
m o l og y II 8 0—4 ; n i n d m l t i
, , u a u h e ri t a nc e of a c q u i re d c h a r a c t e rs ,
a xia l y m m e t y sII 1 4 1 —3 u n i t o f
; r , , II ,
c o m p os i t i n II 1 5 1 h l i ca l
o , , e P h y s i o l og i c a l d i i s i onv of l a b our
g ro w t h II 1 8 1 ; s e c on d a y t h i c k
, , r ( 3 66 L a b o u r ) .
e n i n g II 24 7 ; t i
, e nd l a f d i f
, s su a e P h y s i o l og i c a l S e l e c t i o n, 5 6 9—7 1 I , .
fe e n t i t i on II 24 7 9
r a 38 7 ; a l so , ,
—
, P h y s i o l og y : a nd p s y c h o l og y , , 1 27 ; I
b k a nd
ar m b i um II 2 4 9 5 0 ca , ,
-
, s u b d i i s i o ns , v , 1 28 : i ca ri ou s I v
38 6 ; a l s o o t e ti e II 2 5 2 u r s su , , , fu nc t i on , I , 2 08 ; p r im i ti ve i nte r
2 6 —9 , 2 7 0, 3 8 6 —7 ; w a x d e p os i t o n
5
p re t a t i o n s , I , 41 7 ; m u l t i p l i ca t i on
l e a es , v , II
260- 2 ; d i ffe r e nt i a t i o n of e fi e c t s e xe m p l i fi ed , I , 5 12 , II ,
of i nn e r II 2 7 3 5 38 8 t i s sues , ,
—
, 39 0 ; r e l a t i o ns to m o rp h o l og y , II ,
v c u l s y s t e m d e v e l op m e nt II
as ar , , 3, 2 39 —
4 1 ; e o l u t i o n a r y i nt e r p re v
280—4 38 8 ; i n t e g a t i o n II 29 3 —5
, r , , , t a t i on of p h e n o m e n a , , 24 1 —5 , II
29 6 3 90; i n s e c t fe t i l i
, t i on II -
r za , , 3 84 —9 5 ; u l t i m a t e l n c o nc e i v a b i l i t y
407 ; m l t i p l i c a t i n II 44 1
u 442 ; o , , , of p ro c e s s e s , , 3 72 ; c o r re l a t e d II
g n e s i s a n d g o w t h II 4 5 1
e 45 7 r , , , i nt e g r a t i on a nd d i if e r e n t i a t i on, II ,
a n d d e v e l op m e n t II 4 64 ; a nd nu , , 3 73 .
t i t i o n II 4 7 6 4 7 7 5 1 1 ;
r , b ti , , , su s P h y s i ol og y , A n i m a l : m e t a b o l i s m , I ,
t t i on of
u x i a l fo fol i a r o g a n a r r s, 6 7 —7 7 ; e r t e b ra t e v
i n te r n a l s y m
II 5 4 1 7
,
— . m e t ry , II
1 08 ; t i s s u e d i if e re n t i a
,
P h n m no n l i fe m a n i fes t e d a s I
e o e , , , t i o n i n P r otoz oa , , 299 , 3 8 5 ; p r i II
m a ry t i s s u e d i ffe r e nt i a t i o n , II , 3 00
P h i l o l o g y ( s ee L a n g u a g e ) . 2 , 38 2 , 3 8 9 ; n a t u r a l s e l e c t i o n a n d
P h or oni s , i nd i i d u a l i t y , v II , 44 4 . t i s s u e d i ffe re n t i a t i o n , , 3 04—8 ; II
P h os p h o r e s c e n c e , o rg a n i c , I , 49 . ou t e r t i s su e in C ozl eni er a ta , II ,
P h os p h or u s , 4 ; i n cel l
: a l l ot rop i e , I 3 09 —1 0; r e s p i ra t o ry o rg a ns , II ,
—
p h y s i o l og y , , 2 5 9 6 2 ; c e r e b ra l a c I 3 1 0—1 , 33 3 —8 ; d i ffe r e n t i a t i o n o f
t i vi t y , , I
5 9 6 - 7 ; org a ni c e o l u t i on, v a ni m a l ep i d e rm i c t i s s ue , , 3 1 2
—4 , II
I , 7 03 . 387 ; de v e l op m e n t of t e g u m e nt a r y
P h ot o g e n e s , v i s i bi l i t y o f, I , 21 8 . o rg a n s , II , 3 1 4—6 ; of s e n s ory , II ,
P h y l og e n y : as i n t e rp r e t i n g m or 3 1 7 —2 0: i nn e r a nd ou t e r t i s s ue
p h ol o g y , II fi , 1 0—1 2 ; d i f c u l t i e s o f t r a ns i t i on, 32 1 —2 , 3 8 9 ; a l i m e nt II ,
a ffi l i a t i o n II 34 5 ( 8 6 6 , ,
-
. E m b ry a r y c a n a l d i ffe r e n t i a t i o n, , 3 23 II
o l og y nd E v o l u t i o n )
a . 5 ; g i z z a r d d e e l o p m e nt i n b i r d s , v
P h y s i o l og i c a l U n i t s : ; d e fi n i t i o n, , I II 3 25 ;
, a l i m e nt a r y c a n a l o f r u m i
226 ; g e ne s i s , , I
280 h e re d n a nt s , II 3 2 7—9 ; d i ff e nt i a t i on o f
, er
i ty, I , 3 1 5—9 ; v a r i a t i on I , , 3 3 0, l i ve II 3 29 —3 3 ; f a n i m a l v e n
r, ,
o as
3 3 1 2, -
333, II , 61 9 : s ta bi l i ty , I , l a r sy s tem II 3 3 9 4 4 ; f os eo s , ,
-
o s u
3 4 0: , 6 1 4 ;II
s e l f-
fe rt i l i z a t i on, I , s y t e m II 344 5 5 ;
s , f ne v e t i s
,
-
o r s u e,
3 4 2 —4 , 3 5 3 : i nt e r b r e e d i n g , , 3 45 , I II 3 5 5 —6 1 ; of m l e II 3 6 1 9 :
,
us c , ,
-
35 3 , , II
6 1 5 : r e c a p i t u l a t i o n of h y d i ffe e n t i a t i o n a nd i n t e g
r t i n II ra o , ,
p ot h e s i s , 3 7 3 —6 ; i n v a s c l sy t m II 3 7 6 u ar s e ,
s t r u c t u ra l 9 3 8 3 ; i n n e r v e s II 3 79 8
,
ig in , ,
- ”
or
3 6 9 —7 1 , II o f d e v e l op m e nt II 3 8 4 ; d i ff n , . e re
a na l og y I , , 3 6 4, II , 6 29 ;
/
c om ple x t i a t i on a nd i n s t a b i l i ty of h om o
i ty i n o rg a n i z e d t y p és , I , 3 68 —7 0 g e neou s , , II
3 8 4 —9 , 392 : s u m m a r y
r e -n a m e d c on s t i t ut i o n a l u ni t s , o f d e e l op m e nt , v , 3 84—9 4 ; m u l II
I, 3 69 ; t e l e g o n y , I , 6 5 0;
“
m e
’
t i p l i c a t i on of e fi e c t s , , 3 96 —1 , II
c h a n i c a l t h e or y ,
"
I , 7 01 —6 ; m or q
3 9 2 ; e u i l i b ra t i o n , , 3 9 1 —4
( S ee II .
p h o l og i c a l de v e l op m e nt II , , 7 —9 a lso F u nc t i o n ) .
cel l d o c t r i ne . II , 1 7 -2 l : de vel op P h y s i o l o g y , P l a nt : m e t a b ol i s m , I ,
6 54 S U BJ E CT -
I ND E X .
62 —7 ; t i s s u e d i ff e r e nt i a t i o n i n s ec p l e x i t y o f l i fe , I , 1 03 —4 ; b i o l og i c a l
o nd a ry a g g re g a t e s , II , 2 46 , 38 5 fi
c l a s s i c a t i o n, I , 1 2 5 ; g ro w t h , ,
I
i n p h te n og a m s , , 247 9 , II -
38 6 ; in 1 3 6 , 1 38 , 1 4 0, 1 4 3 , 1 4 5 —9 , 1 5 3 , 1 60
ba r k
a nd c a m b iu m , , II 2 4 9 —5 0, 1 , II , 4 01 —2 ; d e v e l op m e n t I , , 1 63
3 8 6 ; i n f re e a nd fix e d s u r fa c e s , II , 5, 1 6 7 — 70, 2 72 ; w eig ht,
t e m p e ra
25 1 —6 , 2 70, 3 8 6 ; out e r stem a nd t u re, a nd s e l f- m o b i l i t y , , 1 74 ; I
l e a f t i s s ue , II , 2 5 6 —9 , 2 70, 38 6 ; su fu n c t i o n , I , 1 74—8 ; s t r u c t u r e , , I
p e rfi c i a l d i ffe r e nt i a t i o n i n l e a ve s , 1 94 —6 , II , 21 ; a ni m a l s t ru ctu re
II , 2 60—4 ,
2 70, 387 ; fi o r a l t i s s ue c o n t ra s t e d , I , 1 96 ; f un c t i o n a nd
d i ffe re n t i a t i o n , II , 2 65 —9 ; ou t e r t i s s t r uct ure , I , 2 00; v i c a r i ous f un c

s ue, 1 é s um é , II , 2 70: i n n e r t i s s u e t i o n, I , 2 08 —9 ; w a s t e a nd re
d i ffe r e n t i a t i o n , II , 2 73 5 , -
38 8 ; p a i r, I , 2 1 3 , 22 0; p h y s i ol og i c a l
s u p p o r t i ng tis s u e II , , 2 75 9 , 2 85 —8 , u ni t s , I , 2 2 5 —6 , 3 1 7, 3 60; adap
388 ; va s cul a r sys te m de v e l op t a t i o n, I

