G E N ER A L B I O L O GY

BY

L EO NA S LAN C ELOT B U R LINGAM E

AS S O CI AT E P R O F E S S O R O F B OT AN Y A N D G E N E T I CS

HA R OLD HEATH
P R O F E S S O R O F Z OOLO G Y

ER N ES T GALE MA RTI N
PR O FE S S O R OF PH Y S I O LO GY
AN D

G EOR G E ! AM ES PEI R CE
P R O F E S S O R O F P LA N T P H Y S I O L O G Y

AT S T AN F O R D U N IVE R S I TY

W4 1 8

IO .
q .
9 8

NEW YOR K
H EN R Y HOLT AND C OMPAN Y
 9
3 8

Common, 1 922,
BY

HE NRY HOLT AN D COM P A N Y

Decemb er . 1 922
 mut a tion
F A
TH I S B O OK I S RE S P E CT U LLY D ED I C TED

PR ESI D ENT R AY LYM AN WI LBUR

WHOS E SYMPATHE TI C E N C O U RA G EME NT AN D HE AR TY CO O PE R ATIO N

MAD E I T P O S S I B LE T O ATTE MPT

E S TA B LI S HME N T O F S U CH A C OU R S E

AN D LED E VE N T UALLY T O THE

ATI O N
P U B LI C O F TH I S TE X T
 P R E FA CE

T H I S tex t is th e outcome of a c ou rse of l ectu res gi v en

to L owe r D ivisi on stu den ts a t S t a nford U nive rsity i n
s a tisfa cti on of one of thei r requi reme nts The c ou rse.

h a s bee n c on ducted by a c ommittee with the a ssist a nce

of a n umbe r of th ei r c olle a gues a c knowledgme nt o f wh ich
,

is m a de bel o w .

Th e ide a u nde rlyi ng this b ook wh ich wa s a d op ted by

this c ommittee h a s bee n set f orth a t so me le ngth by
Pro fe ssors Bu rli ng a me a nd M a rti n i n S ci ence for M a y
7 1 920 Th e ide a is subst a nti a lly th is Bi ol o gy h a s now
,
. .

re a ch ed such devel opme nt th a t it c omp rises a l a rge

a m ou n t of i nform a ti on a nd k nowledge which is no t
on ly Of th e fi rst p ra ctic a l imp ort a nce to th e l a ym a n
but is a ls o suscep tible of bei ng org a nized i nt o a
c ohe re nt set of p ri nc iples s uit a ble for p rese nt a ti on to
the ordi na ry f resh m a n or to th e l a y p ublic . Th is
necess a rily i nv olv es s t ressi ng p ri nciples ra th e r th a n
fa ctu a l det a ils a nd limits discu ssi onto s uch t opics a s do
not put a n u ndue s t ra in on th e re a de r s k nowledge of
’

rel a ted scie n ces such a s physic s ch emist ry a nd ge ol ogy

, ,
.

An i nspecti on of the t a ble of c onte nts Will S h ow th a t

the th re a d wh ich ru ns th rough th e wh ole is th e inte r
depe nde nce of org a nisms especi a lly i n r el a ti on to ma n
,
.

This it seems to u s na tu ra lly i nv olves a discussi ono f th e

f oll owi ng t opics a t le a st :
I Th e na tu re of th e living subst a nce p rot opl a sm a nd
.
, ,

its divi si on a nd diffe re nti a ti on i nt o th e speci a li zed cells

of c omplex org a n isms
.

I I Th e rOl e of green pl a nts i n th e synth esis of th e

.

p rim a ry f ood a nd fuel c omp ou nds a s a s ou rce of m a te ri a l

an d ene rgy for a ll l iving thi ngs .

VII
v iii P RE FA CE
I I I The m a inte na nce o f li fe of a ll s orts by the utiliz a
.

ti ono f th e m a te ri al s a nd e ne rgy of foods This i nv olves .

o f c o u rse s ome c o n side ra ti ono f the m a chi ne ry by wh ich

me ta b olism is c a rried o n .

I V The mech a nisms by which pl a nts a nd a nim a ls

.

a dj us t thei r v a ri o us o rg a ns to one a no th e r a nd themselves

to thei r e n vi ronme nt b o th physic a l a nd bi ol ogic a l i n ,

v olvi ng a c onside ra ti onof th e ge ne ra l st ructu re a nd fu nc

ti onof ne rv o us systems a nd o f h orm one sec ret i on .

V The i nte ra cti ono f org a nisms wi th one a no the r i n

.
,

v olving such t opics a s f a m ily a nd he rd rel a ti onsh ips

'
,

hum a ns ociety symbi osis p a ra sitism a nd dise a se

, , ,
.

V I D e a th o f o rg a nisms a nd o f p ro t opl a sm
. .

V I I The dec omp o siti on o f org a nic rem a i ns a nd th e

. .

c onseque nt e nrichme nt o f th e s o il inv olvi ng a discussi on ,

o f th e bi ol ogic a l a n d chemic a l p ro cesses i nv olved a nd

thei r rel a tionto the cycles of e ne rgy a nd m a te ri a l .

V I I I The g rowth a nd rep rod ucti on o f org a nisms

. .

IX Th e mech a nisms a nd l a ws of h e redity a nd thei r

.
.

a pplic a ti o n to the b reedi n g of d omestic a ted pl a nts a nd

an im a ls a nd to the p rog ress of the h um a nra ce .

X Th e f a c ts the p ri nciples a nd the results o f ev ol u

.
, ,

ti o n .

X I The fa c ts a nd p ri nciples which a re i nv olved i n

.

th e dist ribu ti ono f org a nisms i ntime a nd sp a ce .

X I I M a ns pl a ce i nna tu re i nv olvi ng onth e one h a nd

.
’
,

a disc ussi o n o f th e f a cts which m a ke ma n a n a n im a l ,

an d onth e o the r h a nd a se tti ng f orth o f th o se eve nm ore

impo rta nt fa c t s which le a d us to believe th a t ma na l o ne
o f a ll a n im a te th i ngs is m a s te r o f th e desti ny o f h is ra ce .

A c ou rse o f th is s ort necess a rily i nv olves a c a refully

org a n ized l a b ora t ory c ou rse illust ra tive o f th e p ri nciples .

devel oped i nth e lec t u res A t S ta nf o rd th e stude nt ma y

.

elec t to ta ke such a c ou rse m a i nta ined for th e pu rp ose or

, ,

to subs titute for it a nequiv a le n t a m ou nt o f eleme nt a ry

bo ta n y or z ool ogy .

Al th o ugh th e a u th ors of this tex t h a ve b orne th e chief

 P R E FA CE ix

bu rde nof these lectu res th ey a re deeply indebted to th ei r

c olle a gues b o th for a ble a nd willing a ssist a nce i n the
.

p rese nt a ti on of the lectu res a nd for h elpful suggesti ons

i nthe p rep a ra ti onof the text Wh e re so m a n . y h a ve bee n
of g re a t a ssist a nce it is p ra ctic a lly imp o ssible to m a ke
’

a ckn owledgme n t of e a ch sep a ra te f a v or Th e ge ne ra l .

na tu re h oweve r of ou r oblig a ti ons will be i ndic a ted by

, ,

th e f oll owi ng P resident Wilbu r h a s regul a rly given

.

th ree lectu res ondise a se i na nim a ls a nd one onthe rel a

ti onof h um a np rog ress to scie ntific disc ove ry P rofess or .

D oa ne h a s give n ge ne rously of his time a nd a dvice i n

c onnecti on with th e le ctu re on i nsects a nd dise a se a nd ,

Profess or M cM u rp h y h a s bee n simil a rly h elp ful with

pl a nt dise a ses D r Alsbe rg of th e F oo d R ese a rch I nsti
. .

tu te h a s bee n h elp ful i n ou r discussi on of th e st ructu re

an d fu ncti ons of p rot opl a sm P rofess or W a lte r Fishe r
.
,

of th e H opki n s M a rine S t a ti on h a s regul a rly p rese nted

,

the subj ect of dist ributi on o f l a nd a nim a ls a s well a s ,

h a vi ng bee nof se rvice i nth e gene ra l org a niz a ti onof the

c ou rse P ro fess or Ab ra ms h a s regul a rly give nth e lectu re
.

on ge og ra phic dist ributi on o f pl a n ts a nd h a s likewise

bee nof se rvice i n th e discussi onof m a ny o the r m a tte rs .

The c ont ribu ti ons o f D r V est a l di rect or of the l a b ora

.
,

t ory h a ve bee nso nume rou s a nd v a ri ed a s to defy speci

,

fi ca ti on bu t thei r v a lue is evide nced by th e freque nt use

,

m a de of th em Th ey ra nge a ll th e wa y f rom th e de
.

live ry of lectu res on e col ogic a l dist ributi on of org a nisms

to th e selecti on o f suit a ble illust ra ti on s D r Becking . .
,

wh o will he ncef orth be a ss oci a ted wi th D r V est a l i n the .

di recti onof th e l a b ora t ories h a s a ls o a l re a dy m a de m a ny

,

stimul a ti ng suggesti ons m a ny o f which h a ve bee nof i m

,

medi a te use i n w riti ng this tex t P ro fess or J P S mith . . .

h a s a ssisted i nv a ri ous w a ys i nc on necti onwi th ge ol ogic a l

dist ributi on of org a nisms We a re a ls o indebted to
.

v a ri ous a u th ors a nd publishe rs for pe rmissi on to rep ro

duce figu res from th ei r b ooks or for origina l ph o t og ra phs .

S pecific a c kn owledgme n t is m a de of e a ch such f a v or i n

 P R E FA CE

c onnecti on with th e figu res i n a dd iti on to this ge ne ra l

on e . P rofessor Te rm a n h a s not only delive red the l ee
tu res onth e rel a ti ono f he redity a nd e nvi ronme nt inma n
b u t h a s a ls o w ritte n ch a pte r 4 2 in e nti rety I t is to be .

u nde rst ood th a t the a u th ors a ss ume full resp onsibili ty

for the st a teme n ts a nd views exp ressed i nth e tex t .

L L B
. . .
,
H H
. .
,
E G
. . M .
,
G !
. . P .

ST AN F O RD U NIVE RS IT Y, C AL I FO RN I A .

D ece mbe r, 1 922 .

 C ON TEN T S

B i ol o gy D e fi n ed

Ai m of Genera l B i ol o gy
T h e L ivi ng S ubs t a nce , P ro t o pl a sm
Green P l a nt s R ol e
’

T h e M a intena nce o f Lif e i n A nim a ls a nd N o n -

Gree n P l a nt
C ells
A dj us t m ent of Or g a nisms t o T h ei r Envi ronm ent
T h e I nt er a ct i o no f Or g a nis ms
D ea t h
T h e D e co mp o si t i o no f O r g a ni c R e m a i ns
T h e R e p r o d u ct i o nof P l a nt s a nd A nim a ls
Here di t y .

E v o lu t i o n
T h e D is tribu t i ono f A nim a ls a nd P l a nt s
M a ns P l a ce i n N a t
’
ur e

PAR T I
SE CT I O N 1 . TH E L I V IN G S U B S TA N CE

A
C H P T ER II
‘

P R O T O P L AS M AN D T HE CE L L
U niv ers a l i t y o f P ro to pl a sm
P ro t o pl a sm A lw a ys O r g a niz e d i nt o C el ls
T h e D is co v er y o f C ells
T h e D is co v er y o f P r o t o pl a sm .

P hysi ca l C h a r a ct er is t i cs o f P ro t o pl a sm .

T h e C h e mis tr y o f P r o t o pl a sm
T h e P hysi o l o gi ca l C h a ra ct er is t i c s o f P r o t o pl a sm
Gro w t h , W a s t e , a nd R e p a i r .

T h e P o w er t o L ib e r a t e E ner gy f r o m Fo o d
T h e P o w er o f C a usi ng C h e mi ca l C h a ng e
S e nsi t iv e ne ss t o S t imuli
S t imul a t i o n, C o ndu ct i on, a nd R e sp o ns e
C ell D ivisi o na nd R e p r o du ct i o n .

T h e S tr u ct ur e of a T ypi c a l C ell
S u mm a r y .
xii CO NT E NT S

C HA P T E R I II

MO I/TI CE LL U LAR O R G AN I S M S AN D D I V IS IO N OF LAB O R

N a t ure of th e S u bj ect
Pa ra me ci u m
I mp o r t a nce o f O ne - C e ll e d O r g a nisms
Co l o ni a l For ms .

V o lv o x
Hyd ra .

D ivisi o no f La b or
Tissu es , Org a ns a nd S ys te ms

S E CTI O N 2 . TH E R OLE OF G R E E N PL A N T S
CH A P T ER Iv

T HE M A N U FACT U R E OF FO O D S AN D FU E L S

The A nnu a l V a l u e of Foo d

Annual P ro du cti on of Fo o d N e cess a r y
T he Fo o d M a t e r i a ls
W a ter a nd M i nera l S a l ts
Th e C o mp o si t i o no f Foo d M a t eri a ls
T h e A bs o r p t i o no f Fo o d M a te ri a ls
Th e W o rki ng O v e r O f Fo o d M a t er i a ls
Ra di a n t En er gy .

P h o to sy nt h es is .

C hl o ro phyll
O c c u rre nce of
Wh a t Limi ts P h o to sy nt h e s i s
T h e P h o t o c h e mi ca l R e a c t i o ns I nP t O
.

sy t
Vo nB a y er s Hyp o t h e sis o f P h o t o sy nt h e sis
’
h es i s n .

T h e Pro d u c ts o f P h o to sy n t h e sis .

C HA P T ER V
T HE B U I LD I N G O F T HE LI V I N G M AC H I N E
T h e Livi ng M a chi n e

T h e C o mp o si ti o no f P ro to p l a s m

P hysi ca l Pro pe rti es o f P ro te i n s

T h e A mi n o- A cids

T h e S y nt h e sis o f Pro te i n s f ro m A mi no- A cids

T h e O rga ni z a ti o no f P ro t o pl a sm ”
T h e Fu n cti o ni ng o f P ro to pl a s m
T h e Gro wt h o f P ro to pl a sm
T h e D ea th o f P ro to pl a sm .

T he Mi n e ra l C o ns ti t u e nts .

P o ta s si u m
Iro n
 C O NT E NT S

C HA P T ER VI

T HE I N CO ME , M O VE ME N T S ,
AN D OU T G O OF M AT E R I A L S IN

P L AN T S

C ells a re B a gs o f
P er m e a bili t y .

O sm o ti c P re ss ur e
Livi ng C el ls a r e O sm o t i c S ys te ms
T h e S tr u ct u re O f R o o ts
M ech a ni ca l R esul ts o f R o o t S tr u ct u re ‘
.

M o v e m ent of W a ter a nd D iss o lv e d S ubs ta nces

T h e S tr u ctu r e o f S t e ms
T h e A s ce nt o f S a p .

T h e S tr u ct ur e o f Le a v e s
Le a v e s a s A bs or bi ng O r g a ns .

Lea v e s a s O r g a ns o f Ga s E x ch a nge
E v a p o ra t i o n
R o o ts a s E x cre t i ng O r g a ns
S e cr e ti o na nd E x cre t i o n
G u tt a ti o na nd B l e e di ng .

C H A P T ER VII

T HE S T O R A GE AN D DI GE S T I O N OF FO O D S IN PLAN TS

S tora ge
T h e C o mm o n P l a ce s o f
T h e P l a ce s o f M or e P e r m a ne nt S t o r a g e
S t a rc h
O th er S t o re d Fo o ds
M e a ns of M o vi ng M a nuf a c t u re d Foo ds
T h e P a t hs o f M o v e m e nt
D ig es t i o n .

T h e M ea ns of D ig e s t i o n
C a t a lyz er s
E nzyms .