is an ln
, 22 7 ; w h at
m e nt, II , 2 73 5 , -
279 —84 , 2 8 5 —8 , v
d i i d ua l ? , 2 4 4 —6 , 2 5 0—1 ; g e ne I
3 8 8 : i nne r t i s s u e , s u m m a r y , , II s i s, , I
2 70, 2 7 1 , 2 7 2 —3 , 2 7 4 , 2 7 6 —8 ,
2 88 —9 1 , 3 8 8 ; i n t e g ra t i o n, , 29 2 —8 II 2 79 8 5 ; -
re l a t i on of n ut ri t i o n to
d i ff e r e n t i a t i on a n d i ns t a b i l i t y of g ro w t h a nd g e ne s i s , I , 2 8 4 9 , 2 94 ,-
h om og e n eo u s , II , 38 4 —9 , 3 9 2 ; m ul 29 5 —3 00, 64 2 , II , 39 ; o vu l e h om o
t i p l i c a t i on of e ffe c t s , II 3 90—1 ,
,
l og n e s , I
2 88 ; n a t u ra l s e l e c t i o n ,
,
392 ; e q u i l i b r a t i o n II , , 3 9 1 —4 ; c i r I , 29 4—8 , 5 32 , 5 33 , , 5 1 ; h e re d i t y , II
c u l a t i on a nd w oo d fo r m a t i o n, II , I , 3 01 4 , 3 08 , 3 8 —6 0;
-
5
a ri a t i o n, , v I
5 64—9 7 ; d y e p e r m ea bi l i ty , II , 5 69 3 2 0, 3 2 3 —4 , 3 2 5 —6 ; fe r t i l i z a t i o n
, , I
7 4 , 5 7 7 —8 1 , 5 84 , 5 86 . ( S ee a l so 3 4 0—5 ; c l a s s i c a t i on , fi , 3 7 7 —8 0
, I
F u nc t i on .
) 3 89 —90; d i s t r i b u t i o n, I , 3 9 6 —4 00,
P ic k e ri ng J W , . .
, on a rt i fi ci a l p ro 4 01 —3 , 4 04 - 1 2 , 4 78 9 , -
5 5 6 ; sp e c i a l
t ei d s , I , 39 . c re a t i o n a nd p a ra s i t i s m , , 42 8 ; I
P ig : c o l ou r of m u s cl e s , I , 3 65 —6 ; e v o l u t i on h y p ot h e s i s , I , 4 3 4 , 44 3 ,
t e l e g o ny , I , 627 ; fe r t i l i t y of d o 44 9 —5 0; r u d i m e n t a ry o rg a ns , I ,
m e s ti c a nd wild s ow , II , 479 8 0 . 4 74 , 4 7 5 , 5 5 6 ; v a ri e d m ed ia , I , 4 84 ,
P i g eo n s : foo d o f s ta r v i ng I , , 21 5 ; II , 32 ; a l i e n a n d na t i ve s p e ci e s in
h e r e d i t y a nd va ri a t i o n, I , 3 05 , New Z e a l a nd I , , 477 ; E . D a rw i n
321 , 61 5 ; a ta ism v , I , 3 1 4 ; fe r t i l i t y , a nd L a m a rc k
-
on e v o l u t i on o f, I ,
II, 4 7 1 —2 , 4 78 . 490 8 ; g e o l og i c c h a ng e s a fi e c t i ng ,
-
’
P ik e,un c e a s i ng g ro w t h , I , 1 54 , 2 9 2 . I, 5 01 —3 , 5 5 7 ; i n t e rd e p e n d e n c e o f
P i q ue g -
o u ffc , c o m m e ns a l re l a t i on s a ni m a l s a nd , I , 5 04 —6 , 5 1 4, II ,
w i t h b u ffa l o , II , 403 . 3 98 ; c o m p l e i t y x of in fl u e nc es a r
P l a g i oc h i l a , e v ol u t i on of s te m , II , t e e t i ng , 5 06 ; I , dir ec t e q i l i b ra u
62 . t i o n, I
5 2 3 —5 ; i n d i r e c t ,
, I , 5 3 2,
P l a na r i a : i n t e g ra t i o n , II , 1 01 —2 ; 5 33 ; seed d i s t r i b u t i o n, I , 546 ;
M o rg a n on r e g e ne ra t i on, II , 1 02 , w ood d e v e l op m e nt II , , 28 5 —7 , 28 9 ,
61 1 ; s e g m e n t a t i on, 1 07 ; s y m II , 5 6 7 9 7 ; i n t e rd e p e nd e n c e ,
-
II 4 02 ,.
m e t ry , II , 1 95 : u ni n t e g ra t e d fu n c 3 4 04 ; i ns e c t e l a t i n
, II 4 06 —7 ; r o s, ,
t i on , II , 373 . a d a p t a t i o n a nd m l t i p l i c a t i on II u , ,
P l a nt s : in fl u e nc e of h ea t, I , 29 ; 4 1 1 —6 ; rh y t h m in n m b r II u e s, ,
e ffe c t of s o l a r ra y s , I , 3 1 —6 , 500, 4 1 9 ; g r w t h a nd o x l g e ne i a se ua s s,
5 5 7 ; c h e m i c a l c o m p o s i t i o n, I , 40 II 4 3 9—42 ; g ow t h a nd e u a l g ne
, r s x e
1 ; h e a t g e n e ra t i o n , , 47 ; p h os I si II 4 4 8—5 1 ; x p e nd i t e II 4 6 7
s, , e ur , ,
p h o r e s e e nc e , I
, 4 9 ; e l e c t ri c i t y , ,
I h o r t i c u l t u r e , n u t ri t i o n , a n d g e n e
5 1 ; s e ns i b l e m o t i o n, , 5 6 —7 5 8 I , s is, , II
4 7 7 ; t re e d e e l o p m e n t , , v II
m e ta b o l i s m , I , 6 2 —7 , 7 0; v i ta l 5 3;
5
c i r c u l a t i o n a nd w o o d fo r m a
c h a ng e s , I , 8 6, 8 7, 9 1 , 94 ; s i m ul a t i on, II
5 6 7 —9 2 ; d y e p e r m e a b i l i t y
,
t i on b y c ry s t a l s , , 96 ; I v i ta l a d a n d c i rc u l a t i on , , 5 69 —7 4 , 5 7 7 —8 1
, II
ju s t m e nt s , , 1 02 : l e n g t h I a nd c o m 5 84 , 586 ; r és m é u on c i rc u l a t i o n
S U BJ E CT—I ND E X 65 5
a nd w oo d for m a t i o n, II , 5 9 2—7 . P re s e r v a t i on : f e rt i l i t y a nd s el f I ,
( S ee a lso M u l t i p l i c a t i on ,
M or 58 1 ; II , 42 3 , 4 30; nu t r i t i o n , II ,
p h o l o g y , a nd P h y s i o l og y ) . 493 .
P l a s m o d i u m , d i s s ol u t i on of, I , 1 85 . P r og r e s s ; i ts L aw a nd C a u s e,
P l a t o, ui é a. o f, II , 550 . t h e o ry of sp ecies d i ffe r e n t i a t i o n,
P l a ty h el m i n th es : t ra n s v e r s e fi s s i on , I , 5 68 .
II, 1 01 ; s e g m e n t e d a n d n o n s e g -
P j
ro e c t i l e s , fa c t o r s i n flight o f, I ,
m ent e d t y p es , , 1 02 , 1 07 ; s y m II 45 0 1 -
.
m e t ry , , II
1 9 5 , 1 9 7 ; m u l ti p l i ca t i on P r ot e i d s : m e t a bol i c fu n c t i o n , I , 6 7,
a nd g ro w t h , II , 4 8 8 —9 . 68 , 69, 72, 76 ; c om ple xi ty of
P l e t h ora , fe r t i l i t y a nd , II , 48 0—4 , m ol e c u l e , I , 12 2 .
51 1 . P r ot e i n : e v ol u t i o n I , , 23 , 2 4 ; i s om er
P le u r ococca ccce , u ni c e l l u l a r fo r m , II ,
ism , I , 7 00, 703 , 7 04 .
21 ,
P r o teu s , d e g e ne ra t i o n of e y e , I , 61 3 .
P l eu r onec tid az: s y m m e t ry a nd l oca P r otod r i l us , i nt e s t i ne s e g m e n t a t i on ,
t i on o f e y e s , II , 2 05 ; ou t e r t i s s u e , II , 1 25 .
II , 38 7 . P r otop h y ta : i n t e r na l m o v e m e nt s I , ,
P l um a te l l a : m e t a g e ne s i s , I , 277 56 ; l im it of g ro w t h , , I 1 3 8 ; d e ve l
s y m m e t ry , II , 1 95 . o p m e nt , I , 1 64 ; s t r u c t u r e , I , 1 73 ,
P od os tem a cew, u nd e v e l op e d c i r c ul a 1 8 1 —3 ; s e l f -
m ob i l i t y , I 1 75 ; i nd i
,
t o ry s y s t e m , , 274 II . vi d ua l i t y , I , 245 ; m u l t i p l i c a t i o n,
P o l a r b o d i e s , h y p ot h e s i s c o n c e r ni ng I , 2 70, 2 76 , 2 79 , 5 8 1 , 58 4— 5 , II ,
x
e tr u s i o n o f, , 2 66
—8 I . 4 3 9 , 4 6 2 ; g e ne s i s a nd nu t ri t i o n I , ,
P o l a ri t y , o rg a n i c , of p h y s i o l og i c a l 2 9 5 ; u ni c e l l u l a r, II , 21 ; c e nt ra l a g
u ni t s , I , 22 6 , 3 1 5 , 3 1 7 , 3 3 2 , 3 504 , g re g a t i o n , II 24 ; y m m , s e t ry , II ,
7 01 -
6 . 1 34 ; t i s s u e s , II 2 44 2 4 9 ; , , p ri m a ry
P ol y a t o m ic com p ou n d s ( s ee C h e m d i ffe r e n t i a t i o n II 3 8 5 ; p , , ri m o r d i a l
i s t ry ) . t yp e, II , 398 ; sy m b i os i s , II , 4 00 .
P ol y ch w tce , a n o m a l ou s de v e l o p m e nt P ro t op l a s m s e l f- i nc re a s i ng func
in M y r ia n i d a , I , 3 61 . t i on p ri m of ord i a l , I , 6 3 —4 ; p l a n t
P o l y cy th a r l a , i n t e g r a t i on, II , 9 0, m e t a b ol i s m , 6 5 —7 ; c o m p l e i t y , I , x
I, 1
2 2 , 25 3 —5 ; d i ffe re n t i a t i o n i n s i m
P o l y g a s tr i ca , a g g re g a t i o n , I , 58 6 .
p l e o r g a n i s m s , , 1 8 2 —3
; c o n t i nu i t y I
P o l y m e ri s m : o f c o m p o und s , I , 9, 1 1 ,
a nd i nt e r -
c i r c u l a t i on, I , 1 9 0—2
25 ; n e r ve t i s s ue, II , 356 . 3 7 1 , 629 , II , 21 , 6 20 s t re a m i ng ,
P o l y p or l , s y m m et ry a nd en v i ron I , 2 5 3 ; s t ru c t u re , I , 25 3 —5 .
( S ee
m e nt, II , 1 39 . a ls o C e l l ) .
P ol y p s ( s e e C azl en ter a ta ) . P r o tozoa : i n o r g a ni c c om p o ne nt s , , I