C l a ssifi c a t i o no f Enzyms
Ger mi na t i o na nd S p r i ng Gro w t h

SE C TIO N 3 . TH E M A I N T E N A N CE OF L I FE
C HAP T E R V I II
T H E U TI L I Z A T I O N OF FO O D
R e a s o ns Why Livi ng Thi ngs R e qui re Fo o d I
M et a b o lism i n Genera l
B a si c M e t a b o lism .

Gro w t h M e t a b o lism
T h e M a nu f a ct u r e o f M a t er i a l
T h e M e t a b o lism o f A ct iv e Fu nc t i on
N erv o us A ct ivi t y
xi v CO N TE NT S

Me ta b o lis m a s a S o u rce o f E nergy .

K inds of Foo d U s ed i n M e ta b o lism .

M e ta bo lis m i n A nim a ls is O f te n W a s te f ul
W a r m bl o o d e d A nim a ls
-
.

C o ld bl oo d e d A nim a ls
-
.

W a s te M a te ri a ls a re Pro d u ce d i nC o nne cti o n Wi t h M e ta b olism

W a s te i n G ro w th Met a b o lism .

Wa s te inFunctiona l Meta bo lism

W a s te M us t b e Go t te n R i d o f if M e ta b ol ism is to C o ntinu e .

M e ta b olism Go e s o ni nE v ery C ell

T h e Servi ces of S u pply

C H A P T ER IX

T R AN S P O R T AT I O N S Y S T E M I N A N I M AL S
T HE

C o ndi t i o ns Fa v or i ng C e ll A c tivi t y
N e e d O f T ra nsp o r t a t i o n
C e ll D i fl u s i o n
’

C el l S u rf a ce a nd B o dy S u rf a ce
W a ter Fronta ge i n Low er A nim a ls
T h e Fl a t w or m T ra n sp o rt a ti o nS ys te m
M o d era te ly C o mpl e x T ra nsp ort a t i onS ys t e ms
C i rcu l a t ory S ys te m o f High er A nim a ls
T h e Bl oo d
Lymph .

Lymph a t i c o r A bs or b e n t V e ss e ls
S umm a ry

C HA PT ER X
T H E FO O D - P R E P A R I N G S Y S T E M

D ig es ti o n
C la sse s o f Fo o ds
Fo o ds P ro p er .

Alim e nta ry T ra c t
M a nuf a ct ure of E nzyms
S a liv a
G a s tri c J ui ce
Pa nc re a ti c J ui ce
T h e C o mp o si ti o no f P ro te i ns
T h e I nte s ti n a l J ui ce

M o v em e nts o f t h e Alim e nt a ry T ra c t
A bs or p t i o n

C H A P T ER XI

R E S P I R AT IO N AN D T HE D I S PO S AL or W AS TE S
R es pi ra ti o n
D i ff usi o no f G s es .

R e spi ra to ry O rg a ns .
 CO N T E NT S

R e spi ra tory O rg a n s of La nd A nim a l s

T h e T ra ch ea l S ys t e m of I n s e ct s
Lu ngs .

R espi ra tor y P igm en ts

E x cre t i o n
E x cre t or y O r g a ns o f Lo w er A nim a ls
K idne ys of H igh e r A nim a ls
K idneys , M o d e of O p era ti on

SE CT I O N 4 . A D ! US TM E N T OF OR GAN I S MS

C H A P T ER X II

Ho w P L A N T S AR E A D ! U S TE D TO T HE I R EN VI R O N ME N T

T h e Envi ron m ent

T h e N e ce ssi t y f o r A dj us t m ent
P l a nts a nd A nim a ls C o ntra s t e d
T h e A dj us t m e nt o f R oo t s t o th e S o il
Geo tro pism
C h e m o tro pism
T h er m o tro pism
O t h e r T ro pisms o f R O o ts

A dj ust m ent o f S te ms .

T endr il s a nd T wi ni ng S te ms
A dj us t m ent of Le a v e s .

A dj us t m ent of R e p ro du ct i on
In t erna l C orr e l a t i o ns i n Pl a nts
T h e M e c h a nism o f T r o pisms
T h e M e ch a n i sm o f C orr el a t i o ni n P l a nt s A .

C H A P T ER XIII
M O VE ME N T IN A N I M AL S
M o v e m ent i nP ro to z oa
Ammb o i d M o t i o n I

C i li a r y M o t i o n
Fl a ge ll a t e M o t i o n”
C o ntra ct i o no f P r o t o m
pla s i c S tra nds
M oti o nI nM a ny C e ll e d A nim a ls
-

M us cl e s
T h e B i o l o gi ca l I mp o r t a nce o f M o t i o n
Th e R e gul a t i o nof M o t i o n
Typ e s of Efi ect ors
S k e l e t a l M u s cl e s
S m o o t h M us cl e s
H ea rt M us cl e .

M us cl e Fib ers
M a k e u p of S k el et a l M us cl e
-

M a k e u p of S m o o t h M us cl e a nd Hea rt M u s cl e
-
.

T h e S tr u ct ure o f S k e l eta l M u s cl e Fib er

xvi CON TE NT S

S m oo t h M u s cl e Fib er
He a rt M u s cl e Fibe r
A t ta chm e nt o f M us cl es
J o i nt M o t i o ns
M us cle Fo rm i nR e l a t i o nt o M u s cl e D u t y
-

S t re n g t h o f M us cl es .

Lo ng M us cl e s
C la s s e s of M o v e m ents

C H A P T ER X IV

A N I M AL S E N S E O R G AN S

A dj us t me nts i nLo w er A nim a ls

R e ce p to rs o r S e ns e O r g a n s
C l a ss e s o f S e n s e O r g a ns .

In te rn a l S en s e O rga ns
H u ng er
Thi rs t .

P a in .

M us cl e S e ns e
E quilib rium .

T h e C o nta ct S e ns e s
T o u ch
T h e T e mp e ra t u re Se ns e
T h e C h e mi ca l S e n s es .

T a s te
T h e D is t a n ce R ece p t o rs

S m e ll
T h e O r g a no f Hea ri ng is t h e E a r
S igh t
f
T h e Perce p t i o no f Fo r m
I m a ge Fo rm a t i o n .

A cce ss ory E ye S t r u c t u re s
T h e P e rce p t i o no f C o l o r
T h e Pe rce p t i o no f D is t a nce
T h e R el a ti o no f S igh t t o T o u c h

C H A P T ER XV
N E R V O U S C O R R E LAT I O N

O ri gi no f t h e N er v o u s S ys te m
C o rre l a ti o ni nP ro to z o a
C o rrel a t i o ni nP ri mi tiv e M e ta z o a
C o rrel a ti o ni nHigh e r Fo r ms .

T h e S tru ct u re o f t h e N e rv o u s S ys te m
S impl e R e fl e x A c ti o n
N e rv o s O rg a niz a ti o ni nJ o i nte d A n
u im a ls
C o mpl e x R e fl e x A c t i o n .

I m po rta nce o f t h e Fro n t En d o f t h e B o dy

T h e A dv a nt a ge o f t h e D is t a nce R e ce p to rs .

Ma ch i nery fo r U si ng P a s t E xperi ence

 CO NT E NT S x vi i

C onta ct R e ce p tors P a rti cip a te i n M e m or y

Asso ci a ti on
T h e C o ntr o l of A ctivi ty i nth e H igh er A nim a ls .

B e h a vi o u r D e t er mi ne d by N er v o u s O r g a niz a t i o n .

Ha bi t For m a t i o n
C ons ci o us ne ss o r A w a re ne ss
T h e Fee li ngs o r E m o t i on s .

B o dily C h a nge s i nE m o t i o n
T h e R e p ro d u ct iv e I ns ti nct

C H A P T ER X VI

C H E M I C A L C O R R E L AT I O N
T h e N e ce ssi t y f or Gra du a l A dj us t m ent s
C h e mi ca l A dj u s ter s ”
B a si c a nd Gr o w t h M e t a b o lism C hi e ly A f f fecte d
C h e mi ca l C o rr e l a t i o nI n P l a nt s .

C h e mi ca l C o rre l a t i o ni n A nim a ls
Ho r m o ne- M a nuf a ct u ri ng Gl a nds
H or m o ne s Whi c h A ff e ct B a si c M e t a b olism .

T h e T hy ro id H o r m o ne
wM
g Gr o th eta b ol ism
.

T h e H o r m on es A ff e c t i n

T h e H or m on e s o f t h e P i t ui t a r y B o dy .

T h e I nter s ti t i a l H o r m o ne
T h e H o r m o ne s A ffec ti ng Fu nct i o na l M e t a b o lism
T h e P a ncr e a t i c H or m o ne
H or m o ne s A ff e ct i ng b o t h Basi c a nd Fu nct i o na l M e t a b o lism
.

T h e H o r m on e of t h e A d r e n a ls .

The R e gul a t i o no f A d r e na li nS e cr e t i o n .

The Eff e ct o f A d rena li no n M us c ul a r P o w er .

The Eff e ct o f A d r e na li no n B a si c M e t a b o lism .

Th e R e l a t i o no f H or m o n e s t o R a ci a l C h a r a ct e ri s t i cs .

S u mm a r y

C H A PT E R X V II
D I S E ASE S D U E To M AL A D ! U S TME N T

D e fi ni t i o no f D is ea s e
M a l a dj us t m e nt s
N a t u re o f D is e a s e
S ymp t o ms o f D is e a s e .

S ymp t o ms D u e t o C h a ng e s i nR a t e o f M e t a b o lis m
S ymp t o ms D u e t o Al t er a t i o ni nt h e C h a ra ct e r o f M eta b o lism .

I mp a i r e d Hor m o ne C o ntro l
U nd e r- A c t ivi t y o f t h e T hy r o id
O v er- A ct ivi t y o f t h e T hy r o id
P i tui t a r y Gl a nd I mp a i r m e nt
D i a b et e s.

Imp a i re d N er v o us C o nt ro l
I mp a i r m e nt s D u e t o Tissu e D es tr u ct i o n
xvi ii CO N T E NT S

I mp a i rm e n t D u e to Al te ra ti o no f N or m a l M e ta b ol is m
N e ura s t h e nia .

D i re c t T rea t m e nt o f N e rv o u s D is o rd ers .

T rea t m e nt of N e rv o us D is o rd ers by Psy ch o t h era py

M a l nu tri ti o n
Po i s o ni ng.

D is ea se D ue to M e ch a ni ca l I nj ur y

S E C TIO N 5 . AS S OC IA T I O N OF O R G AN I S MS

C H A P T ER X V III
A S S O C I ATI O N OF O R G AN IS M S

N a t u re o f Ass o ci a ti o n
Fa mily Gro u ps , He rdS
B ee C o l o n i es
Ants .

A C o mp a ris on Ant a nd M a n
,

S ymbi o se s D e fi n
, i ti o n
C o mm e n s a l ism or M ess ma ti sm
,

P a r a s i t ism
P a ra s i t i c P l a n ts
A nim a l Pa ra si te s
S t ru c t u re o f P a r a si te s .

D e ge n e ra t i o n .

Li f e H is to ri e s
Eff ec ts o f P a ra si te s o nHo s t

C H A P T ER X IX

D IS EAS ES OF P LA N T S

Ec o no mi c I mp o r t a nce of Pla nt D is e a s e
C a use s o f D is ea s e
Th e S o il
Till a ge
Po ll u te d Ai r .

M o is t a nd D ry Ai r
He a t a nd C o ld Ligh t a nd S h a d e
,

Fa s c i a ti o n .

T h e Li v i ng C a u s es o f D is ea s e : I nf ecti o ns

Pi ne G a lls

N a t u re o f D is e a se s D ue to Fu ngi
T h e R u s ts
T h e M il d e ws
S m u ts
B a cte ri a l I nfe c ti o ns
 C O N T E NT S xi x

R oo t T ub ercl es
T re a t m e nt O f D is ea s e i nP l a nts
T h e Pr e v enti o no f D is e a s e
T h e M e di ca l T rea t m ent of D is ea s e
T h e S u r gi ca l T rea t m en t O f D is ea s e
S umm a ry

C HA P T ER XX
D I S E AS E S DU E To H A R M F U L O R G AN I S M S
S o m e K i nds o f O r g a nisms P r o d u ce P o is o ns-
.

W a ys i nWhi ch M i croOrga ni sms C a u s e H a r m i n A nim a ls

Foo d P o is o ni ng
Pre v ent i o no f B o tu l ism .

I ntes t i na l P u tre f a ct i o n
.

T h e M e ta b o li c A ct ivi te s o f P a r a si t i c O r g a nisms
T h e R e l a t i o nof H o s t a nd P a ra si te
C l a ss es of D is ea s e- Pr o du c i ng O r g a nisms
B a cteri a l D is ea se .

Pu s- f o r mi ng O r g a n is ms .

C o mm on C o lds
C a t chi ng C o ld
D is e a s es D u e t o H igh er Fu ngi
D is e a s e D u e t o S pi ro ch e te s
D is ea s es D ue to t h e Fil t er a bl e V i r us e s
P ro to zo a nD is e a s e s
D is ea s e D u e t o M e ta z o a
V i r ul ence .

D ef ens e s o f Or g a n isms
P rev enti o no f I nf ect i o n
T h e O v erco mi ng o f I nf ect i o n
T h e A ct i o no f P h a g o cy t es
Th e I mmu ni t y R ea ct i o n
D iph t h e ri a
Ar tifi ci a l A nti -to xi n
I mmu ni t y
S m a llp o x a nd V a cci na t i o n
D ef e ns e s of C o mmu ni ti e s
C on ta mi n a t i o nof t h e Ai r

D ir ect I n f ecti on

C H A P T ER XXI

I N S E CT S AN D D I S E AS E

B en e fi ci a l a nd H a r mf u l I ns ect s
How I ns e ct s T ra nsmi t D is e a s e
Fli e s a nd T yph o id Fe v er
Fli es a nd S l ee pi ng S i ck ne ss
Fl i e s a nd O t h er D is ea s e s
M o squi toe s a nd M a l a ri a ”
M
M a l a ri a l P a ra si te s Inth e o squi to
xx CO N T E NT S

Y e ll o w Fe v e r a nd M o squi toe s
R e m e di a l M e a s ure s
P l a gu e
Fle a s nd P l a g u e
a .

O the r I nse c ts a nd D is e a s e
M i te s Ti cks a nd D is ea s e
,

Go v ern m e nt S a ni ta t i o n

S E CTIO N 6 . D E A TH

C H A P T ER XX II
D E AT H AN D T HE D U R AT I O N OF LI F E

N a t u re of D ea th .

Hibe rna t i o no r Wi nter S l e e p

S u mm er S l ee p .

E ff e ct s o f Free zi ng
Le ng t h o f Lif e
W e ism a nns Th eo ry’

A ppli ca t i o no f th e T h eo ry
O l d Age M o difi ca t i o n s
I mm e di a te C a us e s o f D e a t h ”
I mm o r ta li ty o f t h e U ni ce llul a r Or g a nisms
D e a t h I nM ul t i ce llul a r O r g a nisms
D ea t h a n A dv a nta g e t o t h e S p e c i e s
S u mm a ry

S E CT I O N 7 . T HE R OLE OF M I CR O O R GA N I S MS

C HA P T ER XX I I I
D E C O M P O S IT I O N

T h e D isp o s a l o f O r g a ni c W a s te s
D eco mp o si t i o n
K i nds o f D e co mp o si ti o n .

T h e A ge nts o f D e c o mp o si t i o n
T h e P ri n cipl e s o f D e co mp o si ti o n .

Alco h o li c Fe rm e nta t i o n
T h e E nzyms C o n ce rn ed

C o mm e rc i a l Us e s o f t h e r o du c t P s
A ce ti c Fe rm e nt a ti o n .

T h e S o u ri n g o f M ilk .

T h e R o t ti ng o f W o o d
C o mm erc i a l Fib e r P ro du c ti o n .

D e co mp o si ti o no f Fa ts a n d O ils
D e co mp o si t i o no f N i tro ge no u s W a s te
T h e S o lid N i t ro ge no u s W a s te s
P u tre fi ca t i o n .