P ol yz oa : I 1 40; m l t i a x i a l d e
s i z e, ,
u 1 7 ; l o c o m o t i o n , , 58 , I 1 75 , , 14 II
v e l p m e nt
o I 1 65 ; t c tu a l i n, , s ru r v i t a l c h a ng e s s h o w n by, I , 94;
d efi nit n I 1 7 3 f n t i o na l d i f
e ess , , : u c l i m i ta ti on of g row t h , , 1 38 ; d e I
f e re nt i t i on I 202 ; t h o p h o l
a , , ro c ra v e l o p m e nt , I , 1 64 ; s t r u c t u r e , , I
k i n h i p I 44 7 ; i nt e g t i n II
s , ,
ra o , ,
1 73 , 1 8 1 — 3 ; i n c i p i e nt d i fi e r e n t i a
9 6 1 2 4 ; s y m m t y II 1 9 4
,
e r , , , t i o n, I, 1 98 , 391 , , 29 9 , 3 09 ; m u l II
2 07 : va c l syst m II 3 40;
s u ar e , ,
t i p l i c a t i o n, I , 2 70, 2 76, 2 79 , 2 8 0,
g m m t i n II 4 44
e a o , , .
II , 442 , 4 5 1 2 ; g e n e s i s a n d
-
P oo r L a w s, a nd na t u r a l s e l e c t i o n, n u t r i t i o n, I , 29 5 ; d i s t r i b u t i on, , I
II , 5 32 . 3 9 6 ; p a r a s i t e s i n fe s t i n g , ,
42 7 I
P op u l a ti on, A Th eory o f , I , 2 65 , 5 7 7 W e i s m a nn s h y p ot h e s i s o f i m m or
'
601 , II , 41 1 . ta l i ty , I 63 7 ; s p o n t a n eo u s g en
P ot a t o : g ro w t h ,
s i m ul a te d I , 1 36 ; e r a t i o n, 6 9 7 4 01 ; I ,
n o n- n u c l e
v i c a r i ou s fu n c t i on o f t u b e r . I , 2 09 , a t ed, , II
2 0; p r i m a r y a g g r eg a t e ,
II , 255 : s ub - s p e c i e s , I , 3 02 ; (l ye II , 8 6 —7
,
1 2 4 : p r o g re s s i n g i n t e g r a
a b s o r p t i o n, II , 2 79 . t i o n, II , 89—91 , s y m m e t ry , II ,
65 6 S U BJ E CT -
IND E X .
1 8 6 ; p r i m ord i a l p l a nt -
a ni m a l ty p e, P ter op oda : b i l a t e ra l s y m m et ry , II ,
II , 39 7 —8 : s y m b i os i s , II , 4 00 . 2 01 ; d e r m al r e s p i ra t i on , II , 31 0 .
P ro t y l e , h y p ot h e t i c a l c h em i ca l uni t, P t y a l i n e, m e t a b ol i c fu n c t i o n , I 69 , .
I , 22 , 2 3 . P un a bi s, j i nh eri t a n ce of ac q i ed u r
P s eu d - a xi a l de v e l op m e n t v e g e t a l , , o s t e o l og i c a l
p e c ul i a ri t i e s , , 6 89 I .
II ,
28 9 , 3 0
-
. P y r os om i daz: p h OS p h or es c e n c e , , 4 7 ; I
P s e u d o - fo l i a r de v e l op m e n t v e g e t a l , , i n t e g r a t i on , , 5 88 , , 97 I II .
II , 2 6 —8 , 30 .
P s y ch i d w : p a r t h e n og e n e s i s , I , 275 ; ! U AG GA , t e l e g on i c t r a n s m i s s i on of
x
s e u a l d i m or p h i s m , , 68 3 I . m ar k i ng s t o o fi s p r i n g o f m a re , 1 ,
P s y c h o l og y : r e a s on i ng a nd d e nn i 6 2 4 , 6 2 7 , 64 6 .
t i on o f l i fe , I , 8 1 —8 ; c or r e s p o nd ! ui l l s , de v e l op m e nt II , , 3 1 4 —6 .
e nc e s h ow n by r ecog ni ti o n, I , 95 ;
c on t r a s t e d p h y s i ol og y , ,w i th I RA B BIT : v i t y a nd m u s c l e c ol
a ct i
1 2 7 ; d e p a r t m e n t s o f, , 1 27 8 ; I — o u r, II 3 65 ; o v e r r u nni ng c h e c k e d
,
-
v i c a r i o u s fu n c t i o n , , 2 09 ; w a s t e I by w ea s e l s , II , 4 05 ; e x p e nd i
a nd rep a i r in s e ns o ry o rg a n s , I , t u re a nd g e ne s i s , II , 4 72 .
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23 1 , 23 2 ; i nh e r i t a n c e of s e n s or y R a d i ol a r i a : unic e nt l d v ra e e l o p m e nt ,
d e fe c t s , I , 3 06 ; m u s i ca l t a l e nt , I , I , 1 63 ; s e c o n d a ry a g g r e g a t i on , II ,
3 1 1 —2 ; i n t e l l e c t u a l p r og r es s a nd 88 ; s y m m et ry , II , 1 87 .
s p e c i a l -c r e a t i o n h y p o t h e s i s , I , 41 7 ; R a d ul a , d e v e l op m e n t o f root s f ro m
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I l ea fl e t s II , , 34 .
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v
e o l u t i o n -h y p o t h e s i s , ,
4 33 —5, I t i s s u e d i ffe r e n t i a t i o n , II 2 7 4 ; nu
,
439, 5 54 ; em b r y ol og y of idea s, I , t r i t i on a nd g e n e s i s , II , 48 6 .
4 5 0, 45 7 p e r s i s t e nt fo r m a ti ve R a t ( s ee R od en t i a ) .
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D a rw i n s
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a nd L a m a rc k s t h e or y o il II ,
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6 92 ; i n c o n c e i v a b i l i t y of t h e n e g a j v
R e u e n e s c e n c e , a n d s e u a l fe rt i l i x
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org a ns , 3 1 7 , ; II
—20 d i ffe re n t i a t i on
Rep a i r : co nt i n u i t y o f, I
2 1 6 - 9 ; a ni
,
of ner ve
II 3 5 5 —6 1 : f nc
t i s s u e, , u m a l i n ur i e s , j I , 21 9 , 2 22 —4 , 1 02 , , II
t i o na l i nt e g a t i on II 3 7 6 ; lso t r , , a u 61 1 ; dedu ct i v e i nt e rp re t a t i on , I ,
t e g a t i o n II 38 0—2 : e q i l i b a t i on
r , , u r 2 2 1 —2 ; t h e or i e s of h er ed i t y a nd
o f n e v e d i s h a g e II 3 9 3 ; h m
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fe t i l i t y
r nd ne v e d v l p m nt a r e e o e , R e p e t i t i on of li ke p a rt s , 1 26 , II .
II 4 6 6 5 3 2 ; fu t e h u m n e ol
, , ur a v u R e p ro d u c t i on ( s ee M u l t i p l i c a t i o n) .
t i o n II 5 2 3 —, 5 5 27 ; h u m a n e o l
, , v u c ti l l a : g r o w t h a n d e x p e n d i t u re of
t i n nd g e n e s i
o a II 5 2 9 3 1 ; f t re s, ,
-
u u forc e , I , 1 42 ; s i zes of o va a nd
m n t l d e v l p m e n t II 5 3 5 ; o i g i n
e a e o , , r a d ul t, I , 1 44 ; l on g e i t y of v c ro c o
f ve t b at
o t y p e II 5 9 8 6 00
r e r e , ,
— . d i l e, I ,
1 5 4 ; t e m p e ra t u r e , I , 1 74 ;
P t i d p hy t
er o i e tt in d by a : I s z a a e , , w a st e, I , 2 1 4 ; d i s t i nc t i ve c h a ra c
1 3 8 1 3 9 ; h o m o l og i
, II 80—1 8 2 ; es, , , t e rs , I , 3 9 2 ; d i s t ri b u t i on i n t i m e ,
f n d s u r fa c e d l if e e n t i t i on II
ro r a , , I ,
4 09 , 41 2 ; e rt e b ra l v s e g m e nt a
t i o n, I , 47 0: r u d i m e n ta ry l i m b s o f
S U BJ E CT —I ND E X 657
s na k es I , , 4 7 3 ; fe r t i l i t y a n d d e e l v 7 1 ; p a nm i i a , x I , 649 , 66 7 ; i n fl u e nc e
o p m e nt , I , 5 8 3 , 5 9 8 , 5 99 ; r e g e n e r a of a p re i ou s v s i r e on p r og e n y , I ,
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2 03 , 204 ; C op e o n s e g m e n t a t i o n i n 34 ; p h y s i o l o g i c a l d i ffe r e n t i a t i on,

x
e t i nc t , , II
2 2 5 , 22 6 ; a c t i i t y a nd v II , 253 —5 , 2 7 0; n u t r i t i on fr o m
m u s cul a r c o l o u r, II , 3 65 ; fu n c l ea v e s II , , 2 74 ; si z e a nd fu nc t i on ,
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R e s p i ra t ory S y s t e m : e ffe c t o f l i g h t , R ou x W , . :
“
i nt r a - s e l e c t i o n , I , 6 76 ;
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-
I fu n c t i o n a l a d a p t a t i o n, II , 354 .
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l u ng s , I , 25 4 ; em b r y on i c b r a n c h i ae of l i fe a nd , I , 1 12 , n a t u ra l s el e c
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t i a t i o n, II , 3 1 0—1 , 3 3 3 —8 ; p h y s i o 3 09 , 6 1 2 —4 , 64 7 —9 , 6 9 3 ; e v o l ut i o n
l o g i c a l i nt e g r a t i on , II 3 7 4—5 , 3 8 2 ;
, h y p ot h e s i s , I , 5 5 6 ; l i m bs of
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.
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Re v ers ed S e l e c t i o n, I , 61 1 , 61 2 . S a lm om
’
d w, r e p r o d u c t i on a nd g ro w t h ,
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R h i z op od a : 1 7 3 ; u nd i f
s t ru c t u r e , I , S a r ci na : c e nt r a l a g g r e g a t i o n , II , 24
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m a ry a g g re g a t e , , 86 ; sy
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”
I , 31 3 .
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R i i nu s , p l a n t c l a s s i fi c a t i on I 3 7 7 , , . 1 08 .
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s p eci es d i ff e r e n t i a t i on, , 569 ; I c h a ng e of m ed i a c a u s ed by, , I
p h y s i o l og i c a l s e l e c t i o n.
"
I : 5 69 “
4 8 1 ; g e o l og i c i n u e nc e , fl I , 5 02 .
88
658 S U B E CT—I ND E J X .
S e a l s : na i l b e a r i ng t oe s , -
I ,
473 ; vi S h a rp , D . : on i ns e c t s om i tes, I , 4 69
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I 2 99 ; v a i a t i ns
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II 48 4 5 ,
-
. F re n c h b re e d s , I , 6 2 5 ; n ut r i t i o n
S e d g w i c k A d a m : on , c o nt i n u i t y of a nd g e ne s i s , II , 480 .
p r o t op l a s m i n a ni m a l s , ,
1 90, I S h e r ri n g t o n, P ro f on e ffe c t s of
62 9 , , II
2 1 ; z oo l og i c a l c l a s s i fi c a ne r ve se v e ra nc e I , , 3 49 .
I
t i o n, , 3 8 7 ; d i s c ri m i n a t i o n o f s p e S h i p b u i l d i ng
-
i n t e r d e p e n d e nc e , of
cies in em b ry o n i c sta g es , I , 461 ; s oc i a l fu n c t i o n s , , 2 3 7 —9 , 2 4 1 I .
p e rs i s t e n c e o f a n c e s t r a l t r a i t s , , I S hi p l ey, A . B . : s e g m e nt a t i on of
4 63 4 ; A rc h i a nn e l i d a n s e g m e n t a
-
M ic r os tom i da , II , 1 02 ; P r ot od r i l us ,
t i o n, II 1 09 , . II , 1 25 .
S e d g w i c k W m : h e re d i t y a n d s x
, . e , S i l i ca, c o l l o i d a nd c r y s t a l l oi d , I , 16 .
I 3 05 3 1 4 ; t l g oni c t a ns m i s i n
, , e e r s o S i l i ci c a ci d : p rop e r t i e s , I , 1 6 ; i s om
of h y p o s p a d i a s I 64 6 , ,
. e ri s m , I , 59 .
S e e d s : n i t rog e n ou s I 4 0; t e m p e a , ,
r S i l i c o n, a l l o t r op i c , I 4 , .
t e o f g e m i n a t i ng
ur I 47 II 6 1 5 ;r , , , , Sil k w or m d i sea s e, I 6 2 2 —3
, .
v i t a l i s m n d l a t e nt v i t a l i t y o f I
a , , Sim u l a t i on : of h om o l og y by a na l
1 1 6 7 ; v a i a t i o n i n e n v i on m n t
— r r e , og y , II
48 5 ; o f s e g m e nt e d
, 1 4,
I 327 ; na t ra l s e l e c t i o n a m ong I
, u , , s t ru c t u re b y m o l l u s c s , , 1 16 II .
53 2 . S i p h onop h or a , s p e c i a l i z a t i o n of c o m
S eg m e nt a t i o n (m et a m e ri c) : s p eci a l p o n e nt p o l y p s , , 95 II .
c r e a t i on h y p ot h e s i s , I , 4 68 —9
; S i r e m a , s i m u l a t e d fi s h for m
’
,
I , 48 5
.
H u l ey x o n nu m b er o f s om i tes i n S i z e ( s ee G r o w t h ) .
h i g h er a rt i c u l a t e s , i h ; i n a nn u
. k
S e l e t o n, e r t e b ra t e v (s ee Ver te
l os e a ni m a l s , II , 9 8 1 1 0, 1 1 1 —5 ,
-
bra ta ) .
601 —5 ; s i m u l a t ed m o l l u s c a n, II , k
S in : r e s p i ra t o r y fu n c t i o n , I , 2 09
1 1 6 ; in v e rt e b ra t e s II , , 1 2 5 —7 , 2 2 5 —7 , a da p ta b i l i ty , I , 22 8 , II , 3 1 2 —4 ,
606 —7 ; i n e l a s m ob ra n c h s , II , 1 26 . 38 7 ; t ra n s m i t t e d p ec ul i a ri t i es , I ,
S e g re g a t i o n : o f g ro w t h , , I 1 36 ; of 3 06 ; W a l l a ce o n d i s t ri b ut i o n of
li ke u ni ts , I 1 7 9 ; o rg a n i c r e p a i r,
, s e ns i t i v e ne s s I , d i fi e re n , 64 6 - 7 ;
I , 22 1 ; v a r i a t i o n, , 3 3 1 , 334 ; I t i a t i o n, II , 2 1 5 , 2 1 7 , 304—7
; t eg u
h e t e ro g e ne i t y , a nd d e fi ni t e n e s s of m e nt a ry v
d e e l op m e n t , , 3 1 4—6 , II
e v o l u t i on I , , 5 1 4—6 , 5 1 7 —8 ; m or 3 8 7 ; d i ffe r e nt i a t i on of s e n s o r y o r
p h o l ogi c a l d e e l o p m v e nt , II , 7 —9 ; g a ns, , 3 1 7 —2 0; a nd II m u c ou s
p h y s i o l og i c a l u n i t s , II , 61 6 . m e m b ra n e, 3 03 —4, 3 2 1 —2 , 3 8 9 II .
S e l f fe r t i l i z a t i o n ,
-
a ni m a l a nd y eg e S k i n fr i c t i o n a n d l o c o m o t i o n , of
ta l , I , 3 4 1 —4 , 3 5 3 . a q u a t i c a ni m a l s I 1 56 , , .
S e n s e s , t h e ( s ee P s y c h ol og y ) . S k u l l ( s e e Ve te br a ta ) r .
S e x : i n A s c i d i a n c ol on i e s , I , 24 7 S l e e p r e p a i r fa v o u re d b y I 2 1 6
, , , .
l i m i ta t i on h e re d i t y b y , , 3 1 4 —6 ;
of I Sm a l l -p o x , b l oo d c h a ng e s fr om , I ,
c orre l a t e d t ra i t s , , 3 7 1 —2 , 5 1 3 ; nu I 22 1 .
t r i t i on a nd d e t e rm i na t i on o f, in S m ith , P ro f . W .
, on fe r t i l i t y of
s oc i a l i ns e c t s , I , 6 5 5—60, 6 78 84 -
, d i a tom a c ecr , II , 440 .
6 8 6 —9 ; ne u ra l a n d h ae m al t ra i t s , I , S m i th , W . P .
, on t e l e g o ny i n c a l ve s
6 8 3 ; d i ffe r e n t i a t i o n o f o rg a n s , II , a nd foa l s , I , 645 .
3 03 ; c a s t ra t i o n a nd g ro w t h , , II S m i th , W W . on h a b i t s o f T etra
459 ; J ul i n on c a s t ra t i o n p a r a s i m or i a m , I , 66 0 .
t a i re in c rus t a c ea ns , II , 4 9 3 —6 ; S na ke s ( s e e R ep t i h a )
‘
the j
ob e c t o f fe r t i l i z a t i o n, II , 61 3 . S oci a l o r g a ni s m ,
”
a u t h or s
’
essa y
( S ee a lso F e rt i l i z a t i on .
) o n, I , 3 63 , 6 7 6 .
Se x ua l S e l e c t i on ( s ee N a t u ra l S e l e c S oc i ol og y : en v i ro nm e nt a nd d e g re e
t i o n) . of l i fe , I , 1 05 - 6 ; fu nc t i o na l dif
S U BJ E CT —I ND E X .
65 9
fe re nt i a t i on, I , 2 04 ; d i i s i o n o f v S p e rm a t o z oa
I ( 8 6 6 F e r t i l i z a t i o n)
3 6 3 —4 , 3 6 7 ; fu n c
.
l a b o u r, , 2 07 , S p e rm -
cel l
t i on a l i nt e r d e p e nd e n c e , I , 23 7 9 , -
S p h e re : t e n d e n c y o f u ni t s t o fo r m ,
2 4 0—2 ; a u t og e n o u s d e e l op m e n t v of I 1 5 ; t h e e m b r y o n i c for m
, ,
I , 1 77 ;
u n i t s i n c o l on i e s , , 3 64 , 3 6 7 —8
, I II , s y m m et r y , , 1 31 II .
6 2 0; b e l i e f i n s oci a l e v o l u t i on , I , S p h e r oi d , s y m m e t ry , II , 1 32 .
4 3 2 ; na t u r a l s e l e c t i o n, I , 553, II , S p i d e rs ( s e e A r a ch n i d a ) .
5 32 ; i n t e g ra t i o n a n d d i ffe r e nt i a S p i ne ( s e e Ver te br a ta ) .
t i on, , 3 7 8 —9 II
; e fi e c t s of p op u l a
’
Sp on g e : s t ru c t u r e a n d d y na m i c el e
t i o n, , II
5 3 5 6 ; e u i l i b ra t i o n , ,
-
q II m e nt i n l i fe , I , 1 1 9 ; m u l t i c e n t ra l
537 . de v e l o p m e nt , I , 1 6 4 ; u n i t s a nd a g
S o i l , d e p e nd e nc e of p l a nt e v o l u t i on g re g a t e , , I
1 85 ; r e p ro d u c t i e t i s v
o n, , II
4 02 . s u e, , I
28 3 ; i n t e g r a t i on , , 5 8 6 ; ,
I II
S ol a ri um ja sm i noi d es , org a n s of at 90, 38 3 ; p h y s i o l o g i c a l d i ffe r e n t i a
t a ch m ent, II , 276 . t i on , II , 3 00, 386 ; de v e l op m e nt
S ol a r s y s t em ,
a u t og e no u s de v e l op a nd g e ne s i s , II ,
4 63 ; a n a l og y fr o m ,
m e nt i l l u s t ra t ed by d i s t r i b u t i on II , 5 76 .
of fo r c e s i n , I , 3 66 . S p o n t a n e ou s g e ne r a t i o n : a nd h et e
S ol e, s y m m e t ry a nd l o c a t i on of r og e ne s i s , , 2 7 0; a n d I e v ol u t i o n ,
ey es, II , 2 05 . I , 6 9 6 — 7 01 , 7 03 .
p l a s m , W e i s m a nn s t h e or y o f St a g ,
’
S om a -
h or n s a nd cor re l a t e d s t ru c
d i ffe re n t i a t i on fr o m g e r m p l a s m , -
t u re s , I , 5 6 7 , 6 7 0, 6 7 6 —7 , 6 9 2 .
I , 3 5 7 , 6 22 , 6 2 8 3 0, 6 3 3 —4 4 -
. Sta m e ns , a nd fo l i a r h o m o l og y , II ,
S o m i t e s ( s ee S e g m e nt a t i on ) . 44 .
S p e c i a l c re a t i on : a nd e v ol u t i on I , , S ta rch e s : p ro p e rt i e s , , 1 1 ; t ra ns I
4 1 2, 41 5, 4 3 1 ; i m p ro b a b i l i t i e s , , I for m a t i on s , , 6 6 , 68 , 6 9 , 7 0, , I II
4 1 8 —9 , 4 3 0, 4 3 9 , 5 5 4 ; i n c o n c e i v a 593 .
b i l i ty , I , 42 0, 42 9 , 4 3 1 , 5 5 4 ; of ia S t a r fi s h e s ( s e e A s ter oi d ea )
-
.
d i vi d ua l s a nd s p ec i e s , I , 4 2 1 —4 ; S t a t ob l a s t s , o f P l um a te l l a , I , 277 .
t h e i m p l i ca t i o n o f b e n e fi c e n c e , , I S t e e ns t ru p , on A l t e r n a t e G e ne r a
4 2 5 —9 ; s u m m a r y , 1 , 42 9 , 5 5 4 ; V on t i on, I , 592 .
’
B a e r s fo r m u l a , , 4 5 1 —6 ; e rt e I v S t e r i l i t y ( s ee M u l t i p l i c a t i on) .
b ra t e s k e l e t on , II , 551 , 5 5 6, 5 65 . k
S t i c l e b a c : ov a , , k
4 5 4 ; both r i a II
S p eci es : a d a p t a t i on a n d s t a b i l i t y , I , cep h a l u s i n, II , 49 0 .
242 ; h e r ed i t a ry t ra n s m i s s i o n , I , S tom a ch ( s ee A l i m e n t a r y c a na l ) .
3 01 —4 ; v a r i a t i on in wi ld a nd cul ti S t om ata , d i s t r i b u t i o n, II , 2 6 0—1 .
va t ed , I , 3 2 3 —5 , 3 26 , 693 ; g am o S t r a i g h t l i ne , a nd e v o l u t i o n h y p ot h
g e n e s i s a n d l i fe o f, I , 3 4 7 —9 ; p h y e sis , I ,
433 .
s i o l og i c a l u n i t s , I , 3 62 , 3 64, 3 69 S t ra i n : com p r e s s i o n a n d t e n s i o n of,