P u tre fyi ng S u bs t a n ce s
T he P ro d u cts of P u tre f a cti o n .
 CO NT E N T S

T h e P to m a i nes

T h e P u tre fyi ng P ro cess

S u mm a ry

C H A P T ER XX I V
T H E FE R TI LI T Y O F THE S OIL

E xh a us ti ono f th e S o il
N a t u r a l Fer tili t y
Fer tiliz er s .

A nim a l W a s t es
M i cro Orga ni s ms ”
T h e N I trI f y I ng B a cter i a
T h e N i tri te B a ct er i a .

T h e N i tr a te B a ct e r i a .

N i tr ifyi ng O r g a nisms a nd High er P l a nt s

E ff e c ts o f C ul t iv a ti o n
Fr ee- livi ng S o il B a cte ri a
B a cteri a S ymbi o t i c wi t h Algae ]
T ub ercl e B a cteri a .

M y corrhiz a
D en i trifyi ng B a c te r i a
S ulphu r B a cteri a . .

T h e C a r b o n C y cl e
Th e N i tro g e nC y cl e
S u lph u r a nd P h o sph or us C y c l e s .

T h e Ener gy C y cl e

PA R T II

S E CT I O N 8 . GR O W TH AN D R E P R O D U CT I O N

C H A P T ER XX V
C E LL- D I V I S I O N
The Grow t h of Or g a nisms
C e ll Gro w t h an d D ivisi o n
The M ech a nism o f C e ll D ivisi on
Th e R e s ti ng N u cl e us
Th e
The
The
B e h a vi or o f th e C h r o m a t i n
For m a t i ono f th e N u cl e a r S pi
S pli t t i ng o f th e C h r o m o s o m e s
.

n
dl
e

The A na ph a s e
The T el o ph a s e a nd R eco ns tr u cti on of th e N u cl e us
The I ndividu a li t y o f t h e C h ro m o s o m e s
Th e D ivisi o no f th e C e ll B o dy -

S u mm a ry
x xii CON TE NT S

C HA P T E R XX V I
R E PR O D U CT I O N IN A N I M ALS
R e p ro d u cti o n
S pon ta n eo u s Genera t i o n
B udd i n g
Fissi on
R es ul ts o f As e xu a l R e p ro d u ct i o n
R e ge n era ti o n

R e ge n er a t i o nin High e r A ni m a ls
S e xu a l R e p ro d cti o n
u
T h e S e x C e lls
A ccess ory Se x O rg a ns
Fe rt ili z a t i o n
R es u l ts o f Fer t ili z a t i o n
Individ u a li t y of th e C h ro m o s o m e s
Fert ili z a ti o nO f U ni cell ul a r A ni m a ls
E a rly D e v e l o pm e nt
T h e Ga s tr ul a a nd B o dy L a y ers
La ter D e v e l o pm e n t
V erte b ra te D ev el o pm ent
Gro w t h
D i ff ere nt i a ti o n
M e ta m orph o sis
C a re o f E ggs
N o u r ishm e nt o f th e Y o u ng
S eco nd a ry S e x u a l C h a ra c ters
B o dy a nd Ge r m C e lls .

Ea rly H is t o ry o f Ger m C ell s .

R e du c i ng D ivi si o n
R a nd o m Ass o r t m e n t o f C hr o m o s o m es
M a t u ra t i o no f t h e E gg
D ete r mi n a t i ono f S e x

C H A P T ER XX V I I
R E P R O D U CTI O N IN P L AN T S

Se xu a l a nd As e x u a l R e p ro d u cti on
Typ es o f R e p ro du ct i o ni nP l a nts
Ase xu a l R e p ro d u c ti o ni nth e Al gze
Ess e nt i a ls o f As e x u a l R e p ro d u c t i o n
S e xu a l R e p ro d u c tio nby m e a n s o f Ga m e t es
T h e D i ff ere n t i a t i o no f Se x
Al te rna t i o no f Ge ne ra t i o ns
AnE x tra Ge n e ra ti o n .

Ga m e to phy te a nd S poro phy te

Fe rns
T h e S tr u ctur e of Ferns
He te ro sp o ry .

Re p ro du c ti o nby See d .

T h e O vu le a nd Fe m a l e Ga m e to phy te
T h e An t h ers a n d M a l e Ga m e to phy te
Fert ili z a t io n, E mb ry o, a nd E nd o sp erm
 CO N T E NT S x x ii i

T h e S ee d
D o r m a ncy o f S ee ds
T h e C h ro m o s o m e H is to ry i nP l a n ts a n d A nim a ls .

R e du cti o na nd Se gr e g a t i o no f C h ro m o s o m e s i nP l a nts
V e geta t iv e R ep ro d ct i o n
u

S E CTI O N 9 . H E R E D I TY
C H A P T ER XX V I I I
MEN DEL ’
S LA W S
Va ri a ti o n
P u re Li nes a nd Hyb rids
M o difi ca t i on
s
M u ta t i o ns
R eco mbi na ti ons
Here di t y .

M end el ’
s D is co v er y o f t h e La ws o f H ere di t y
T h e La w of S e gre g a t i on
A S impl e I llus tra t i o n .

T h e P u r i t y o f Ga m et e s .

Th e R eco mbin a ti o n R a t i o

T h e B ea rers o f H er e di t y a r e
T h e C h ro m o s o m e s C o n
.

h ro o
C
t a i n th e Gene s
so s mm
e .

S e gre g a t i o no f C h ro m o s o m e s a nd G e n es

D o mi n an ce a n d t h e 3 1 R a ti o
T h e L a w of S e gre g a t i on .

T h e R e l a t i on of Gene s t o C h a ra c ter s
Are Gene s t h e S o l e B e a rer s O f H er e di t y ? .

Are Acq u i re d C h a ra ct ers I nh er i t e d ?

C H AP T E R XX I X
T H E L AwO F I N D E P E N D E N T A S S O C I AT I O N

The I nd e p end ence o f C h a r a ct er s .

The C o mbi na t i o ns o f I nd e p e nd ent C h a ra ct e r s i nF 2

T he M e ch a nism o f R ea ss ort m ent .

Th e C h r o m o s o m e s o f t h e T a ll P u r pl e H yb rid
The Ga m e te s of th e T a ll P u r pl e H yb r id .

T he C o mbi na t i o no f th e Fo u r K i nds of Ga m e t es
The 9 : 3 : 3 z l
Geno t yp es a nd P h e no t yp e s
T h e La w o f I nd e p end ent A ss o r t m e nt of Ge ne s a nd C h a ra cters

C H A P T ER XXX
LI N KA G E P H E N O M E N A
T h e Thi r d L a w
T h e Fo u rt h L a w
T h e V ine g a r Fl y

T h e SEX C h r o m os o m e s of D ro s o phil a
T h e D et er mi n a t i o no f S e x
xx vi CO N TE NT S

Li n k a ge inV i ne ga r Fly C ross e s .

The Ba ck C m
B a c k C ro ss I nv o lvi n g I nd e p e nde nt C h a ra cte rs
Li nked Cha ra cte rs inth e Firs t Hybrid Ge nera tion
Ba c k - C ro ssi ng L ink e d C h a ra c t ers
T he Se grega t ionof Link ed Ge n es
T he S t ren gt h of Lin a ge k

The Fif th La w Of Heredi ty

The S ixt h La w of Here di ty

Re d- Ey ed Fema le X W hi te - Eyed M a le
Red- M ed Ma le X Whi tG Ey ed Fema l e
S u mm a ry o f Pri ncipl e s o f He re di ty

C HA P T ER XXX I
T HE C HAR A C T E R EX P R E S S I O N OF GE N E S
Th e D i ff e re nti a l Ge ne s o r
T h e R e l a t iv e Po si t i o no f Ge n es .

T h e M u ta t i o n s of Ge nes .

T h e Ga m e ti c R a t i o s o f H yb rids
T h e N umb er o f Ge n o t yp e s o f t h e S eco n d Genera t i o n
T h e P hen o ty p e s o f t h e S eco n d Ge ne ra t i o n .

C o mpl e m e n t a ry Ge n es .

S uppl e m e nt a ry Ge n es

I nhibi t o ry Ge n es .

C umul a t iv e Ge n es

S umm a ry

C H A P T ER XXX I I
PL ANT BRE ED I N G

T he His to ry o f P l a n t B re e di ng
T h e E co n o mi c I mpo rt a n ce o f P l a nt B ree di ng
T h e P ro bl e ms o f P l a nt B re e ding .

T h e M e t h o ds o f P l a nt Bre e di n g .

P u re Li n e S e l ec t i o n
I mp ro vi ng I n di a nC o rn
B re e di n g C o t to n
S e e dl e ss O ra n ge s
D is e a s e R e s ist a n ce
Fros t R e sis ta nce .

S u mm a ry

C HA PT ER XXX II I
A N I MA L B R E E DI N G

R e p ro du cti o ni nHigh er A nim a ls

O nI n te rb ree di ng .
 CO NT E NT S

O nth e E ff ect s o f S el e ct i o n
I nh eri ta nce O f C o a t co l o r i nGui n
-
ea P igs 0 0 0 0 0 0 0 0 0 0 0

I nh er i ta nce o f H o o d e d P a t terni nR a t s
Le t h a l Ge ne s i nY e ll o w M i ce . .

h
In e tr i a nce o f Wi n t e r E gg P ro du ct o n
i by Fo wls ’ . 0 0 0 0 0 0 0 0 0 0 0

I nh er i ta nce i nD a i r y C a t t l e 0 0 0 0 0 0 0 0 0 0 0

I nh eri ta nce o f S p e e d i n H o r s e s 0 0 0 0 0 0 0 0 0 0 0

S umm a ry

SE CTIO N 1 0 . EV O L U T I O N

C H A P T ER X XX I V
EV I D E N CE S OF EV O LU T I O N

E v ol u t i o nD e fi ne d 0 0 0 0 0 0 0 0 0 0 0

S o u rces o f E vid e nce 0 0 0 0 0 0 0 0 0 0 0

E vid ence s f r o m P a l eo nt o l o gy o o o o o o o o o o o

His t or y o f th e Hor s e .

E v o lu t i ono f C a m els
C l a ssifi ca t i o n”
T h e M o d e r nS s y t e m o f C l a ssifi ca t i on
S ci ent ifi c N a m e s
C o mp a ra t iv e A na t o my .

R u dim e nta r y o r V e s t igi a l O r g a ns 0 0 0 0 0 0 0 0 0 0 0

E vid e nce f r o m E mb ry o l o gy
’

D e v el o pm e nt o f V er t e b r a t e s . 0 0 0 0 0 0 0 0 0 0 0

Geo gr a phi ca l D is tr ib u ti o n
A rtifi ci a l S el ect i o n
Genet i cs

C H A P T ER XXX V
T HE OR IE S OF E V O LU TI O N

Pre- D a rwi ni a nS ci ence .

T h e I nflu ence o f D a r wi ns W o r k
’
.

Fa ctors o f D a r wi ns T h eo r y
’

O v er p ro du c ti o n 0 0 0 0 0 0 0 0

S ta t i o na r y N umb er s o f I ndividu a ls
T h e S tr uggl e f or E xis t e nce
In terrel a t i o n s of O r g a nisms . 0 0 0 0 0 0 0 0 0

V a ri a ti ona nd H ere di t y .

Th e S u r viv a l of th e Fi t t es t
A ppli ca t i onof th e T h eory
S e xu a l S e l ect i o n.

La m a r cki sm
T h e N e w L a m a rcki a nS ch o o l
Ort h o g ene sis o r D e t er mi na t e E v o lu ti o n 0 0 0 0 0 0 0 0 0 0 0
x xvi CO NT E NT S

C HA P T E R XXX V I
T H E R E S U LT S OF E V O LU T I O N

A d a p ta tio n
C on v erge nt A d a p ta ti ons i n A nim a ls .

C on v e rge nt A d a p ta t i ons a m o ng P l a nts

O t h er C l a ss e s o f A d a p t a ti o ns
Foo d S e cu r ing A d a p ta ti o ns
- .

P l a nts A d a p te d to A bs o r b Ga s e s W a ter , , a nd S a l ts
I ns e c t iv o ro u s P l a nts
A d a p ta ti o ns for S el f D ef ens e
-

D ef e ns e o f Y o u ng
S elf D e f e ns e i nP l a nt s
-

A d a p ta ti ons f or R iv a l ry
R iv a l r y a m o ng P l a n ts
A dj us tm ent A d a p ta ti o ns i n A nim a ls
P l a nt
C o l o rs o f A n im a ls
Pro te ctiv e R es e mbl a nce
C o l ors o f P hysi o l o gi ca l V a lu e
W a rni ng C o l ora ti o n
M imi cr y .

Fl o w ers a nd I ns ect s

SE CTI O N 1 1 . TH E D I S TR I B UTI O N OF O R GAN I S M S

C H A P T ER XXX V I I
P L AN T D I S T R I B U T I O N

T h e Fa cts o f D is trib t i o n u
Th e Gen era l C a ses o f D is t rib t i on
u u
E v o lu ti o na n d D is tribu t i o n
M igra ti o na nd D is tribu t i o n
A d a p ta t i o na nd D is tribu ti o n
Fl o ra s a nd Fa na s u
V e g e ta t i o n
T h e B a rri e rs to U niv ers a l D i s tri b ii ti o n
T a ngibl e B a rri ers .

I nta n gibl e B a rri ers .

B i o l o gi ca l B a rri e rs
T h e G rea t P l a n t R ea l mS u
T h e Ho l a rc ti c R ea l m
V e g e ta ti o no f th e U ni te d S ta te s a nd C a na d a
D e cidu o s Fo re s t
u
We s te rnC o nif ero us Fo re s t
Gra ssl a nds
D es e rt
C ha p a rr a l
S w a mps .
 CO NT E NT S

C H A P T ER XXX V I I I
D IS TR IB U TI ON OF LA N D A N I M A L S
Exte nt of th e S u bj ect
M ea ns of D isp ers a l
T a ngibl e B a rri ers
I nt a ngibl e B a rr i ers .

La ws a nd R es u l t s o f D is trib u t i on
A nim a ls of O cea ni c I sl a nds
Z o Ol o gi ca l R e a lms
H ol a rct i c R ea lm
N e o tro p i ca l R ea lm .

Ethi o pi a nR ea lm
O ri ent a l R e a lm
T h e A us tra li a nR e a lm
R ea lm S ubdi visi ons .

M igra t i o n s of A nim a ls

C H A P T ER XXX I X
D I S TR IB U TI O N OF LI FE IN T HE O CE A N
P hysi ca l C h a r a ct er s
T e mp era t u re of th e S e a
E ff e ct o f O ce a nC urrent s
P r e ssur e
Ligh t .

D is tr ibu t i o nof S h ore P l a nt s

Fl o a ti ng P l a nt s
P l a nk t o n
U ni ce ll ul a r Or ga nisms a nd B o tt o m D e p o si t s
S o m e P e l a gi c M a ny - C e ll e d A nim a ls
O r g a ns o f Fl o t a t i on
A nim a ls of th e S ea Fl o or .

P h o sph ore s cence

S umm a r y

SE C T I O N 1 2 . MAN
C H A PT E R XL

TH E EVO L U TI O N OF M AN
Tim e R e qu i re d f o r M a ns E v o lu t i on ’

P l a ce o f O r igi n”
Ea rli er H is t or y o f theH um a nR a ce
T h e J a v a M a n Ap e
Th e P il t d ownMa n
Oth er T ra ces of Ma ns A nt iquity
’

Th e N e a nd ert h a l M a n
M o d ernM a n
C ro M a gno nM a n
-
.

La t er E u ro p ea nR a ce s
M a ni n A m er i ca
S u b S p ec i e s o f M o d ernM a n
-
.

M a ns Fu t ure
’
xx viii C O NT E NT S

C HA P T ER X LI

H U M A N I N H E R IT AN CE

Hu m a nT ra i ts
D iffi c ul t i e s o f S tu dyi ng Hu m a nI n h eri t a nce
Heri ta bl e C h a ra c te rs i nM a n
P hysi ca l C h a ra c t e rs .