71 , 458 , II ,6 1 3 ; i n d e fi ni t e n e s s , I ,
I , 1 51 , II ,
2 0 9 —
1 2 ; re l a t i o n t o
3 8 9 , 4 45 , 5 7 2 ; s p e c i a l c re a t i o n , I , m a ss , , 1 5 I
5 —7
; e g e t a l s t r u c t u re , v
4 22 —4 ; i n s t a b i l i t y o f h o m og e n e o u s , II , 5 7 4—88 , 5 9 2 —6 ; o r i g i n of v e r
a nd d i f f e r e n t i a t i o n o f, I , 5 09 — 1 1 , t e b ra t e t y p e , II , 6 00 .
5 1 5, 5 1 7 —8 , 5 5 0, 5 5 7 ; p e rs is t ence S t r a w b e rr y : m ul ti a xia l de v e l op
o f, I , 5 1 6, 5 1 8 , II , 1 0—1 1
; n a t u ra l m e nt , I , 1 66 ; m u l t i p l i ca t i on, II ,
s el e c t i on a nd eq u i l i b ra t i o n I , ,
441 .
553 , 557 n on a d a p t i v e c h a r
-
S t r e n g th , a v i ta l a t t ri b u t e , I , 5 78 .
a c t e rs , I ,5 65 ; m o r b i d p ro d u c t s a s S t ru ct u re : a p p l i a nc e s fo r g e n e r a t
m ar ks o f, ,
I
5 6 7 ; m i g r a t i o n a nd i ng m ot i on , I , 7 5 —7 ; b i o l og i c a l
i s ol a t ion as ca us es of d i ffe r e n t i a cl a s si fi c a t i on , I ,
1 2 5 —7 , 12 9 ; s i z e
t i on, I
568 — 9 ; i n c r e a s i n g m
, ulti a nd or g a ni c, I , 1 37 ; g ro w t h a nd
fo r m i t y o f a g g r e g a t e , , 396 II . co m p l e ity, , 1 38 , x I 1 45 , 1 6 1 ; re l a
fi
S p e c i c g r a i t y , o f o rg a n i s m s v a nd t i o n t o e n i r o n m e nt , v I
1 7 2 —8 , 1 9 5 —6 ;
,
en v i ronm e nt I , , 1 74 , 1 7 7 . o f u ni c e l l u l a r or g a n i s m s , , 1 8 1 —3
; I
6 60 S U BJE CT—I ND E X .
m u l t i c e l l u l a r, I ,
—
1 83 9 6 ; H e r t w g s
i
'
s i on , I , 5 3 0, 61 0 .
( S ee N a t u ra l
cl a s s i fi ca t i on of t i ss ue s, 1, 1 89 ; S e l e c t i on ) .
c o nt i nu i t y o f u ni t s , I,
1 9 0—2
; sy s S w a n, v
e r t e b r ae of n e c k II , , 1 23 .
t em s of o rg a n s , I ,
1 92 ; d i i s i on v S w i ft n e s s , a v ita l a t t ri bu t e , I ,
i nt o u ni v e rs a l a nd p a r t i c u l a r, , I 5 78 .
1 9 3 —4 ; g e n e ra l t ru t h s , , 1 94 5 ; I -
S yl lis r a m os a , l a t e r a l b ra n c h i ng , I ,
p l a nt a nd a ni m a l , c on t ra s t e d , I ,
1 66, 3 61 , II 1 05 1 08
, , .
1 9 5 6 ; p r e c e d e n c e o f fu n c t i o n o r , Sy m b ios i s, II 3 99 4 00 , , .
I , 1 9 7,
2 1 1 ; c o r r e l a t i e c om p l e i t y v x Sy m m e t ry ( s ee M o rp h ol og y ) .
o f fu n c t i o n a n d , , 2 00, 2 1 1 ; p ro I S y p h i l i s , h e r e d i t a r y t ra n s m i s s i on, I ,
g re s s i v e c on c om i t a nt d i fl e re n t i a
t i o n, ,
I
2 01 4 ; p h y s i o l og i c a l u n i t s ,
-
I , 2 25 —6 , 3 62 , 3 6 4 , 3 6 9 — 7 1 , , 61 3 II C
T A T U AL P e rc e p t i v e ne s s , h e re d i t y
s oci a l a nd o rg a ni c i nt e r d e p e n d a nd t h e d i s t ri b u t i on o f, I , 6 02 —8 ,
e nc e , I ,
2 3 5 —4 2 ; a r i e d b y fu n c v 6 3 3 , 6 65 , 6 6 6 , 6 7 2 , 69 2 .
t i on , I , 3 34, 5 3 5 ; , 2 1 7 ( s ee A c II T ce m
’
a (s ee E nt oz oa ) .
q ui re d C h a ra c t e rs ) ; z o o l og i c a l T a ns l ey , A . G .
, I , II vi , , vi ; a da p t a
c l a s s i fi c a t i on I , , 3 90 2 ; 4
e quil i b ra t i on of r ep r od u c t i v e a c ti vi ty to
t i o n, I , 5 2 1 , 5 5 7 ; p r og re s s o f, a n d c on d i t i o ns in A l y ce , 1, 28 8 —9 ;
g e ne s i s , I , 5 9 0—1
; , 4 6 2 ; c oop e r a II sh a p es of C a u l er p a , II , 22 ; stem
t i on w i t h fu n c t i o n , II , 3; e vol a th ic k e n i ng in e x t i nct T ha llo

t i on a nd i nc r e a s e d , II ,
4; re t ro p h y te s , , II
5 6 ; na t u ra l s e l e c t i on
g ra d e m e t a m or p h os i s , II , 12; a n d l e a f- d i s t ri b u t i o n , , 1 79 II .
s i m u l a ted h om ol og i e s , II , 13 14; -
T a p e -w orm (s ee E n tozoa ) .
e a rl i e s t o rg a n i c fo r m s, II , 19; T a s t e, d e p e nd e n t on ch em i ca l ac
c y l i nd ri c a l v e g e ta l II 5 7 —6 2 ; , , per t i o n, I , 54 .
m a ne n c e a nd c o m p l e x i t y II , , 29 5 , T ee th : h e r e d i t a ry t r a ns m i s s i o n, I ,
29 6 ; fun ct i o n a nd e p i d erm ic, II , 3 06 ; s u p p r e s s i o n of m a m m a l i a n , I ,
3 12 - 4, 387 ; a nd m u s c u l a r, II , 3 69 , 457 ; of u nc i vi l i z ed a nd c i v i l i ze d , I ,
3 91 ; a d a p t a t i o n a nd e q u i l i b ra t i o n , 5 41 , 69 3 .
II ,3 9 2 ; p e rs i s t e n c e o f fo r c e a n d T e g um e nt a r y org a n s , o ri g i n o f, I ,
p h y s i o l o g i c a l a d a p t a t i o n, , 394 ; II
e v o l u t i o n II , , 5 01 4 -
.
(8 66 a lso T e l e g ony , or th e i n fl u e nc e of a p re
M orp h ol og y ) . v i ou s s i re on o ffs p r i ng , I , 6 2 4—7 ,
S t ru g g l e , fo r n u t r i m e nt a m o n g c om 64 4—6 , 6 4 9 —5 0 .
p o ne n t s of an o r g a ni s m ,
I , 562, T e m p e r a t u re ( s ee H e a t ) .
6 76 ; for e x i s t e nc e ( s ee N a t u ra l T e n s i o n ( a ce S t r a i n ) .
S e l e c t i on) . T er m i tes : fe r t i l i t y , I , 58 3 , II 4 9 3 ,
S t ru t h e r s , Sir J . : on h e re d i t y , I , l ate de v e l o p m e nt of s ex a l or u
3 05 , 3 1 4 ; d i g i t a l a r i a t i o n, , 3 21 ; v I g a ns , ,
I
68 0; nu t r i t i o n a n d d i flfe r
‘
ru d i m e n t a r y l i m b s of w h a l e , ,
I e nt i a t i o n o f fo r m s , , 681 I .
66 8 u ti l i z a t i on of a p h i d e s
‘
. T e tr a m om u m ,
S t r y c h n i ne , e ffe c t s o f, I , 5 4, 5 5 . by , I , 6 6 0—1 .
S t u r g e o n, s i ze of ova a nd a d ul t , I , T h a l l op h y ta : s i z e, I ,
1 38, 1 39 ; l ow
1 44 .
c o - o r d i na t i on of p a r ts , , 1 64 ; I
S u g a r s : p rop e r ti e s , 1 0 1 1 ; t r a ns , I -
p s e u d o-
fol i a r , II, 28 ; t ra ns i t i o n
fo rm a t i o n s , , I
3 8 , 4 0, 6 6 , 6 9 , 7 0, p l a ce, II , 3 0; s i m ul a t ion of
II , 593 .
h ig her II typ es , , 3 2 ; s e c o nd a ry
S u i c i d e , h e re d i t a ry t e n d e n c y t o, I , t h i c k e n i ng i n e x t i n c t s p eci es, II ,
307 . 5 6 ; s e x ua l a nd a e x a l s u g e ne s i s , II ,
Sul p hu r : a l l ot r op i c , I , 4, 5 9 ; o rg a ni c 84 . ( S ee a l so A l g ae .
)
e v o l u t i on I , , 7 03 . k
T i c l i n g , p h y s i o l og y o f, I , 76 .
S u n ( s ee L i g h t ) . T i d e ( s ee S e a ) .
vv
S u r i a l of t h e F i t t e s t, th e e x p re s Tim e, a s a fa c t or i n g ro w t h , II , 77 .
S U BJ E C T—I ND E X .
6 61
T i s s ue, H e rt w i g
’
s cl a ssi fi c a t i on I , , s e g re g a t i on a n d o r g a n i c r e p a i r , I ,
1 89 .
( S ee P h y s i o l o g y .
) 22 1 — 2 , 22 2 —6 ; c h e m i c a l , m o r p h o
T o ng u e , p e rc e p t i v e ne s s of tip, I , l og i c a l , a n d p h y s i o l og i c a l , , 22 5 —6 ; I
606 8 , -
66 5 , 6 7 2 —3 . II 6 12 ; s t a b i l i t y ,
, , 3 3 9 ; i ns t a b i l I
T o r t oi s e : c o nt ra s t e d l i fe of d og i t y a nd h e t e r og e n e i t y o f o rg a n i c ,
a nd , I , 1 03 — 4 ; na t u ra l s e l e c t i on I, 35 0; D a r w i n s g e m m u l e s ,
’
, I
I 5 34 356—
60 3 62 , 3 72 ; W ei s sm '
a nd c a ra p a c e , , .
, a nn s
T r a n s c e n d e nt a l P h y s i o l og y , g e rm pla sm
-
(q . v .
) ih ; . s o c i o l og i c a l
1 76 . c o m p a r i s on , I, 3 6 3 —
8 ; s p eci fic p ro
T re e , as sym b o l i z i n g p h y l og e n y , I , c l i vi t i e s in e m b r y o g e ny , I , 458 ;
42 8 , 4 5 2 — 3 .
( S ee P l a nt s ) .
p h aen o g a m i c , , 73 , 1 51 ; II a nn u l o s e ,
T r em a t od a : a g a m og e n e s i s , I , 277 ; II, 1 05 ; i n c i d e n t fo rc e a nd ho
p a ra s i t i s m , , 428 ; I a l t e r na t e g en m o l og o u s , II , 1 5 9 ; m o r p h ol og i c a l
e ra t i o n , , 5 92 I . s u m m a ry , II , 23 3 ( S ee a l s o P h y
.
T re m b l e y , A .
, t h e polyp ,
on , 223 I . s i o l og i c a l u ni t s .
)
T r i c h i n os i s , i n G erm a n y , , 4 2 8 I . U ni v e rs a l P os t u l a t c , I , 6 75 .
T ro c h op h or e , p h y l e t i c r e l a t i o n s h i p s U ns y m m et r i c a l , fi
d e n i t i o n, II , 1 31 .
s h ow n b y , ,
4 47 , , 1 I
0 8 —
9 II . U re a , m us cul a r e n e rg y a nd e x ore
T u b l col az: d eve l o p m e nt , II , 1 00; b i t i o n, I , 72 .
l a t e ra l s y m m et ry , II , 1 97 .
T uni c a ta : gem m a tio n I