T h e I nh e ri ta n c e o f Ey e - C o l o r

B ra c h y d a c ty fy .

Po lyd a c t yly a n d S y nd a c tyly

S k i nC o l o r
S t a t u re a n d W e igh t
C o l o r- B li n d ne ss
C re ti nis m

C HA P T ER X LII
I N HE R IT A N C E OF M E N T AL T R AI T S
(B y Le wis M T er m a n)
.

M e nta l a s w e ll a s P hysi c a l T ra i t s a re H eri t a bl e

A m o u nt a nd N a t ur e of V a ri a bili t y i n I nt e llig ence
V a ri a bili t y i n O t h er T r a i t s
S t a t is ti ca l M e t h o ds Us e d i n t h e M ea su re m e n t o f R e s e mbl a n ce

Fa mily R e s e mbl a nce i n I nt e llige nce

T h e La w o f Fili a l R e gre ssi o n
M e nta l R e s e mbl a nce o f T wi ns
Fa mi ly R e s e mbl a nce no t Wh olly A c co unt e d f or by Envi ro n
.

m e nt
T h e I nh e ri ta n c e o f Ge n ius .

I nh e ri ta n ce o f Fe e bl e- M i n d e dne ss
T h e K a llik a k Fa mily
In h e ri ta nce o f O th e r Fo rms o f M e nta l D ef e ct
En d o wm e nt o f t h e D i ff e re nt S o ci a l a nd O cc up a t i o na l C l a ss es
T h e D ecre a si ng Fert ili t y o f S u pe ri o r S to c k .

T h e I m po r ta n c e O f E ug e n i cs
T h e E f e c t o f U nd e s i ra bl e I mmig ra t i o n
f
E ge ni c s a nd M a rria g e
u
T h e I nlu e nce o f En
f vi ro nme nt a nd E du ca t i o n p o n M e nta l Per u
m
f o rma n ce

He re di t y S e ts t h e L i ts i
M o difi ca ti o ns o f M e nta l T ra i t s no t T ra nsmi t te d

C HA PT E R X LI II
M AN P LA C E
'
S IN N AT U R E

M a nth e A p e x o f t h e A nim a l S e ri e s
M a ns S imil a ri t y to t h e High e r A nim a ls
’

The Fa c t of Progre ss
H um a nR a ce U nch a nge d
Ex pl a na t i o no f Pro g re ss .
 CONT E NT S x x ix

P hysi ca l C h a ra c t er is ti cs
In t ell ect u a l C h a r a ct er is ti cs

C l o t hi ng .

T h e C ul t iv a t i o no f t h e S o il
L a ngu a g e
La ngu a g e a s a Fa c to r i n I ndividu a l P ro g re ss
T h e S i gn i fi ca nce of La ngu a ge t o R a cia l Pro gress
E m o t i o na l C h a ra ct er is t i cs .

T he In t era ct i o no f th e E m o t i o ns wi t h I nte llig e nce

C H A P T ER X LI V
HU M A N P R O G R E S S
Pa s t P r o gre ss t h e R e a liz a t i o nO f I nh er e nt P o ssibili ti e s
I nh e re nt P o ssibili t i e s a re no t y e t f ully R e a liz e d
A nim a ls Ex ercis e a Limi te d C o ntro l o v er th e E nvi ro nm ent
M a nL a r g e ly C o nt ro ls Hi s E nvi ro nm ent
T h e D e t a ils o f P r o g r e ss
T h e S a t isf a ct i o no f t h e P r im a r y N ee ds a nd t h e E n ri c h m ent of

L if e
T h e Fo o d S upply
S e cu r i t y O f L if e
T h e D e s tr u ct iv e C o mp e t i ti o no f M a nwi t h M a n
C oOp er a ti o na B i o l o gi ca l N e ce ssi t y f o r P r o g re ss
Le ss ene d C hil d M or t a li t y
E du ca t i o n
I s o l a t i on
P r i nti ng
T e le ph one, T el e gr a ph , a nd R a di o
T ra nsp ort a ti o n
A d e qu a te P ro du ct i o n
H igh I d e a ls
T h e I mpul s e t o C o mb a t
S p or t s
C o mm er ci a l S u ccess
Fu t u re P ro gr e ss
R a ci a l B e t t er m e nt
 G ENE R AL BIO LO GY

C HAPTER I

I N T R O D U C TI O N

B I OLOGY is th e scie nce which t re a ts of livi ng thi ngs .

Th e numbe r a nd dive rsity of org a nisms i n th e w orld is

so g re a t a s to be a lm o st in credible I t is s a id th a t the re
.

a re m ore th a n a h a lf milli on ki n ds or S pecies o f insect s

a l on e a l re a dy k now n a nd th a t th e re a re u nd oubtedl y a
g re a t m a ny a s yet u ndisc o ve red Of D i a t oms a s ort of
.
,

on -
e celled pl a n t th e re h a ve bee nna med m ore th a n ten
,

th ou sa nd spec ies Alt ogeth e r i t is well with inth e limit

.
,

to sa y th a t th e re a re m ore th a na milli onki nds of a nim a ls

an d pl a nts
.

N o t only a re org a nisms imme nsely nume rous a nd

,

highly dive rsified b u t th ei r a ctivities a re a ls o ext remely

nume rous a nd c omplica ted E ve n a pp a re ntly S imple
.

p ro cesses frequ e ntly tu rn ou t to be c omplex a nd ve ry

difficult to unde rst a nd Th is m a kes it necess a ry i norder
.

to a cc omplish mu ch i n th e w a y o f n ew kn owledge for

i nvestig a t ors to c onfi ne th ei r a tte nti on not me rely to a

few ki nds of org a nisms bu t a ls o to a S ingle one or a few
of thei r a ctivities T o a dd to th e diffi culty of the m a tte r
. ,

it is usu a l to h a ve a gre a t m a ny di ffe rent ch emic a l re

a cti on s g oi ng oninone sm a ll bi t of livi ng m a tte r so cl ose
t ogeth e r th a t it is difficult to re a lize th a t th ey ca np os
Sibly a v o id i n te rfe ri ng with one a no the r For th is s a me
.

re a s o nit is di ffi cult to is ol a te a sin gle a ctivity for study .

I t is v e ry obvi ou s th a t no pe rs onc o uld p ossibly bec ome

f a mi l i a r with a l l th e di ffe re nt kinds of livi ng bei ngs
‘

It is necess a ry the refore to subdivide so v a st a subj ect

, ,
.

3
4 I NT R OD U C TI ON
One ofthe m ost na tu ra l of th ese su bd ivisi ons sep a ra t es
an im a ls a nd pl a nts int o two disti nct scie nces Z oOlogy ,

an d B ot a n y V e ry lit tle i ntellige nt w ork c ould be d one

.

wi th such l a rge numbe rs o f pl a nts u nless th ey we re fi rst

c a refull y cl a ss ified The re must a ls o be a d efinite a nd
.

fix ed pl a n of na ming them so th a t they ca n be re a dily

an d ce rt a i nly refe rred to Th us the re a rises a scie nce
.

of C l a ssific a ti on s ometimes c a lled T a x on

,
omy .

B o th B ot a ny a nd Z oOlogy a re fu rthe r subdivid ed

a c c ord in g to the p oi nt o f V iew int o seve ra l scie nces ,
.

Whe re th e chief i nte rest lies i n f orm a nd st ructu re the

an im a l or pl a nt must usu a lly be cu t up for ex a mi na ti on ,

wh ich o fte n requ ires speci a l t ools a nd S peci a l t ra i ni ng .

Th is c onstitutes th e scie nce of M orph ol ogy a s distin ,

gu i sh ed f rom the study o f fu n cti on or Physi ol ogy S tu ,

.

de nts of he red ity requi re a d iff e re nt s ort o f equ ipme nt

for thei r i n vestig a ti ons They must h a ve g a rde ns a nd
.

b reed ing pe ns to re a r thei r pl a nts a nd a nima ls a nd ma y

-
,

requi re a l a b ora t ory i nwh ich to c a rry ou t mic ro sc opic a l

o r chemic a l in vestig a ti ons This c onstitutes the scie nce

.

of G e n etics S ome i nvestig a t ors a re c once rned with th e

.

rel a ti on s of pl a nts a nd a nim a ls to one a no th e r a nd to

thei r e nvi ronme nt a nd so do m ost o f th ei r w ork I nthe
,

field They a re c a lled Ec ol ogists Th e obj ect i n men

. .

t i on ing these speci a l scie nces is to S h o w th a t e a ch re

qu ires a speci a l equipme nt a nd m uch t ra i ni ng so th a t ,

it is no t e a sy for one i nd ividu a l to do a ctive w ork inm ore

th a n one field o f study .

Besides th e pu re scie nces o f Bi ol ogy th e re a re m a ny

fields i n wh ich bi ol ogic a l p ri nciples a re a pplied p ra cti
c a lly M edic ine is one o f th e m os t c onspicu ous o f such
.

a pplic a ti o n s o f th e results o f pu re scie nce to th e a ff a i rs

o f eve ry da y life S a nit a ry bi ol ogy c once rned wi th
-
.
,

d ra ina ge the a b a teme nt of i nsec t pests a nd simil a r

,

m a tte rs is a noth e r Public h e a lth w ork i nv olvi ng th e

, .

regul a ti on o f build in gs the h a ndli ng of food a nd m a ny

, ,

othe r th in gs is still a no th e r Pl a nt b reedi ng a nd pl a nt

,
.
 T HE LIVING S U B S TAN CE

p a th ol ogy a re b oth of g re a t imp ort a nce i nm ode rn a gri

cultu re the one to p rovide new a nd imp roved s orts of
,

pl a nts a nd the o the r to fi nd me a ns of checking the e nor

mous l osses due to the a tt a cks of p a ra sitic fungi a nd
b a cte ri a on c rop pl a nts A list of this s ort migh t be
.

gre a tly exte nded bu t it is p rob a bly a lre a dy cle a r th a t

,

m ode rn B i ol ogy h a s bec ome a v a st a nd c omplic a ted

subj ect one in wh ich the ordi nary citize n ca n h ope to
,

know only th e ge ne ra l f a cts a nd p ri nciples .

G EN ER A L B I OLOGY a ims to b ri ng t ogethe r a nd p rese nt

to the n onspeci a list th ose a spects o f Bi ol ogy which it is
-

imp ort a nt th a t eve ry one S h ould unde rst a nd i n orde r to

pe rform h is du ty a s a citize n i n a n i ntellige nt m a nne r .

Wh ere a s the p ro fessi ona l bi ol ogist must limit his field

an d speci a lize i norde r to keep th e ra nge o f i nvestig a ti on
with in such b o u nds a s will p e rmi t h im to bec ome a
m a ste r o f a ll th a t is know n c once rni ng it th e ge ne ra l
,

bi ol ogist st rives to b ri ng t ogethe r th e m ost imp ort a nt

f a cts a nd p ri nciples a nd so rel a te them th a t the a ve ra ge
pe rs on ca n useful l y a pply them to h is ordi na ry a ff a i rs .

Ina neleme nt a ry c o u rse m a ny inte resti ng a nd imp ort a nt

subj ects must be omitted or cu rt a iled bec a use they a re
too di ff icult or th e stud ent s p revi o us t ra i ni ng h a s bee n
’
,

i na dequ a te or b ec a use the time is not su fficie nt In

,
.

c onformity with th ese limit a ti ons the f oll owi ng subj ects
h a v e beench ose nfor c onside ra ti onint h e belief th a t th ey
a re of such imp ort a n ce th a t no educ a ted pe rs on ca n
a ff ord to be ign oran t o f them °

.1 T H E L I V I N G S U B S TA N CE P R OTOP LA S M
, . I t will be
-

S h ow nth a t eve ry livi n g thi ng is a live bec a use it is com

“

p osed i n wh ole or p a rt o f a peculi a r subst a nce c a lled

p rot opl a sm Fu rthe rm ore this p rot opl a sm is no t only
i

of u n ive rs a l occu rre nce but it is ve ry S imil a r eve ni nthe

m ost dive rse org a nisms I t is ce rt a i nly w orth ou r while
.

to u n de rst a nd the physic a l chemic a l a nd physi ol ogic a l

, ,

ch a ra cte ristics of this m ost w onde rful of a l l subst a nces .

.2 GREEN P LA N T S pl a y a peculi a r a n d m ost imp ort a nt

6 IN T R O DU CTI O N
rOle inna tu re i nth a t th ey a re the ch ief s ou rces d i rectly ,

or i n di rec tly o f ne a rly a ll th e e ne rgy (a side f rom th a t

,

ge ne ra ted by f a l ling w a te r a nd w ind p owe r ) wh ich ma n

uses i n his Ope ra ti ons F rom them a re de rived a ll the
.

foods a nd a ll the fuels besides a h os t o f o the r u seful

,

p rod uc ts I t is su rely imp ort a nt the refore to u nde r

.
, ,

s ta nd h ow they a re a ble to c a tch a nd st ore up th e e ne rgy

o f the su n ligh t in foods a nd fuels .

3 T H E M A I N T E N A N CE O F L I F E I N A N I M A LS AN D
.

N O N G REEN P LA N T C ELLS is wh olly depe n

-
de nt on f ood
m a nufa ctu red by g ree n ce l ls Th is they use to build
.

thei r b odies a nd f ro m i t they de rive th e e ne rgy to do a ll

s orts of w ork Inlibe ra ting th e e ne rgy st ored up i nf oods
.

an im a ls d ec omp ose them int o th e c a rb on di oxid a nd

w a te r f ro m wh ich they we re origi na lly m a nuf a ctu red .

S ome f ood s a re used to bu ild living p ro t opl a sm This .

ve ry g ra d u a lly b re a ks d ow ni nth e c ou rse of its life a cti vi

ties a nd a p orti on c ont a i ni ng th e eleme nt nit ro ge n is
th row noff a s w a ste a nd exc reted f rom the b ody E v entu .

a lly it a ls o retu rn s to th e s oil to be used ove r a g a i n .

4 AD J U S T M E N T O F OR GA N I S M S T o T H E I R E N V I R O N
.

M E N T is n ecess a ry to th ei r c onti nued existe nce A pl a nt .

must be a ble to d i rect its roots int o th e s oil a nd its le a fy

stems i nt o th e a i r if th ey a re to p e rf orm th ei r p rope r
fu ncti ons Inlike m a nne r a n a nim a l must be a ble to
.
,

fin d a nd rec o gn ize suit a ble food i nsuffi cie nt qu a ntity or ,

it st a rves Th e ve ry f a ct th a t they ca ndo these th i ngs

.

sh ows th a t the re is s ome s ort o f mech a n i sm for secu ring

p rope r a d j ustme nt of org a nisms to th e e nvi ronme nt I t .

will be sh ownth a t this d epe nd s once rt a i nch a ra cte ristics

p osses sed by a ll p ro t opl a sm bu t wh ich h as bec ome h ighly
,

speci a lized i n ce rt a i n org a ns of th e h igh e r pl a nts a nd

an im a ls I t ma y a p p e a r s omewh a t st a rtli ng to th e b e
.

ginni ng stude nt to re a lize th a t h is org a ns of speci a l se nse

(sigh t t a ste smell ) a nd th e system o f muscles a nd b ones
, ,

an d j oi nts th e org a ns o f l o c om o ti on a re bu t th e ul tim a te

, ,

devel opme nts o f a food getting mech a nism a nd th a t they

-
 IN T E R A CTI O N OF O R GANI S M S 7

r est fu nd a ment a lly on the p owe r of a ll p rot opl a sm to

resp on d to ce rt a i n o utside i nflue nces a nd to execute
f orcible m ovements by the utiliz a ti onof e ne rgy libe ra ted
from foo d Th e ne rv ous system a nd th e b ra ini tself a re
.

likewise to be i ncluded i n this mech a nism of a dj ustme nt

of the b ody to its su rro u n dings a nd o f its v a ri ous p a rts
to on e an o the r .

5 T H E I N TER A CT I O N O F OR GA N I S M S onon
. e an o the r is

likewise a m o st imp ort a nt subj ec t E ve ry pl a nt a nd .

an im a l is m ore or less i nflue nced by its neighb ors but i n ,

s ome c a ses th e a ssocIa ti on is so cl ose th a t b oth be nefit

from i t or one be nefits a t th e expe nse of the o the r The
, .

forme r c onsti tu tes th e rel a ti on of symbi o sis (from th e

G reek sym with a nd bi os life ) a nd th e l a t te r th a t of
,

p a ra sitism The fi rst is less c omm ona nd of S ligh te r i m

.

p ort a nce Th e l a tte r r el a ti on very c omm only gives rise

.

t o a c on diti ono f dise a se i none of the p a rties to the as s o

ci a ti on This is of e norm o us imp ort a nce i nb o th pl a nts
.

an d a nim a ls E ve ry ye a r milli ons of d oll a rs w orth o f

.

c rops a re l ost th rough the a tt a cks o f p a ra sites The l oss .

an d su ffe ri ng d ue to the a c ti on of p a ra sitic b a c te ri a on

an im a ls i nclud ing ma n is too f a mili a r to need m ore
th a nb a re me nti on .

6 D EAT H c o mes to a ll highly org a nized livi ng bei ngs

.

s oone r or l a te r E a ch begi ns a s a si ngle sm a ll bit o f liv

.

i ng m a tte r (a cell ) which by repe a ted divisi ons a nd

S peci a liz a ti on fi n a lly g rows i n to th e m a tu re i ndividu a l .

This ma y live bu t a few h o u rs or it ma y li ve thi rty cen

tu ri es bu t i n the en
, d i t re a ch es its sp a n o f life Of .

co u rse m o st o rg a n isms meet de a th i n na tu re from v i o

n
le ce — In j u ry o r dise a se — but n eve r theless it h a s n ot

been p ossible so fa r for the bi ol ogist to p rol ong life i n

defi nitely i n a ny highly org a nized t ype I t is i n conse .

q u en ce a ve ry i nte resti ng a nd imp ort a nt p roblem to

a sce rt a in j ust wh a t a re th e c on diti ons wh ich dete rmine
th a t p ro t opl a sm is a live one m ome nt a nd de a d th e next .

7 T H E D E CO M P OS I TI O N OF ORGA N I C R EM AI N S is on
. e
8 I NT R O DU CTI O N
of the p rime necessities for the c o nti nued existe nce of
life one a rth O the rwise th e e a rth w ould S h ortly bec ome
.

c ompletely c ove red with de a d b odies The ra w m a te ri a ls .

which a re c ombi ned by t h e g ree np l a nts must eve ntu a lly

be re tu rned to th e a i r a nd s oil Alth o ugh these decom .

p ositi o ns a re chemic a l re a cti ons like th ose occu rri ng i n ,

living b odies they do not o ccu r sp ont a ne o usly a ny m ore

,

th a n w a te r a nd c a rb on di oxid w ill sp ont a ne ously form

sug a r In b o th c a ses th e p rese nce o f living org a nisms
.

is necess a ry u nde r ordina ry na tu ra l c onditi ons D ecom .

p ositi on is b rough t a b ou t by sm a ll nong ree n pl a nts -

chiefly a l th o ugh a nim a ls i n libe ra ti ng e ne rgy for d oi ng

,

w ork retu rnl a rge qu a nti ties o f w a te r a nd c a rb ondi ox id

to th e s o il a n d a ir The p rope r t re a tme nt o f s o ils to
.

secu re the m o st desi ra ble g rowth of th ese sm a ll pl a nt s

an d th e m a x i mum e n richme n t o f the s o il th ro ugh their
a cti on f o rms a m ost i nte resti ng ch a pte r i n m ode rn
a chieveme n t .

8 T H E R EP R ODU CT I O N O F P LA N T S AN D A N I M A L S is j u st
.

as n ecess a ry a s th e ir a bility to live a nd m a i nt a in them

selves The me a ns by wh ich this is a tt a ined a re m ost
.

v a ried inte res ti ng a nd imp ort a nt In f a ct th ey a re so

, ,
.

dive rse th a t they ca n be t re a ted in a b ook o f this s ort

on ly i n a m o st ge ne ra l wa y .

9 H ERED ITY
.
— Alth o ugh so f a mil i a r a s to h a ve l ost
.

i ts novelty i t is neve rth eless a m o st c u ri o us a nd i nte rest

,

in g f a ct t h a t o ff sp ri ng cl o sely resemble thei r p a re nts but

a re n eve r ex a ctly like them Th is is s a id to be due to .

h e red ity Alth o ugh o u r k nowledge o f the l a ws o f h e

.

redi ty h a s bee na cqui red a lm o st wh olly i nth e l a st twe n ty

ye a rs the l a st teno f th em h a ve see nm ore ra pid a dv a nce
,

i nthe u n de rst a ndi ng o f the me ch a nism o f he red ity th a n

i na lm ost a ny o th e r li n e o f bi ol ogi ca l i nvestig a ti on N o t .

a l on e f rom its h um a na nd scie ntific i nte rest bu t a ls o on

a cco u n t o f its gre a t p ra ctic a l u tili ty (i nb reedi ng ec onomic
pl a nts a nd a nim a ls ) d oes this subj ect dese rve discussion
inthis c ou rse .
 E V O LUTI ON 9

1 0 EV OLUTI O N
. .
— This te rm is used by bi ol ogi sts to
desc ribe th e f a cts th a t the nume ro us v a rieties of b oth
an im a ls a nd pl a nts ca nbe re a dily a rra nged i nt o m ore or
less c omplete se ries g ra du a lly i nc re a si ng in c omplexity
from Simple one celled f orms a t one end to the m ost
-

h ighly org a nized pl a nts a nd a nim a ls a t th e o the r end ;

an d fu rth e r th a t these se ries cl osely p a ra llel the a rra nge
,

me nt o f the f ossil rem a i ns o f p a st ge ol ogic a ges which

a re f oun d embedded i nth e rocks Si nce it is obvi ous th a t
.

these se ries rep rese nt a g ra du a l de vel opme nt of org a n

isms m a ny the ories h a ve bee np rop o unded to expl a i nthe
,

f a cts of ev olut i on Th e ory a nd fa ct must be c a refully

.

disti nguished by th e stude nt Th e se ries of pl a nts a nd

.

an im a ls a re u nquesti oned f a cts wh e re a s th e expl a na ti ons

,

a re on ly m ore or less p rob a ble the ories Th e Obse rv a ti ons

.

an d expe rime nt a l d a t a wh ich h a ve bee n c ollected withi n

the l a st twe nty ye a rs h a ve te nded to elimi na te m a ny of
these the ori es bec a use they do no t a gree with ou r new
knowledge a nd to s tre ngth e n o the rs bec a use they a re
,

m ore i na cc ord with it .

E v oluti onis li n ked in th e mi nds of m ost p e ople with

th e na me o f Ch a rles D a rwi nbec a use h e wa s the fi rst to
pu t cle a rly before th e w orld evide nce i n supp ort of the
ide a th a t a ll p ro t opl a sm is r el a ted ; th a t the p resent
forms of life a re desce nded f rom p r evi ously existing
forms ; a nd th a t i nth e c o u rse of desce nt v a ri a ti ons h a ve
occu rred wh i ch we re i n he rited ; a nd fi na lly th a t th ose
,

i nhe rited v a ri a ti ons wh ich we re well fitted to th ei r en

v i ron ment h a ve su rvived Th is ide a h a s p roved a nd
.

still is a m ost p ote nt a nd fruitful stimulus to bi ol ogic a l

i nvestig a ti oni na l l fields .

1 1 T H E D I S TR I B UTI O N OF A N I M A L S AN D P LA N T S h a s
.

a ls o l on g offe red i nte resti ng p roblems b o th to th e l a ym a n

an d the scie ntist . Th e bi ol ogist must a cc ou nt for two
s orts o f fa cts In th e first pl a ce h e fi nds th a t widely
.

sep a ra ted regi ons of th e e a rth with simil a r clim a tic a nd

s oil c onditi ons h a ve a nim a ls a nd veget a ti onof ve ry S imi
10 I N T R O DU C TI O N
la r appe a ra nce a l th ough th e a ctu a l species a re ra rely th e
,

s a me or evencl osely rel a ted For ex a mple the re a re no

.
,

ca cti i n th e S a h a ra D ese rt bu t th e re a re pl a nts cl o sely

resembli n g them which a re no t ne a rly rel a te d O n th e .

o the r h a n d rel a ted species a re usu a ll y f ou nd i nthe s a me

,

ge ne ra l regi ona lth ough they ma y no t resemble one a n

o the r i n a ppe a ra n ce o r live i n simil a r h a bit a ts This .

study o f th e a d a p t a ti o no f org a nisms to th ei r su rrou nd

ings h a s a p ra c tic a l side a ls o i n th a t successful a gri cul
,

tu re c onsists ve ry l a rgely i np ro vidi ng th e m ost sui ta ble

e nv i ronme nt for e a c h p a rticul a r c rop Th us th e U S .
,
. .

D ep a rtme nt of Ag ricultu re expe rts a re o fte na ble to p re

dict wh a t s ort o f c rop s will do well u nde r cultiv a ti on
onwild l a n ds by obse rvi ng th e pl a nts growi ng on th em
i nna tu re . Th is is o f c ou rse o nly a begi nni ng for th e
, , ,

chief obj ec t o f th e scie ntific a g ricultu rist is to devise

still m ore fa v ora ble c onditi ons o f g rowth for h is a nim a ls
an d pl a nts .

1 2 M A N S P LA CE I N N ATU R E i s ce rt a i n ly one o f th e
’
.

subj ects o f g re a test imp ort a nce a nd i nte rest to h imself .

I t is the refore well to u nde rs ta nd in wh a t w a ys h e is

, ,

subj ect to th e s a me limi t a ti o ns a s o th e r a nim a ls a nd i n ,

wh a t w a ys h e di fi ers f rom them a nd is th ei r supe ri or i n

m oldi ng a nd t ra nsf ormi ng h is e nvi ronme nt to suit h is
needs or c onve nie nce I t ma y be dis a pp oi nti ng to find
.

th a t h e h a s fa r m ore i nc omm onwi th th em th a no th e r

wise H oweve r the di ffe re nces sm a ll a s th ey a re m a ke
.
, , ,

him l a rgely th e m a s te r o f th e destiny of h is ra ce a nd a n

imp ort a nt influe nce ona ll na tu re .
 PAR T I

S E C T I ON 1

T HE LI V I N G SUB S TA N CE
 CHAPTER II

P R O T OPLA S M AN D T HE CE LL

U N I VER S A LI T Y P R OTOP LA S M
OF — I t is
a cu ri ous f a ct
.

th a t th ough th e su rf a ce of th e e a rth 1 s e norm ously di

v ersi fi ed the re I S a lm ost n o squ a re f oo t o f its a re a th a t

is with o u t life I t is a lm ost a s st riki ng a f a ct th a t

.

th rough ou t the e norm o us ra nge of livi ng th i ngs the re is

p rese nt a si ngle livi ng subst a nce p ro top la sm wh ich is , ,

fu nd a me nt a lly a like whe reve r f ou nd S ome org a nisms .

c onsist enti rely of p ro t opl a sm s ome highly org a nized

,

beings c onsist only in sm a ll p a rt of livi ng p rot opl a sm

but chiefly of th e p roducts formed by it .

PR OTO P LA S M A LWAY S OR GA N I Z ED I N TO CELL S I t will .

be seeni na subseque nt p a ra g ra ph th a t the physic a l st rue

tu re of p ro t opl a sm is such th a t it c ould not exist u n
supp orted i nl a rge m a ss From th is it f oll ows th a t a ny
.

l a rge org a nism must h a ve s ome meth od of dividi ng th e

p rot opl a sm i nt o sm a lle r p orti ons In f a ct it is f ou nd
.

to be t rue th a t whereve r th e livi ng subst a nce i n

,
c re a ses
l a rgely i n m a ss it is org a nized i nt o disti nct ve ry sm a l l ,

p a rts c a lled cells Th e te rm cell signifies a nindepende nt

.

bi t o f p ro t opl a sm .

T H E D I S COV ER Y OF CELLS — Alth o ugh livi ng org a n

.

isms h a ve beenknownfrom the e a rliest times it is only ,

with i n th e l a st two hu nd red a nd fifty ye a rs th a t th ey

h a ve bee nknow nto c onsist of cells Th e re a s onfor this .

is th a t p revi o us to th a t time th e re we re no me a ns o f
obse rvin g so sm a ll a n obj ect Ordi na ry cells a re too
.

sm a ll to be S ee nwith the na ked eye or eve nwith a sm a ll

le ns They v a ry gre a tly in size but a n a ve ra ge cell i n
.
,

a growi n g roo t tip for i nst a nce is a b ou t one ten

-
, ,

13
 4 P R O T O P LA S M AND THE CE LL

th ous a ndth o f a ni nch i ndi a mete r Inthe h um a nb ody .

the re a re p rob a bly seve ra l h u nd red m illi oncells .

Th e disc o ve ry o f cells the ref ore h a d to a w a it th e p er

, ,

f ecti ono f th e mic ro sc ope In th e S eve n

. tee nth C e ntu ry
a H oll a n de r na med v a n Le uwenh oek h a d so fa r p ro
g re ss ed i n g ri nd in g le n ses th a t it bec a me p o ssible to see
sm a l l obj ec ts These le nses a ro used a gre a t de a l of cu ri
.

osi ty a n d we re e a ge rly s ough t by scie ntific men Am ong .

them wa s R obe rt H ook a n En ,

glish m a n In 1 667 h e .

h a ppe ned to slice a piece o f c ork with h is ra z or a nd ex

a mi n ed i t with a mic rosc op e a nd d isc ove red to h is a st on
,

i sh men t th a t it c on t a i ned a g re a t m a ny ti ny c a vities .

Bec a use o f thei r a ppe a ra nce s ometh ing like the cells
o f a p ris on— h e g a ve th em th e n a me c el l This te rm .

we still u se a l th ough it is pe rfectly evide nt th a t it is a

,

misnome r Th e imp ort a nt p a rt of th e cell is not wh a t

.

R obe rt H ook sa w na mely th e h a rd cell w a ll bu t the

, ,
-
,

livi ng p ro t opl a sm i nsid e th e w a ll wh ich h e did no t see ,

.

T H E D I S COVE R Y O F P R OTO P L A S M — I t wa s m ore th a n .

on e hun d red ye a rs a fte r H ook s d isc ove ry of th e cell b e

’

fore a ny obse rve r sa w a nd publish ed a na cc ou nt of wh a t

we now k no w to be th e livi ng subst a nce Th e fi rst a o .

c o u nt which one ca n rec og nize a s desc ribi ng p rot o pl a sm

wa s w ritte n i n 1 7 55 . A few ye a rs l a te r i n 1 7 73 a n , ,

I t a li a na bb o t na med Corti obse rved i nth e c omm onw a te r

pl a nt na med Cha ra th a t th e c onte nts o f th e cell we re ,

u nde r ce rt a i nc ondi ti ons i n m oti on Alth ough h e p ub

,
.

l i sh ed a na cc o u nt o f h is obse rv a ti oni t seems no t to h a ve

a t tra cted a n y p a rticul a r a tte nti o n u n til 1 80 7 wh e n th is
fa c t wa s a g a i nobse rved i nthe s a me pl a nt by a G e rm a n
i nvestig a t or na me d T rev i ra nu s S till th is obse rv a ti on.

did no t suggest to a ny one th a t th e m ovi ng subst a nce

was a live . A t th a t time th e re we re m a ny th e o ries to
a cc ou n t for th e growth o f org a n isms a l l o f wh ich a ppe a r ,

to us to da y to be f a n
-
ciful bu t it is p rob a bly due to th is
.

fa ct th a t T rev i ra nu s s o bse rv a ti on d id not suggest th e

’

c onnec ti on betwee n m o ti on a nd life I t wa s no t u ntil .