, , 5 88 , II , 44 5 VA N B E N E D E N , P . J .
, on T aeni a , II,
a l t e r na t e g e n e ra t i o n, , 5 9 2 ; i nt e I 1 03 .
g r a t i o n, II , 9 3 —4
; t e r t i a r y a g g re Va r i a t i on : d i gita l,
33 1 ; e fi e c t s I ,
'
of
g a t i on , II , 1 24 ; s y m m e t ry , II , p a r e nt a l c o n d i t i o n s , , 3 24 ; o f I ai
1 94 5 -
. t e re d fu n c t i o n , I , 325, 3 3 4, 69 3 ;
T u nn y , size of o va a nd a d ul t , I , d i s s i m i l a ri t y of i ni t i a l c on d i t i on s ,
1 44 . I ,327 32, 3 3 3 ; -
s p o n t a ne ou s ,
”
I ,
T u r be l l a r i a : s e g m e n t a t i o n, II , 1 02 ; 3 2 8 , 5 1 3 , 69 7 , II 5 2 9 ; p e r s i s t e nc e
,
sym b i os i s , II , 4 00 . of fo r c e , I , 3 3 5 ; p h y s i ol o g i c a l
T u rni p : c h l or op h y l l in ro ot s , I ,
2 09 , u ni t s , I , 34 8 —5 4 , 3 6 0, 3 69, 3 7 1 —3 ,
II , 25 4 ; va s cu l a r s y s tem , II , 28 1 , II , 6 1 4—7 , 6 22 — 3 ; W eism a nn s
’
28 4 , 5 7 8 , 5 9 1 , 5 96 .
g erm p l a s m
-
t h e or y , I , 35 7 —8 ,
T w i ns : sim i l a ri t y o f, I , 32 4 ; t r a i t s 3 7 2 —3 , 6 7 1 , 6 7 7 ; II , 62 2 ; e q u i l i b ra
of w om en b e a r i ng , II , 45 7 . t i o n a nd eg eta l , v I , 5 2 3 —5 ; W ei s
T y p e s , p e r s i s t e nt ,
”
H u l ey x on, I , m a nn s p a n m i
’
xi a t h eory , I , 5 6 1 —3 ,
64 9 , 6 6 7 —9 , 6 7 1 , 6 8 5 ; r e p ro d u c t i e v
or g a n s , , I
5 7 0; n a t u ra l s e l e c t i o n
UL CE R, d erm a l s t ru c t u r e , II , 3 06 . a nd c o nc o m i t a n t, I , 6 1 4—2 1 , 65 3 ,
U l ti m ate R e a l i t y , i nc om p re h e n s i b i l 6 64 , 6 7 4 , 6 9 2 ; a nd d i s us ed o rg a ns ,
i ty o f, I , 1 20 . I , 64 8 , 6 68 ; p l us a nd m i nu s . I ,
U l va c el l m u l t i p l i c a t i on, II , 26 6 6 7, 6 85 ; M a s t e rs on c orr e l a t e d ,
out e r ti s s ue, II , 25 6 . in p l a nt s , II , 29 8 ,6 21 —2 ; e q u i l i
Um be l l i fer az: fl or a l s y m m et ry , II , b ra t i on of ,
f v a ou r a b l e , , 39 4 II .
1 71 ; a xi a l a nd fo l i a r o rg a n s , II ,
Va s c u l a r System : f e c t s of
ef veg eto
5 4 1 —6 . a l ka l i es I 5 5 ; n u t r i t i on ,
, ,
I , 1 46 ,
U ni t e d S t a t e s : c a s e s of t e l e g o ny , I , 1 48 ; e m b ry o n i c d e e l op m v e nt , I ,
b i rth ra t e , II , 5 20 . 1 69 ; s t ru c t u r a l t ra i t s , I , 1 92, 1 9 3
U ni t s : d i if e r e nt i a t i o n a nd dissim i fu n c t i on , 1 9 9 ; of A s ci d i a ns I , , , I
l a ri t y , I , 20 p rot y l e , I , 22 —3 2 02 ; fu n c t i o n a l d i ffe r e nt i a t i o n a n d
sh ap es in h igh er t y p es , I , 1 64 ; i nt e g ra t i o n , , 2 05 —6 ; o r g a n i c r e I
d i ffe r e nt i a l a ss im i l a t i o n, I , 1 8 0; p a i r, I , 2 1 7 , 22 1 —2 ; e ffe c t of fu n c
p r i m or d i a l org a ni c , I , 1 81 ; m or t i o n, I ,
2 2 9 , 2 34 —5 , 2 3 6 ; e u i l i b r a q
p h o l og i c a l com p os i t i o n, I , 1 8 4 —7 , t i on , I , 5 3 5 ; c om m u n i t y i n c o m
1 94 , 25 2 , II , 5 , 7 - 9 , 2 1 , 7 9 , 8 5 —6 ; p ou nd org a ni s m s , I , 5 88 ; d e v e l op
6 62 S UBJ E CT —I ND E X .
m e nt o f v eg e t a l II 27 3 5 , 2 7 9 — 84 , , ,
-
s o ry o rg a n s , II , 3 1 8 ; a i r c h a m b e rs ,
28 5 —8 , 3 8 8 ; d i ffe r e n t i a t i on o f, s u m II , 334 ; o s s e ou s d i ff e r e nt i a t i o n, ,
II
m a ry , II
, 2 8 8 —90 38 8 ; d i ffe r e n t i a ,
3 44- 5 5 ; a ct i vity a nd m u s c u l a r c o l
t i o n o f a ni m al, II , 339 —4 4 ; o s s e ou s o u r, II 3 65 —9 ; h
, e a rt - m ot o r appa
de v e l op m e nt , II , 3 4 7 - 5 1 ; m u s eu ra t u s , II 3 7 4 ; c o , st of g e ne s i s , II ,
l a ri t y , II
3 64 ; m ,
u s c u l a r c o l ou r , 4 3 6 ; a g a m o g e ne s i s ,
u n no w n , k II
II , 3 6 5 9 ; h e a r t m ot o r a p p a r a t u s ,
- -
44 5 ; g ro w t h a n d g e n e s i s , , 4 54 ; II
II , 3 7 4 ; d i ffe re n t i a t i on a n d i n t e x
h e a t e p e n d i t u r e a n d g e ne s i s , ,
II
g ra t i on in a ni m a l , II , 3 7 6 —9 , 3 8 3 ; 4 68 —9 , 4 7 4 ; O w e n , t h e o ry o f s e i e k
w ood fo r m a t i on , II , 5 6 7 —9 2 ; r é t on, , 5 48 II
—6 6 e o l u t i o n o f e rt e
; v v
s um é of w oo d fo r m a t i on, II , 5 9 2—7 . b rae , II , 5 63 —6 ; o ri g i n o f typ e, II ,
Va u ch er i a , re p r o d u c t i o n , I , 2 7 9 , 28 9 .
Ve g e t a t i v e S ys tem , c o - o r d i na t i o n o f Ves t i g es of C r ea ti on, I , 49 1 .
a c t i o ns i n, I , 5 78 . Vi b r i s s a e , fu nc t i o n of, I , 75 .
Ve g e t o -
al ka l i es , p h y s i ol og i c a l ef Vi t a l i s m ,
h y p ot h e s i s e x a m i ne d I , ,
fe c t s o f, I , 5 4—5 . 1 1 4- 7 .
Ve l o c i t y , of m o v i ng bod i e s , II , 21 9 Vi t t a d i n i , C .
, on s i l k w or m d i s ea s e ,
20 . I , 6 22—3 .
Ver te br a ta : s i z e,
1 39 ; si ze a t I ,
Vi v i p a ro us g e n e s i s, I , 27 1 , 2 74—5 ,
b i r t h a n d m a tu ri t y , , 1 44 : a i a l I x 2 78 .
s t ru c t ure, , 1 65 ; e m b r y on i c d eI Vo i c e , c or r e l a t e d se x ua l t ra i t s , I ,
v e l op m e n t a nd s e l f- m o b i l i t y , I , 3 7 1 —2 .
1 7 5 ; f u n c t i o na l d i ffe r e n t i a t i o n, , I Vo l c a n o , de fi n i t i on of l i fe a nd , I ,
206 , 5 9 1 ; r e p a r a t i e p o w e r , , 2 1 9 , v I 85 , 8 9 .
2 2 3 , 5 8 9 ; h o m og e ne s i s u n i e r s a l , v Vo l voc i ncw : u ni c e n t ra l de v e l op m e nt ,
I , 2 7 1 ; d i s t i n c t i e t ra i t s , , 392, v I I 1 63 ; i n d i i d u a l i t y ,
,
v I , 245 ; d i s i n ~
II , 3 5 ; d i s t ri b u t i o n i n t i m e , , I t e g ra t i o n o f g e n e s i s , I , 2 76 , 5 8 7 ;
4 08 ; c l a s s i fi c a t o r y a l ue , , 446 ; v I s p h e r i c a l a g g r e g a t i on , II , 24 ; s ym
e m b r y o n i c m a m m a l i a n r e s p i ra t o r y m e t ry , II , 1 3 7, 1 87 ; fe r t i l i t y , II ,
s y s te m , I , 456 ; emb ry o l o g i c a l p r e 44 1 .
a d a p t a t i o n, I , 4 6 1 ; e ol u t i o n a nd v Vom i t i ng , a l i m e nt a ry c a na l de vel
v e rt e b ra l c o l u m n, I , 4 7 0; ru d i m en op m e n t , II , 32 8 .
t a ry o rg a n s , I ,
4 73 ; e o l u t i on a n d v Vor ti ce l l a : s e c o n d a r y a g g r e g a t e , II ,
v a ri ed m ed ia , I , 4 7 9 —85
; s i z e of 90; s y m m e t ry , , 1 88 II .
h ea d v e rt e b r a! I 5 1 2 5 3 7
a nd , , ,
s e g reg a t i on a nd e v ol u t i o n of W AL L A C E , A . R T he O ri g i n of
v e rt e b rae I 5 1 5 ; fe rt i l i t y a nd
, , th e H um an Ra c es , I , 5 5 3 ; th e ex
d e v e l o p m e n t I 5 8 3 5 9 8 —9 ; W e i s v va l
“
, , , p re s s i o n Sur i of t h e F it
m a nn o n r e p o d u c t i v e I I
”
cel l s r , , t e s t, , 5 3 0; his a s s oc i a t i o n of
6 35 ; l i m b l oc om o t i o n, II , 1 5 ; a da p n a t u r a l w i t h a rt i fi c ia l s e l e c t i on, I ,
ti e v s e g m e n t a t i o n, II , 1 1 7 23 , -
609 ; c o -a d a p t a t i o n in g i ra ffe , I ,
1 2 5 —7 , 22 3 , 602 ,
605 7 ; s u p e r nu -
61 5 ; s kin s e ns i t i v e ne s s I , , 646 .
m e ra r y v e r t e b rae, , 1 23 ; b i l a t II W a sp : c o -o r d i na t i o n o f i n s t i nc t s i n
e ra l 203 - 6 ; i n t e r na l
s y m m e t ry , II , M a s on 1 , 5 7 4 , (37 9 —8 0; g e n e s i s of
o rg a n i c s y m m e t r y , , 2 08 ; g e n e II w or ker I , , 65 4—7 .
s i s o f r u d i m e n t a ry a i s , , 2 1 2 —6 ; x II Wa ste , a ni m a l , I ,6 9 , 2 1 3 - 5 , 228 ; r e
na t u r a l s e l e c ti on a nd g e ne s i s of l a t i on t o acti v I
i t y , , 1 9 6 , 22 0—1 ; i n
s t ru c t u re , II , 21 6, 22 7 ; o ri g i n of p l a nt s , I , 2 1 3 , 220 .
no t o c h o r d , II , 2 1 6 —8 ; s p i na l s eg W a ter : p ro p e r t i e s , I , 7 , 9 ; c o l l oi d a l
m e nt a t i o n , II , 2 1 8 —22 , 22 4 ; k ul l
s fi
a f n i t y fo r , , 28 ; I o r g a ni c c h a ng e
de v e l op m e nt II 2 22 , , , 2 27 ; r és um é fro m , I , 29 ; o rg a ni c ne e d fo r, I ,
of axis d e v e l op m e n t , II 22 4
, 1 47 ; p rop o r t i o n in m a m m al ia n
C op e o n a ut h or s
’
t h e o ry , II 2 2 5 —7 ;
, a d u l t a nd foe t u s , , 1 54 ; m I ot i o n
v
ne r e d i ffe re n t i a t i o n , II , 3 04 ; sen t h ro u g h , I ,
1 5 6 ; o rg a n i c d e v e l op
S U BJE CT—IND E X .
663
m e nt a nd en v i r o nm e n t , I , 1 73, d a t i on s of s oc i a l i n s e ct s , I , 654 ,
1 7 7 , 4 79 ; t e r r e s t r i a l o r g a n i s m s l n 65 8 —6 5 , 6 70, 6 7 5 , 6 7 8 — 84 , 68 5 ; foo d
h a b i t i ng , , I
4 00 ; a d a p t a t i o n o f o r k
s e e i ng i n s t i n c t i n A m a z o n a n t s ,
g a n i s m s t o c h a ng e of m e d i a , , I I , 66 0, 6 7 0; t h e c o -a d a p t a t i on of
4 79 —8 5 ; v
e g e t a l t i s s u e d i ffe re nt i a c o - op e r a t i ve p a rt s , I , 6 63 —4 , 6 7 0,
t i o n, , II
25 3 ; m ol ecu l a r rea r 6 7 4 , 6 75 , 6 7 6 ; t a c t u a l d i s c r i m i n a
r a ng e m e nt , II , 35 9 ; co l l oi d a l c on t i v e ne s s , I , 6 65 , 6 72 ; i nt r a - s e l e c
t ra c t i o n , II , 3 6 1 —2 . t i o n, 1 , 6 7 6 —8 ; e ffe c t of nu t ri t i o n
W a ter -
w eed , A m e r i c a n, in v a s i on o f, o n fe r t i l i t y o f b l o w - fl y , ,
6 78 —9 I .
I , 39 9 . W h a l e : w e i g h t o f b r a i n, , 5 99 ; I
Wa tts , D r .
, on T he P r i nci p l es o f Bi r u d i m e n t a ry l im bs, I , 6 68 - 9 , 68 5 ,
o l og y , I , ix . 69 3 .
Wax fo l i a r d e p os i t ,
, , 2 60—1 II . W h e a t a d a p t i v e va r i a t i o n s
,
’
,
II , 29 8 .
! Ve b e r, o n t a c t u a l d i s c ri m i n a t i ve W h i s t l i n g d e fi ni t i o n o f l i fe , a nd , I ,
ne s s , I , 602 . 1 12 .
W eight : r e l a t i on to env i ro n m e n t of White -