 P H Y S I C AL C HA R A C T E R I S TI CS 15

fifty ye a rs l a te r th rough th e l a b ors of a g re a t m a ny men

, ,

th a t it wa s g ra du a lly re a lized th a t th e cells i nthe livi ng

p a rts o f pl a nts a nd a nim a ls a re a lw a ys filled with a j elly
like subst a nce a nd th a t this is re a lly th e livi ng subs ta nce
, .

In 1 84 4 C a rl N a egeli fi rst e nu nci a ted wh a t we rec og

nize to da y to be th e c orrec t th e ory of g rowth : Tha t
-

e very li vi n g ce l l i s deri ved fro m a no th e r li vi ng cell by

di vi si on Th e full f orce of this st a teme nt ca n only be
.

a pp reci a ted whe n it i s re a lized th a t the f a n ciful the ories

i n f a v or a t th a t t i me supp osed th a t cells migh t be p ro
du ced i ns omethi ng like th e s a me wa y th a t the p ores i n
a l oa f o f b re a d a re f or m
ed by th e exp a ndi ng g a ses Th us .

it is see n th a t th is wa s a ve ry imp ort a nt disc ove ry I t .

wa s not l ong u ntil it wa s rec ognized a s a c orrect st a te

ment of f a c t b o th f or pl a nts a nd a nim a ls A t a b out th e
, .

s a me time i t beg a n to be re a lized th a t the c onte nts of

the cell a re o f imp ort a nce a nd i n 1 84 6 Hug o v onM ohl
,

rec og n ized this f a ct by givi ng th e na me p ro top la sm to

th e slimy cell c onte nts Withi n th e next twe nty ye a rs
.

it bec a me a n est a blish ed f a c t th a t th e imp ort a nt p a r t

o f the cell is the p ro t opl a sm a n d th a t it i s the se a t o f a l l
life a ctivities .

P H Y S I CA L CH AR A CTER I S TI CS O F P R OTO P LA S M It .

migh t be supp o sed th a t a subst a nce so imp ort a nt to a ll

livi ng thi ngs a s p ro t opl a sm w ould be disti nguish ed by
e a sily rec ogniz a bl e a nd st riki ng ch a ra cte ristics Thi s .
,

h oweve r is not t rue P ro t opl a sm f rom di fferent s o u rces

,
.

a ppe a s ve y much th e s a me
r r —
I t is a j elly lik e subst a nce
.
,

th a t is to sa y it is neith e r a s olid nor a liquid bei ng i n

, ,

c onsiste ncy s omewh a t l ike th e wh ite of a n egg I t is .

eithe r c ol orless or S ligh tly yell ow i n a ppe a ra nce I t is .

not sufficiently fluid to ru nl ike w a te r nor is i t sufficie ntly ,

rigid to st a n d up like a s olid j elly I ts own weigh t is

‘

sufficie nt to c a use i t to sp re a d ou t on th e su rf a ce of a
gl a ss S lide I t ca nbe pulled to One side like rubbe r a nd
.

will r et ra ct to its origi na l f orm wh e n rele a sed S h owi ng

n
,

th a t it h a s a ce rt a i ndegree of el a sticity S ome ki ds o f .

16 P R O T OP LA S M AN D THE CELL

p ro to pl a s m a re cle a r a nd t ra nsp a re nt a nd h om oge ne ous

i n s truc tu re wh ile o the r kinds a ppe a r to h a ve a f oa my
,

s truc tu re like s oa psuds P ro t opl a sm ma y s ometimes

,
.

a ppe a r op a que bec a use i t c on t a ins c onside ra ble qu a ntities

o f f oo d .

T H E C H EM I S TR Y O F P R OT O P LA S M P rot opl a sm i s n ot .

a simple chemic a l c o mp o u n d nor i ndeed is i t simply a

,

mix tu re o f chemic a l c omp o u nd s ; i t i s i n re a lity a n , ,

o rga n i z ed st ructu re c omp o sed o f m a n

,
y d i ff e re nt com
p ou nds Th e l a rge r p a r t o f its bulk c onsists o f w a te r
.
,

two th i rds or m o re i nf a c t
-
The s olid p orti ono f it con
,
.

sists chiefly o f th e ch emic a l subst a nce k now na s p ro tei n ,

wi th s ome f a ts s ome sug a r a nd v a ri ou s m i ne ra l s al ts

, ,
.

In a d diti on to these subst a nces wh ich p resum a bly con

s ti tu te a p a r t of the livi n g m a ch i ne th e re a re u su a lly ,

p rese nt f ood s or o the r m a te ri a ls f ormed by th e p roto

pl a s m .

The a ctu a l numbe r o f elements which e nte r i nt o th e

st ru ctu re o f p rot opl a sm i s sm a ll Ca rb on hyd ro ge n .
, ,

o xy ge n a n d ni troge n a re a lw a ys p rese nt i n th e p ro teins

,

an d u su a lly i n a dditi on sulphu r or ph o sph orus or b o th ,

.

The fa ts a nd sug a rs c onta i nc a rb on hyd roge n a nd oxy , ,

g e n ; the w a te r c o nt a ins hyd roge n a nd oxyge n; th e Va

ri o us s a l ts p rese n t c onta i n c a lcium m a g nesium p o ta s , ,

sium sulphu r ph osph o rus a nd i ron I t is d ifficul t to

, , , .

decide a s to wh ich o f the eleme nts e nume ra ted a re a b a o

l u tely i ndi spe ns a ble to th e livi ng subst a nce I t w ill be .

sh ow n i n a subseque nt ch a pte r th a t living o rga nis ms

c a nno t m a i nt a inlife wi th ou t a ll of th e eleme nts we h a ve
na med bu t i t d o es not foll ow f rom th is th a t e a ch a nd
,

a ll c on stitute a n a ctu a l p a rt o f the livi ng m a chi ne I t .

seems ce rt a i nth a t no thi ng ca nbe a l ive with o u t p rotei n ,

w a te r a nd ce rt a i ns orts of foods I t is a ls o ce rt a i nth a t

, .

i f th e livi ng p rot opl a s m be dep rived of th e p rese nce of

s ome o f th e m i ne ra l s a l ts me nti oned it w ill p rese ntly
ce a se to exhibi t th e livi ng fu ncti ons This m igh t me a n
,
.

th a t they a re a ll a p a rt o f it ; bu t m ore likely it me a ns

 P H Y S I O L O GI C AL C HA R A C T E R I S TI C S 17

th a t s ome of them a t le a st me rely fa cilit a te the w ork of

th e living m a ch ine by thei r p rese nce In a c rude wa y .
,

on e migh t supp ose th a t s o me o f these s a lts e n a ble the

livi ng m a chi ne to fu nc ti on i n s omethi ng th e s a me wa y

th a t oi l e na bles a n a ut om obile e ngi ne to fu ncti on In .

the l a tte r c a se the oi l d oes not c onstitu te a p a rt o f the

,

mech a nism bu t th e engi ne W i ll no t runl ong with o ut it .

T H E P H Y S I OL OGI CA L C H AR A CTER I S TI CS OF P R OT OP LA S M .

I t h a s a l re a dy bee n me nti oned th a t wh e reve r p ro t o

pl a sm is fou nd it a lw a ys exhibits ce rt a in ch a ra cte ristics .

Of these we ma y enume ra te six

.1 I t h a s th e p o we r o f g rowth w a ste a n ,
d rep a ir
,
.

.2 I t h a s th e p o we r to u tilize f oods a n d to libe ra te

ene rgy from them to do w ork .

.3 Th e e ne rgy th us libe ra ted ma y be used to exe cute

forcible m oveme nts .

.4 Th e e ne rgy libe ra ted ma y b e used i nc a rryi n g ou t

ch emic a l re a cti ons i n th e m a nuf a c tu re of o th er sub
st a nces.

.5 I t h a s th e p owe r o f resp onse to externa l stimuli .

.6 It h a s th e p owe r o f a u t om a tic divisi on a n d re

p roducti on .

G RO WT H W A S TE AN D R E PA I R
, ,
Whe reve r living p ro
.

tOp l a sm is f o u nd i t is a lw a ys i n a st a te o f ch a nge The .

p rot opl a sm o f th e growing p a rts of pl a nts a nd a ni ma l s

is a ble to u tilize f oo ds to c onst ruct new p rot opl a sm .

P rot opl a sm wh e reve r it exists a lw a ys w a stes a w a y u nder

its own a ctivities All p ro t opl a sm h a s th e p owe r to
.

utilize f oods to rep a i r this w a st a ge Th ese t opics need

.

little m ore th a nme nti on a t th e p resent time i na smuch ,

a s th ey f orm th e subj ect m a tte r o f a subseque n t ch a pte r .

TH E P O W ER T O LI BER ATE E N ER GY F R O M F OOD F ood .

is used by p rot opl a sm not only for g rowth a nd rep a i r bu t

a ls o a s a s ou rce o f e n e rgy I t will be Sh ow n l a ter th a t
.

I nth e f orm a ti ono f f o od the e n e rgy of su nligh t is l ocked

up i nth e chemic a l c omp ou nds of sug a r a nd othe r S imil a r
subst a nces Th ese whe n used a s food by p rot opl a sm
.
, ,
18 P R O T O PLA S M AN D T HE CE LL

ca nbe dec omp osed a nd the ene rgy libe ra ted This e ne rgy .

ma y be empl oyed to ch a nge the sh a pe a nd form of th e

p ro t opl as m forcibly The p recise mech a nism wh ich i s
.

empl oyed to tu rn the chemic a l e ne rgy o f f ood int o me

ch a n i ca l m oti onby th e l i v ing subst a nce is no t well u n de r
st ood a nd is a t best a difficult one to study .

T H E P OW ER O F CAU S I N G C H E M I CA L C H A N GE — The
e ne rgy libe ra ted f rom f oods ma y be empl oyed to c a rry
o u t o th e r chemic a l ch a n ges Th e seque nce of eve nts is .

s omewh a t a s foll ows : Ene rgy is st ored up in f ood sub

st a nces only by th e g ree ncells o f pl a nts ; but once th is
e ne rgy is st ored i nfood s th ey ma y u nde rg o fu rth e r chemi
ca l ch a n ges Th ese ch a nges ma y be o f two s orts In
. .

th e fi rst pl a ce they ma y be o f th e s ort wh ich libe ra tes

,

e ne rgy This ty pe of ch a nge is of c ou rse necess a ry whe n

.

eve r e ne rgy is to be u sed for f orcible m o veme nts On .

th e o the r h a nd the re a re ch a nges i nwh ich th e resulti ng

,

c omp ou nd c ont a i ns m ore e ne rgy th a n th e one ou t of

which it wa s m a d e N ow no ch a nge o f this s ort ca nbe
.
,

b rough t a b ou t with out th e a pplic a ti on o f e ne rgy f rom

th e outside This e ne rgy must th e nbe de rived f rom th e
.

d ec omp ositi ono f food s It will not be p ossible i na b ook

.

o f th is limited sc ope to discuss i nd et a il th e g re a t v a ri ety

,

o f chemic a l ch a n ges b rough t a b out i nlivi ng p rot opl a sm .

I t will be sufficie nt pe rh a ps me rely to c a ll a tte nti on to

, ,

the f a ct th a t org a nic subst a nces p roduced i n th is wa y

v a stly o ut numbe r th e i norg a nic ones S uch v a ri ous sub .

st a nces a s w ood the textiles d rugs rubbe r gums a nd

, , , , ,

th ous a nd s o f o th e r a rticles o f c omme rce a ll h a ve th ei r ,

o rig i ni nch emic a l ch a n ges which occu r i nlivi ng m a tte r .

S E N S IT I VE N ES S T o ST I M U LI — P rot opl a sm is no t on ly .

c a p a ble o f executi ng ch emic a l re a cti ons a nd f o rcible

mo veme nts but it is a ls o ext remely se nsitive to ou tside
i nflue nces The ch a nges i nth e e nvi ronme nt to which it
.

resp on ds a re c a lled s ti mu li Th ey ma y be of a v a riety

.

o f s o rts . M e re c o nt a c t the p rese nce o f ch emic a l sub

,

s ta nces o f a p a rticul a r ch a ra c te r elect ric cu rre nts ligh t , , ,

 S TI M ULATI O N , C O ND U C TI O N , R E S P ON S E 19

or o th e r f orms o f ene rgy a re a l l c a p a ble o f p roduci ng

ch a nges i np ro t opl a s m I ts p owe r to resp ond to stimuli
.

i s depe nde nt up on its physic a l a nd chemic a l st ructu re .

N o t only d oes it resp ond a t the p o i nt o f c ont a ct bu t the ,

stimulus ma y p rod uce the resp onse a t a dist a nt p oi nt .

S TI M U LATI O N C O N D U CTI O N AN D R E S P O N S E — I t will

, ,

be obse rved th a t th ese a re re a lly th ree sep a ra te a nd di s

ti nct p rocesses Th e fi rst is th e se nsitive ness of the p a rt
.

o f th e p ro t opl a sm which fi rst r eceives th e stimulus the

;
sec ond p a rt is th e c onducti onof th e stimulus th rough the
p rot opl a sm th a t is to sa y th e p rop a g a ti onof s ome s ort
, ,

of ch a n ge ; a nd thi rd th e re is the resp onse i ns ome o the r

,

p a rt of the p rot opl a sm Th e resp onse d iffe rs from se nsi

.

ti v en ess a n d c onducti on ordi na rily i n th a t it is a ccom

, ,

pa n i ed by much g re a te r e ne rgy ch a nges If th e resp onse .

is to be one of m ovement it is obvi o us th a t e ne rgy must

be libe ra ted by th e dec o mp ositi on o f s ome e ne rgy
c ont a i ni ng subst a nce i norder to a cc omplish it We h a ve .

a nexh ibiti ono f this inou r ownb odies i na ve ry Spe ci a l

i z ed f orm wh e re th e resul t is b ro ught a b o ut no t withi n

, ,

a Si n gle cell ; but by th e coOp era ti on of m a ny cells I f .

on e t ouches his fin ge r to a h o t st ove th e fi nge r will be

a ut om a tic a lly withd ra w n In orde r th a t th is ma y
.

h a ppen a l l o f th e th ree thi ngs wh ich we h a ve noted

,

i nthe si ngle cell must o ccu r i n th e hum a nb ody Th a t .

is to sa y the re mu st fi rst be se nsitive ness to this h ea t

,

e ne rgy ; the n s ome s or t of stimulus must be c onducted

a l on g the -ne rve t ru nk to th e ne rve cente r a nd b a ck a g a in
to th e muscle ; a n d fina lly a libe ra ti on o f e ne rgy must
,

be b rough t a b o u t wh ich will e na ble th e muscle to con

t ra ct a nd d ra w th e fi ng e r a w a y fro m th e h ot st ove
CELL D I V I S I O N AN D R EP R OD U CT I O N Wh e ncells g row .

they tend to re a ch a m a ximum size which is dete rmi ned

by th e ki nd of pl a nt or a nim a l i n wh ich th ey o ccu r .

Wh en th ey h a ve re a ched this S ize divisi on a u t om a tic a lly

occu r s. Th is is a s omewh a t c omplic a ted p rocess a nd
will be desc ribed i n a nothe r pl a ce I t is inte nded h e re.
20 P R O T O P LA S M A ND T HE C E LL

Simply to c a ll a tte nti on to th e f a c t th a t this p rocess is

necess a ry i n o rde r to keep th e cells o f th eir p rope r size
to m a i n t a i nthe st ructu re o f the org a nism a nd th a t it is ,

a n a u tom a tic self regul a ted p rocess wh ich is e nt irely

,
-

ch a ra c te ristic of livi ng m a tte r .

T H E S TRU CTU R E OF A TY P I CA L C E LL I t is not p os .

sible to ch oose a ny S i ngle cell which w ill S h ow a ll the

ch a ra cte ristics of c ells In orde r to exhibi t th e m a i n
.

fe a tu res o f S imple cells i t will be necess a ry to ch oose two

or m ore ill ust ra ti on s .

If a cell be ch ose nf ro m th e g row ing p orti onof a y ou ng

root 1 ) or stem of a pl a n t it will be f ou nd to con ,

FI G 1
. A Typi ca l Pl a nt C e ll f ro m th e ro o t ti p of th e S piderw ort
.
,

(T ra desca ntia vi r gi ni ca ) c w ce ll w a ll ; cyl cy to pl a sm ; n nu cl e us ;

. . .
, , ,

chr chr o m a ti n
, ; n s n u c.l e.a r,sa p ; n nu c l eo lus ; v v a cu o le fill ed wi th
’
, ,

w a ter a nd di ss olv ed subs ta nces .

t a i nor to c onsist of the foll owi ng p a rts : a ro u nd the ou t

side the re is a delic a te w a ll c omp osed of a nonlivi ng -

subst a nce cellul o se ; with inth is w a ll th e re is a s olid m a ss

,

o f p ro t opl a sm I n th e ce nte r o f th e p ro t opl a sm the re

.

is a rou nd ed b ody m ore or less sphe ric a l c a lled th e

, ,

nucleus P ra c tic a lly a ll cells c ont a i n p rot opl a s m whi ch

.

is diffe re nti a ted i nt o th ese two p a rts for c onve nie nce o f ,

refe re n ce c a lled nu c leu s a nd cy top la sm I n a dd iti on to .

th ese two st ructu res m a ny cells p ossess othe r org a nized

,
 S T R U C TU R E OF A T Y P I C AL CE LL 21

p a rts I f we ex a mi ne a cell f rom the olde r p a rt o f a

.

pl a nt we S h a ll fi nd th a t the w al l is thicke r th a t the cell ,

is l a rge r a nd th a t the p rot opl a sm is c onfi ned to a thi n

,

l a ye r f ormi ng a cl osed sa c i nc ont a c t with the wa ll The .

middl e of the cel l is filled by a wa ter va cu ole c ont a i ni ng

a v a riety of subst a n ces i ns oluti on Th e fu ncti onof this .

v a cu ole is a n i nte resti ng one I t will be sh ow n l a te r .

th a t the m a te ri a ls which a re diss olved inth e w a te r exe rt

a p ressu re up onth e p ro t opl a smic sa c to keep it i n cl ose

c ont a ct with the su rrou nd i ng cell w a ll This is the chief -

.

me a ns of m a int a i ni ng th e rigidity o f th e s ofte r p a rts of

pl a nts A nim a l cells on th e c ont ra ry do not usu a lly
.
, ,

FI G 2 . Am oeb a a one celle d a ni m a l n nu cl eus ; cyt cyt opl a sm ;

.
,
-
.
, ,

f f oo d v a cu o l e ; c v contra ct il e or e x cretory v a cu o l e ; g gra nul a r

,
. .
, ,

m ore fluid i nner p a rt of cy topl a m ; e o u ter s olider l a y er of th e

s
,

cy t o pl a sm th e ecto pl a s t ; p ps eudo p o dium or p ro j ecti on of th e

, ,

protopla sm .

h a ve fi rm cell w a lls
-
,
a nd do not h a ve v a cu oles of th is
type a t al l .

A S i ngle celled a nim a l will illust ra te o the r cell st rue

- -

tu res The one c a lled Amoeba Sh ow ninth e a cc omp a ny

.
,

in g illust ra ti on (Fig is a ve ry simple a nim al Its .

b ody c onsists of nucleus a nd cyt opl a sm The re is no .

w a ll a b out it nor d oes it exh ibit a ny o the r defi nite cell

,

org a n s N eve rth eless if it is w a tched it will be f ou nd

.
, ,

th a t it is a ble to ch a nge its o utli ne th a t is to sa y it , ,

22 P R O T O P LA S M AN D T HE CE LL

exhibit s m oveme nt I t ca n p rot rude on one side a n

.

exte nsi on of i ts b ody a nd e nl a rge th a t so th a t a ll the

p rot opl a sm fl ows i nt o i t ret ra cti ng the opp osite side
,

me a nwh i le By so d oi ng it h a s ch anged its p ositi on

.
, .

This typ e of m oveme nt is k now na s a mceboi d mo vement ,

an d is p rob a bly c omm on to ne a rly a ll p rot opl a sm .

Th e one celled g ree n pl a nt Ch la mydo mona s (Fig 3 )

-
.

exhibits a s omewh a t highe r deg ree o f org a niz a ti on ,

i n th a t it h a s a w a ll su rrou ndi ng it wh ich p reve nts

FI G 3 . . ne cell ed pl a nt n nu cl e us wi th i ts
C hl a myd o m o na s , a o -
.
,

chr o m a tin n e tw o rk nu cl ea r sa p a nd nu cl e o lus n; chr the green

, , ,
’
,

chro m a to ph o re o f s o li d e r p ro to pl as m co n ta i ni ng the green col o ri ng

m a t te r chl o rophyll ; cyl cy to pl a sm ; p y the py renoid o r s ta rch
, , ,

f orming o rga no f the chrom a to ph ore ; s s ta rch gra i ns ; e ey e sp o t

, ,
-
,

a so li d e r m a ss o f p ro to p la s m co n ta inin g a reddish b row n pigm ent ,

supposed ly a n orga n se nsi tive to li gh t ; c v co ntra ctil e v a cu o l es ; . .

,

c w . cell w a ll ;
.
, c fl a g ell a h a i rlik e p ro j ecti o ns of the cy to pl a sm by
, ,

m ea ns o f whi ch th e pl a nt p rope ls i ts elf th rough th e w a te r .

th e o rg a nism f rom ch a ngi ng its sh a pe a s a wh ole .

An d i t h a s a ls o a t one end two p ro j ecti ng oa r like ,

-

p rocesses which ca n be m oved ra pidly th e reby p ro ,

du ci ng m oveme nt o f th e e nti re cell I t h a s a ls o a .

gree n c ol or Th is g ree n c ol or will be f ou nd to be

.
 S UMMA R Y 23

limited to ce rt a i n p a rticul a r p a rt of the livi ng sub

a

st a nce and the p rese nce o f this g ree n pigme nt chl o

rop h y l l is tha chi efldm
, ,

,
ti ncti o nbetweenthe pl a nt a nd the
an i ma l kingdoms for i t is by th e a cti ono f this pigme nt
,

on ligh t th a t p ro t opl a sm is e n a bled to b ri n g a b ou t th e

m a nufa ctu re of sug a r f rom c a rb ondi oxid a nd w a te r I II -
.

the cell o f Chl a myd om ona s we h a ve th us two a dditi ona l

org a n s not f o u nd in Amoeb a org a ns of l oc om o ti on a nd
,

org a n s of f ood m a nu fa ctu re This s a me pl a nt exh ibits

.

a l s o a S peci a l org a n f or the recepti on o f stimuli N ea r .

the end o f th e cell to wh ich th e l o c o m o t or org a ns a re

.

a tt a ched is a ti n y sp o t of reddish b row npigme nt I t h a s

-
.

bee n f o u nd th a t org a nisms With a p i gme nt sp o t o f this

s ort a re m ore pe rfectly a d a pted to re a ct to ligh t th a n
th ese with o ut it .

S U M M AR Y — . P ro t opl a sm is th e u nive rs a l livi ng sub

st a nce I t is a lw a ys org a nized i nt o cells or u nits These
. .

exhibit a rel a tively u nif orm physic a l a nd ch emic a l com

p ositi ona nd a re a ble to c a rry on a de finite a nd ch a ra c
teri sti c set of physi ol o gic a l fu ncti ons
,
A typic a l cell .

m n n
/

p os sesse s ore o r less d i ffere t i a te d o rga s a d a p ted t o

pe rform these fu ncti ons E ve ry cell h a s a nucleus a nd

.

cyt opl a sm I t ma y or ma y not h a ve a w a ll of cellul ose

.

or o the r n onlivi n -
g m a te ri a l su rrou ndi ng i t I t ma y .

h a ve a w a te r v a cu ol e i n the ce nte r I ts p ro t opl a sm is .

usu a lly a ble to ex ecute m oveme nts eith e r by a ch a nge ,

i nth e f orm of the cell a s a wh ole or by th e i nstru men

t a lity of speci a l org a ns of l oc om o ti on All cells a re se nsi .

tive to exte rna l stimuli a nd a ble to resp ond to them i na

v a riety o f w a ys a s for ex a mple m oveme nt Pl a nt cells
, ,
.
,

i ndisti ncti onfrom o th e r cells ma y p ossess speci a l org a n

,
s
c ont a i ni ng chl orophyll for th e m a nuf a ctu re of sug a r .

C ells a re a ble to u nde rg o a ut om a tic divisi on the reby ,

b ri nging a b ou t fu rthe r growth or rep roducti on .

 C HAPT ER III

M ULTI C E LL ULA R OR G AN I S M S AND DI VI S I O N

OF LA B OR

N ATU RE O F T H E S U B J ECT — S ince p ro t opl a sm is the

on ly livi ng subst a nce it f oll ows f ro m wh a t we h a ve
,

le a rned i nthe f oreg o ing ch a p te r th a t livi ng i na bi ol ogi ,

ca l se n se c onsists of se ns a ti on m o veme nt w a ste rep a i r

, , , , ,

g row th a nd th e multiplic a ti on o f i ndivid u a ls At th e

,
.

s a me time i t i s equ a lly t rue th a t we rec og nize ce rt a indif

ferences a m ong livi ng thi ngs th a t le a d u s to sep a ra te
them int o two g re a t g roups a nim a ls a nd pl a nts We a ls o
,
.

dis ti nguish ce rt a i n peculi a rities th a t e na ble u s to sub

divide th e membe rs of these g roups i nt o m a ny diff e re nt
species or kind s o f l i vi ng thi ngs We k now th a t for s ome .

re a s o nth e squi rrel is diff e re nt f ro m th e oa k t ree up wh ich

it sc a mpe rs a nd the bee is u nlike th e cl ove r f ro m wh ich
,

i t g a th e rs h oney .

S ince these org a nisms a nd h u nd red s o f th o us a nds o f

o the r species a re livi n g a nd he nce a re ca rryi ng on th e
,

s a me vi t a l p rocesses i t is well to a sk o u rselves th e ques

,

ti on on wh a t do th ese diff e re nces rest ? Th e a nswe r is

, ,

o n st ructu re Al l o f th e d iff e re nt ki nds of livi ng th i ngs

.

a re vi ta l e n gi nes c a rryi ng onp rocesses wh o se sum t ot a l

,

we c a ll li fe So me a re fitted to w ork i n w a te r o the rs

.
,

o n the su rf a ce o f th e e a rth or on th e g ro u n d wh ile still ,

o the rs spe n d a p orti ono f thei r existe nce i nth e a i r Th e .

pl a nt m a ch i nes a re usu a lly st a ti ona ry wh ile th e g re a t ,

m a j ority of a nim a ls m ove a b o ut Fu rth e rm ore m a ny of .

,

th ese m a chi nes a re c omp a ra tively simpl e th e w ork th ey ,

do is ra th e r c rud e a nd we spe a k o f th em a s l ow fo rms o f

,

li fe Onthe o the r h a nd th e oa k t ree or th e squi rrel for

.
, ,

24
 A M (E B A 25

ex a m p le a re ve ry c omple x th ei r fu ncti ons or a ctivities

, ,

a re m ore e ffi cie n tly pe rf ormed a nd they a re desig na ted

high org a nisms This f a ct i nv olves th e p ri nciple o f the
.

divisi ono f l a b or th a t will be bette r u nde rst oo d a fte r we

h a ve ex a mi ned a few ex a mples illust ra ting S imple m ore ,

c omplex a nd h ighly c omplic a ted types .

A M (E B A Am ong th e best k now nex a mples of a p rimi

.

tive org a nismis th e a m oeb a (Fig livi ng i n the mud .

an d slime a t the b o tt om o f p onds a n d ge ntly r u n ni ng

st re a ms L a rge i ndividu a ls a re b a rely visible to th e u m
.

a ided eye , bu t u n de r the mic rosc ope th e b od y is see n to

FI G . 4 .
—T h e Am oeb a ,
S h owi ng pro cess of m ul tipli ca ti on .

c onsist o f d s mi flu i d p rot opl a sm with o ut a nen

a rop o f e -

cl osing w a ll but su rrou nding a gl obul a r nucleus This .

an im a l is th us a S i ngle cell Inm ovi ng a b out one p a rt

.

of the b ody is sl o wly pushed ou t i n the fo rm of a blu n t

p rocess wh ile a c orresp ondi ng p a rt o f th e cell is with
,

d ra w n th e a nim a l thus fl owi ng a l ong like a S heet of

,

quicksilve r m ovi ng a c ross a n u npl a ned ge ntly sl opi ng ,

b oa rd Whe re a p a rticle of f ood lies in its p a th the

.

a m oeb a pushes it i n t o its s o ft i nte ri or whe re it is digested

with out the a i d of a ny st ructu res c omp a ra ble to a m outh
or digestive system B re a th i ng t a kes pl a ce th rough the
.

genera l su rfa ce o f th e b ody Cold he a t a nd v a ri ous .

, ,
26 M ULTI CE LLUL A R O R GANI S M S

chemi c a l a ge nts p rod uce ch a nges inth e ra te of l ocomo

ti onor a lte r its d irec ti on; he nce th e a m oeb a feels The re .

is a ls o a defi nite sp ot whe re th e w a te r a ccumul a tes u ntil

it forms a d roplet a b out th e size of the nucleus whe re ,

up on i t is emptied to the o utside ; I t is ge ne ra lly be

l i eved th a t w a ste p rod ucts e nte r this contra ctil e va cu ole ,

wh ich th us a cts a s a S imple kidn ey or exc ret ing org a n .

FI G . 5 Pa ra mmci u
.
m
o r S li pper a n im a l c l e cv,
u . co ntra ctil e
v a c u ol e or simple ki d ney ; m, m ou th a nd g ll e t ; n, nu cl e
u us B two
.
,

i ndi vidu a ls conjuga ting, a type of fertiliz a ti on C, indi vid

. u a l di vi ding
s xu a lly
a e .

Whe nth e b ody re a ches a ce rt a i nsize it divides i nt o two

equ a l h a lves e a ch of which th e nceforth bec omes a ni n
—
,

n n
depe de t o g a ism
r n a simple typ e of multiplic a ti on

o f individu a ls .

P AR AM CECI U M In th e p a ra m oecium or slippe r a n

.
-
i
ma l cul e (Fig livi ng i n th e s a me situ a ti ons a s the
.
 —
O N E C E LL E D O R GANI S M S 27

a m oeb a the b ody is likewise o f mi crOSCOp i c p rop orti ons

, ,

but ret a i ns a defi ni te sh a pe owi ng to a perma nent cell .

memb ra ne Th e b ody i s th i ckl y c o ve red by a c oa t o f

.
'

delic a te p ro t opl a smic h a i rs or cili a wh ose rhythmic a l

be a ting d rives the org a nism f rom pl a ce to pl a ce At one .

p o int th e cel l w a ll is p e rf ora ted a nd th rough th is m ou th

-
,

Ope n ing food p a rticles a re d rive nby speci a l cili a into the
inte ri or In o the r respects th e life p rocesses resemble
.

th ose of th e a m oeb a I t is to be noted , h oweve r th a t th e

.
,

p a ra m oecium p ossesses defi nite org a ns of l oc om otiona nd

a m o uth ; a n d a s will a ppe a r m ore cle a rly i na succeed
!

i ng p a ra g ra ph this c onstitutes a re a l a dv a nce bey ond the

,

a m oeb a .