C o op e r , Mn , on i nh e r i t a n c e
or g a ni c , I , 1 74 , 1 7 7 ; v a ry i ng as of a b n or m a l v i s i on I , , 3 06 .
c ub e of dim e n s i ons , 1, 1 51 , II ,
W i l l ow , nu t r i t i o n a nd g ro w t h , I ,
4 34 , 4 7 0 .
Wei s m a nn , A ug . : r e p ro d u c t i ve ti s W i l s on , E . B . : c om p o s i t i o n o f e b ro
s ue l n M ed u s a , I
28 1 ; i n D a p h ni d az,
,
m a t i n, I , 260; s e p a ra t i o n o f s e g
I , 29 0; his t h eo r y o f t h e d i ff e r m e nt a t i o n s p h e re s of A m p h y owus
e nt i a t e d g erm p l a s m
-
and i t s fu n o vum I , , 69 1 .
d a m e nt a l u ni t s , , I 35 7 , 62 2 —3 , W i nd : a nd v eg et a l b i l a t e ra l sym
6 2 8 —3 0, 63 3 —44 , 64 6 , II , 6 1 8 —9 , 622 m et ry , II , 1 42 ; a nd i nn e r veg et a l

the d i fi e r e nt i a t i o n
a l l e g ed
'
a nd ti s sue d i ffe r e n t i a t i o n , II , 2 7 5 —9 ,
p l a n t -
p h e n o m e n a , , 3 5 9 —6 0
; I a nd 28 5 , 28 8 , 3 8 8 ; a nd p r o l i fe ra t i o n o f
r eg e ne r a t i ve p r oc e s s e s , , 3 6 0; I B r y op h y l l u m , II , 2 9 5 ; a nd eg et a l v
fa l s e j oi n t s I , , 3 6 2 ; i m p l i e d c om sap m o v e m e nt II , , 5 83, 5 84 , 587
p l exi ty of d e t e rm i na n t s , I , 3 70 r es u m e, 5 9 2 —6 .
t h eo ry i na d e qua te to e xp l a i n c or W o l df , C . : veg eta l fr u c t i fi c a t i on a nd
re l a t i on of I 3 72 ; s e xu a l t ra i t s, ,
n u t ri t i on, I , 28 3 , II , 1 7 9 —8 0; v eg e
a n d v a i t i o n s i n p e a c oc k s t i l
r a
’
a ta l v a s cul a r s ystem ,
II , 28 3 .
fe t h ea I 3 7 2—3 69 5 II 6 1 8 ; h i s
r, , , , ,
W om e n ( s ee M a n) .
v i e w o f n a t a l l c t i n a s s ol e ur se e o W ood ( s ee P l a nt s ) .
fa c t o i n or g n i c e v o l t i o n I 5 5 9 ;
r a u , ,
t h e d o c t i ne o f p a n m i x i
r I 5 61 3 a, ,
—
, YE A ST : fe r m e n t a t i o n, I , 38 ; fe rt i l
6 1 2, 63 2 , 6 49 , 6 6 7 —9 , 671 , 68 5 , i ty , I , 581 , II ,
44 0; l i ne a r a g g re
68 9 ; a rg u m e nt s a g a i ns t i nh e r i t g a t i o n, I , 5 87, II , 23 .
a nc e of ac q u i re d c h a r a c t e rs , 1,
6 12 —3 , 65 1 —65 , 66 9 —7 1 ; b l i n d n e s s o f ZEBRA m a ks i n h o es I r rs , , 31 4 .
v
c a e - a ni m a l s , , 6 1 3 ; c u r r e nt a c I Z l g y c l a s s i fi c a t i on 1
oo o , , , 1 24 5 , -
c e p t a nc e of h is vi ew s , I , 63 1 , 38 0—9 .
690; c a nn o t e xp l a i n the p r oc e s s Z oop h y t e s s t c t ur a l i nd e fi ni t e , ru
of na t u r a l s e l e c t i o n, I , 65 1 ; t h e n ess, I 1 73 , .
d e g ra d a t i on of th e l i ttl e t oe in Z oos p ore s u n i t l i fe of I 1 85 ,

-
, , .
m a n, I , 6 5 2 , 66 9 , 6 73 ; ca st e g ra Z y g ot e of c onj g a t i ng A l yce I 283

, u , , .
THE E ND .

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T HE P R IN C I P L E S OF

RE VIS E D A ND E NL A RGE D E DITIO N

T o t he s tate m ent s m a d e i n the p r e f a c e t o th e ﬁ r s t v ol u m e

O ne of t h e m i s th at the re v isi on has n ot b e e n c a rr i e d

m ark e d as n ew, th e e ss enti a l id ea s se t f or th a re th e sam e as

c our s e s om e wh a t li k e t h at s u g g est e d by P r of Mor g an .

Th e a ddi ti on s m a d e t o t his v ol u m e a re l e ss nu m er ou s and

g ani e W or ld , s er v e s t o r oun d off th e g ene ra l t h eory of

E v ol uti on i n i ts a p p li cat i on t o li vi ng t hi ngs B e y on d a n e w .

X I I i n P a rt V I .Mor eo v e r t h ere a r e t h re e f u rth e r a pp en

say t h a t the ai d n e e d e d for b ri ngi ng up to d a te th e c onte nts

o f th is v ol u m e f a ll P l an t Morph ol o gy and P l ant P hysi

l gy h a v e b een o v e r s een by Mr A G T an sl e y Oriti

cism e u p on p art s d e ali n g wi th A nim al Mor ph ol og y I owe

A n d the s ta te m ents incl u d e d u nd e r A n im al P hysi ol ogy

r e sp on sibl e for s om e e rr or which i s n ot hi s b u t mi n e I t is .

d one t his i n re sp e c t of m a tte r s of i n f e ren c e , but in s u ndr y

P e rh a p s I m ay be e xc u s e d for e xp re ssi ng s om e sa t i sfa c

t i on t h a t I h a ve n ot b een obli g e d t o re li n q u i s h th e v i e ws set

as I am a w a re , the a ll e g e d g en e ra l l a w of or a nic symm e try

not I b e li e v e b een c ontr o v ert e d

w ork s on b ota n y h a v e gi v en c u rre n c y to th e d o c tri n e s et

p ro v e i t The only v i e ws c ont e ste d by c ert ai n of th e g e n

T HE p r oof sh eets of this v ol u m e , l ik e th os e of th e l a s t

P rof H u xl e y ; a n d I h a v e , a s b e for e , to t h a n k th e m for

t h e i r v a l u a bl e c ri ticisms , a n d for th e t rou b l e t h e y h a v e

P a rt I V , wit h which this v olu m e c om m e n c e s , c onta i n s

ti on s , m ostly a l t e re d i n sc al e a nd s impliﬁ e d i n e x e cu ti on, of

o us B otan is t s a n d Z ool ogis t s A m ong th e a uth ors wh om I

h a v e l aid u n d er c ontri bu ti on, I m ay n a m e B er k e l e y , C ar

E d w a rds , R a lfs , S mi th T he r e m a i ni ng ﬁ gu re s , n u mb eri ng

T h e s u cc e ssi v e i n st a lm e nts which c o mp os e this v ol u m e,

V — T HE M O R P H O L O G I CA L C O M PO S ITIO N O F A N I MA LS— Conti nu ed

X III — M O R P H O L OG I CA L DI FF E R EN TIA TIO N I N A N I MA LS

VI —D I FF E R EN T I A TIO NS BE T WEEN T H E O U T E R A N D I NNE R TI SSUES

X — T H E I N T E GRA TIO N O F T H E O RGA N I C W O R L D

V II I — AN T AG O N I S M BE T WEEN E X P EN DIT U R E A N D G ENES I S

I X — C OI N C ID EN C E BE TWEEN H I GH NU T R ITIO N A N D G ENES I S

X — S P E C I A L ITI ES O F T H ESE R EL A TIO NS

t/ XII I — HU MA N POP UL A TIO N I N T H E F U T U R E

G —T H E I N H E R IT A N C E O F F UN C TIO N A LLY CA USE D M ODI FI CA TIO NS

§ 1 75 . T H E di visi on of Mor phol ogy f rom P hysi ol ogy i s

one whi c h m ay be t ol er a bly w e ll p r e s er v e d s o l on g as w e do

not c a rr y ou r i n qu i r i e s b ey on d th e e mpi r ic a l g enera li z a ti on s

of th e i r res p ectiv e ph enom e na ; bu t it is one w hich b ec om e s

i n grea t m ea su r e nom i na l wh en th e ph enom en a are to be

ra ti on a lly i n t er p ret e d I t wou ld be p ossibl e a ft er a na lyzi n g

ou r S ol ar S y s t e m to set d own c ert a i n g enera l t ru th s re sp ect

in g th e sizes a n d di st a nc es of its p ri m ary a nd s ec ond ar y

be p oss ibl e to set d own c ert a i n oth er g ener a l t ru th s resp e ct

p os iti ons fu rth er th an as gr eat er or l ess ne arer or m ore r e

m ot e Bu t on seeki ng to a ccou nt for th es e genera l t ru th s

a rr iv e d a t by i n d u cti on w e ﬁ nd ou r se lv e s oblige d t o c on sid er

s im u lt a neou sly th e r e l a tiv e s i z e s a n d pl aces of th e m a ss e s ,

a nd th e re l a t iv e a m ou nts a nd di recti on s of th e i r m oti on s .

S imil arly with orga n i s m s T h ou gh w e m ay f ra m e s u n d ry

We m a y est a bli s h s ev era l w id e c on cl u s i on s r esp ecti ng th e s ep a

a nyt hi n g d eﬁ n it e res p ecti n g th e f orm s a nd p os iti on s of th es e

th e hyp oth esi s of E

s tru ctu r es a nd f u n c ti on s i n th ei r m u tu a l r el a ti ons E v ery .

w h er e st ru ctu res i n g r ea t m ea su re d et erm i n e fu n cti ons ; a nd

I n N atu r e the two ar e i ns ep a r a bl e co—Op er a tor s an d S ci enc e

the i r co—op er a ti on c ons t a ntly i n Vi ew A n a ccou nt of org anic