I P RTA N CE OF ON E CE LLED OR GA N I S M S
M O - — Wh ile .

th ese two ex a mples a re i nsufficie nt to illust ra te th e b e

wilde ring dive rsity of st ructu re of th e th ous a nds of na med
Species of th ese on e celled a n im a ls a nd pl a nts th ey ma y
-
,

be used to emph a size th e f a ct th a t a ll le a d successful

lives M i nist eri ng to th ei r needs a v oiding e nemies a nd
.
, ,

le a vi ng offsp ri ng to t a ke th eir pl a ce th ey exist a nd h a ve ,

p e rsisted f or th ous a nds of ye a rs a nd th is c onstitutes a ,

successful life Als o i n p oi nt of numbe rs of i ndividu a ls

.

th ey f a r ou tra nk al l othe r livi ng th ings In m a ny re .

gi ons ce rt a i n a qu a tic f orm s settl e to th e b ott om a nd ,

thei r S kelet ons of lime or fli nt f orm dep o sits of gre a t ex

te nt a nd th ick ness Ch a lk t rip oli e a rth a nd oth e r simi
.
, ,

l a r subst a nces a re a ncient a ccumul a ti ons th a t a re o f high

ec onomic imp ort a nce Th e Oi l e ncl osed in th e bo dies
.

o f the mi n ute a qu a tic di a t om pl a nts h a v e f ormed it is ,

believed th e g re a t b eds of na tu ra l oil i nth e weste rnp a rt

,

o f N orth Ame ric a Fi na lly th e re a re m a ny species th a t

.
,

de rive th ei r f ood supply f rom oth e r orga nisms a nd a s ,

p a ra sites p roduce dise a ses th a t ra nk a m ong th e m ost

s e ri o us sc o u rges k n ow n to ma n .

COLO N I A L F OR M S Am ong th e gre a te r numbe r of one

.

celled org a nisms th e divisi on of th e b ody results i n two

off sp ri n g th a t sep a ra te a nd th e ncef orth le a d i ndepe nde nt
28 M ULTI C E LL U LA R OR GANI S M S

lives Inth e high e r typ es o f S imple a nim a l s a nd pl a nts

.
,

h oweve r the divisi onp roduc ts rem a ina tt a ched a nd form

, ,

c ol onies inthe sh a pe of th re a ds sheets or s olids o f v a ri , ,

o u s f orm depe n di ng up on th e di recti on o f th e cle a v a ge

,

an d the subseque nt a rra n geme nt of th e cells Am ong .

th e S implest of these c ol oni a l s p ecies is G onium (Fig .

o ccu rr in g c omm only i n f resh w a te r H e re the si ngle .

p a re nt a l cell divides the two p roducts rem a i n a tt a ch ed

, ,

re divide
-
an d i n this f a sh i on re a ch a six tee ncell st a ge
,
-

with th e cells f orming a pl a te E ve ry membe r of th e .

c ol ony bea ring l oc om ot or whips or fl a gell a th a t ope ra te

, , ,

FI G 6
. .
—C oloni a l O rg a nisms . A, V olv o x ; B , C, to p a nd sid e
views of Go nium .

like cili a is p recisely li ke i ts fell ow i nf orm a nd fu ncti on

, .

I t is imp orta nt to note th a t th e c ol ony ma y be b rokenup

an d e a ch cell c o nti nue to exist A s a m a tte r o f fa ct th e
.

cells do bec ome diss o ci a ted a t th e tim e of rep rod ucti on ,

e a ch f ormi ng a d a ugh te r c ol o ny .

VoLvox In V o lv ox (Fig
. a h ighe r type o f .

c ol o i a l o rg a nism th e b ody c omp rises m a ny cells

n ,
— in
s ome spe cies th o u sa nds i nnumbe r imbedded i na j elly
like subs ta nce a nd a rra nged i n th e f orm o f a h oll ow
sph e re Ea ch cell is egg sh a ped a nd th e gre a te r numbe r
.
-

c a rry fl a gell a wh o se c ombined a cti onrolls th e a ss oci a ti on

 H YD R A 29

th rough the b odies of fresh w a te r Wh e nth e c ol ony h a s .

re a ched its full size a di ffe re n ce a m ong the cells bec omes
a pp a re n t A few i nc re a se i n v olume to a m a rked de
.

gree ow
,
i ng to f oo d m a te ri a l th a t they st ore up a nd ,

th ei r futu re devel opme nt occu rs a l ong one o f two li nes .

Some divi de repe a tedly a nd si nki ng i nt o th e i nte ri or

,

form d a ughte r c ol onies th a t esc a p e with the ruptu re of

the p a re nt w a ll U nde r o th e r c onditi ons ce rt a in cells
.

divide repe a tedly to f orm bu ndles of sle nde r fla gel l a te

cells c a lled spe rms a ny one of which ma y u nite with a
,

p a rticul a r s ort of l a rge cell or egg which the ne nte rs up on

essenti a l ly th e s a me c ou rse of devel opme nt a s the one
just desc ribed .

It is evide nt th a t Volv ox lies a step a h e a d of Gonium

i n reg a rd to c omple xity I n th e f orme r the re a re
.

the begi nni ngs of a divisi on o f l a b or a m ong th e co m

p one nt cells b a sed up on st ructu ra l di ffe re nces C e rt a in .

cells t a ki ng no p a rt i n l o c om o ti on se rve a s rep roductive

eleme nts while th e gre a t m a j ority fu ncti on in l ocomo
,

ti on se ns a ti on a nd feeding bu t a re wh olly u na ble to

, , ,

rep roduce . Th e imp ort a nce of this fa c t will a ppe a r with

g re a te r cle a rness a fte r we h a ve ex a mi ned a nothe r type
of still g re a te r c omplexity .

HYDRA .
— Th e l a st ex a mple th a t we sh a ll ex a m i ne In
its e nti rety is Hyd ra (Fig a f resh w a te r a nim a l re
.

l a ted to th e sea a nem ones c ora ls a nd j elly fi sh es of the

, ,
-

o ce a n . Th e b ody b row nor g reeni nc ol or p rese nts th e

, ,

f orm of a S le nde r h oll ow cyli nde r a b ou t one fou rth o f -

a ni n ch i nle ngth One end of th e b ody is a tt a ch ed by

n
;

a sticky subst a ce to s o me subme rged le a f or st on e while ,

th e Othe r be a rs th e m outh su rrou nded by five or S ix

S le n de r o utgrowth s te rmed te nt a cles Th e b ody w a ll .
,

th rough ou t its e nti re exte nt is c omp o sed of two l a ye rs ,

o f cl osely grouped cells i n wh ich v a ri o us types ca n be

disti nguished In th e o ute r l a ye r the re a re cells th a t

.

se rve for p rotecti on se ns a ti on a nd m oti on while speci a l

, , ,

nettle or st inging cells p a ra lyze the sm a ll org a nisms

30 M UL TI C E LLULA R O R GAN I S M S

th a t se rve a s f ood This nu t ritive m a te ri a l is con

.

v ey ed by a m oveme n t of the te nt a cles int o the h oll ow

in te ri or o f the b ody a nd is th e re d iss olved by th e a cti on
,

o f a sec reti on f ro m th e cells o f the i n ne r l a ye r th a t a re

th us diges tive eleme nts R ep rod ucti on is e ffected by.

eggs a nd spe rm s f ormed f rom ce rt a i noute r l a ye r ce lls or ,

by h oll ow outg rowth s o f th e b ody w a ll th a t devel op ing ,

a m o uth a n d te nt a cles fina lly pi nch off f rom the p a re nt

,

an d bec ome i n depe nde nt org a nisms .

FI G 7
. . Freshw a te r Hydra , nti re a nd l ongi tudi na lly S pli t
e

vi d u a ls . e, e gg ; m m o u th s ur ro u nd e d by te nta cl es ; s spe rm s
, , ,
co n
s idera b ly m a gnifi ed .

D I V I S IO NL A BO R OFI t is p rob a ble th a t th e n

. umbe r
o f cells i nm a ny V olv ox c ol onies is g re a te r th a ni nHyd ra ,

an d ye t Hyd ra is high e r i n th e sc a le o f livi ng thi ngs .

The re a s onfor this is a di ffe re nce i n th e c omplexity of

 DIVI S I O N OF LA B OR 31

or ag niz a ti on The re a re m ore diff e re nt ki nds of cells i n

;

Hyd ra th a ninV olv ox a nd e a ch ki nd is a d a pted to c a rry

,

on a p a rticul a r d uty esse n ti a l to the ex iste nce of the

wh ole And not only is e a ch st ructu ra lly fitted for a
.

defi nite t a sk bu t i n bec omi ng a speci a list it fu ncti ons

,

m ore successfully T o re a li ze the imp ort a nce of thi s f a ct

‘

let us ex a mi ne it f rom a nothe r a ngle .

Let us supp ose th a t a ma nm a kes a na ut om obile He .

c onst ructs th e esse nti a l p a rts a nd it ru ns bu t h a ving to , ,

pe rf orm a ll of the ope ra ti ons i nv olved he is not especi a lly

p roficie nt i na ny one a nd th e m a chi ne is a c omp a ra tively
c rude a ff a ir This st a te o f th ings is c omp a ra ble to wh a t
.

exists i nth e a m oeb a wh e re th e life p rocesses a re c a rried

on equ a lly well by a lm o st a n y p a r t o f the b o dy The .

origi n a l a ut om obile m a n u f a ctu re r ma y bec ome a ss oci a ted

with othe r w orkme n wh o likewise m a ke a ut om obiles
, ,

an d th e s i tu a ti on is duplic a ted by Gonium whe re the ,

membe rs o f the c ommu nity a ll beh a ve i n the s a me

m a nne r Th e re is h e re little if a ny r e a l a dv a nce In
. .

Volv o x onth e o th e r h a nd we fi nd th a t th e cells of the

, ,

c ol ony c om p rise two cl a sses a nd a na na l og ous situ a ti on

,

i n th e f a ct ory a ppe a rs wh e n th e w orkme n a re divided

int o two g ro u p s e a ch with disti nct typ es of w ork An .

i nc re a se i n th e numbe r of w orkme n is not a n essenti a l

fe a tu re I t is now obvi ous th a t to re a ch the level of the
.

Hyd ra org a niz a ti on wh ere seve ra l types of cells a re

,

co Op era ti n g i n the p e r f or m a n ce o f th es e fu ncti on s esse n

ti a l to the life of th e c ol ony th e w orkme nrequ i re to be
,

divided i nt o a c orresp ond ing numbe r of g roups e a ch with ,

its p a rticul a r t a sks .

I nth e high est typ e o f f a ct ories th e di ff e re n t cl a sses of

w orkme na re m a ny a nd e a ch h a v mg bu t one th ing to do
, , ,

bec omes u nusu a lly p roficie nt so th a t th e result of their ,

coOp era ti onis a sm oo thly ru n ning fi nished p roduct I n

,
.

p recisely th e s a me wa y th e high e r species of a nim a ls a nd

pl a nts occupy thei r p ositi on by vi rtue of th e degree to
which the v a ri ous cells Of the b ody a re d ivided i nt o g roups
32 M ULTI C E LL ULA R O R GANI S M S

th a t a s Speci a lists skillfull y pe rform the ir p a rticul a r

, ,

fu ncti ons Insuch c a ses th e individu a l cell l oses i ts i n

.

depe nde nce to a l a rge deg ree but i ncoOp era ting with i ts ,

fell ows p rod uces a norg a niz a ti on th a t is a s much a b ove

Hyd ra for ex a mple a s m ode rn h um a ns ociety is a b ove
, ,

a s a v a ge c omm u n ity .

T I S S UES ORGA N S AN D SY S TE M S
,
In Hyd ra the divi .

si onof l a b or is a b ou t a s c omplete a s i nth e highe r typ es

FI G 8 . .
—B od y C e ll s f rom li ni ng o f i nte s ti ne ;
. (1 ,
fro m skin; b,
0 ,
fa t ce ll ; d mus cl e cell fr o m s to m a ch ; e whi te a n
,
d red bl oo d ,

cel ls ; f n erv e cell ; g ce lls f ro m sa liv a ry gl a n

, ,
d ; h b one cell , .

of pl a nts bu t a b ove this p o int i n th e a nim a l ki ngd om

,

the g re a t m a j ority o f species a m ou nti ng to fully h a lf a ,

mil li on a re of gre a te r c omplexity I n th e dog or ma n

, .
,

for ex a mple th e org a niz a ti ono f th e b ody i n

,
v olves a ve ry
gre a t numbe r o f cells o f diff e re nt types (Fig e a ch of .

wh ich is usu a lly g rouped t ogeth e r to form a tissu e Th us .

we h a ve th e highly c ont ra c tile cells th ose resp onsible for ,

th e m o veme nts of th e b ody fa shi oned i nt o mu scu la r ,

.

tis sue Th is is b ou nd t ogeth e r by cell s a nd fib res f orm

.

in g con nec ti ve tissue is a tt a ched i nm a ny c a s es to bony ,

tissue is impelled to a ct by impulses se nt ou t by ner vou s

,

tissue a nd d i rectly or ind irec tly is p rotected by

,
 TI S S U E S , OR GAN S AND S Y S TE MS 33

ep i theli a tissue of wh ich the ski n a nd li ning of the

l ,

m outh a nd digestive t ra ct fu rnish g ood ex a mples .

While a norga nma y be defi ned a s a ny st ructu re fitted

to pe rf orm a defi n ite functi on a nd th us i ncludes th e
,

nucleus cili a a nd o the r p a rts of a si ngle cell it inv olves

, , , ,

i nhigh e r org a nisms a g ro up ing of tissues Th e a rm for .

,

ex a mple is a norg a n a nd c omp rises a ll of the tissues j ust

, ,

me nti oned Th e na g a in seve ra l org a ns ma y be e ng a ged

.

i nrel a ted w ork a n d so f orm a sys tem Th e oes oph a gus .

,

st om a ch i ntestine live r a nd p a nc re a s a re a ll org a ns

, , ,

se rving i n th e p rocess of digesti on a nd i n c ombi na ti on

,

c onstitute the di ges ti ve system Othe r org a ns c ombi ne

.

to f orm the n er vou s mu scu la r ci rcu la tory a nd othe r

, , ,

systems th a t will be desc ribed i ns ome of th e succeed ing

ch a pte rs .
S ECTI ON 2

OF GR EEN P LA N T S
 C HAP TER IV

T HE M AN UFA CT U R E OF F OOD S AND FUE L S

TH E A N N UA L VA L U E OF F OOD S i nce f ood is i nd is .

p ens a ble its v a lue c a nnot be a dequ a tely rep rese nted i n
,

defi nite te rms I t c onstitutes th e l a rgest item in com

.

me rce Th e food p roduced i n Ca lif orni a a l one i n 1 91 9

.

wa s w orth i n rou nd numbe rs

,
th a t p ro
,

duced i n the U nited S t a tes and i n the

,

wh ole w orld a ni nc onceiv a bly l a rge sum .

A N N UA L PR OD U CTI O N OF F OOD N E CES S AR Y Th e re is .

no st ore of f ood i nna tu re a s the re is of w a te r ; the re is

no fixed a m ou nt for it is a nnu a lly m a nuf a ctu red from th e
,

ra w m a te ri a ls i n g ree n pl a nts
,
Th e qu a ntity v a ries .
,

b oth i n the wild st a te a nd u nde r cultiv a ti on from ye a r

Th ere is ra rely a ny c on
,

to ye a r . side ra ble excess ; th e re

is frequently sh ort a ge ; the re is gene ra lly e nough M a n .

an d a few o the r a nim a ls h a rvest a nd st ore f ood for futu re

use but the su rplus o f supply ove r dem a nd is ge ne ra lly
,

ve ry S ligh t a nd l oc a lly ma y not exist a t a ll The result

, .

of S h ort a ge is a t times dis a st ro us .

TH E F OOD M ATER I A L S — Th e f ood m a te ri a ls s ome ,

t i m es errone o usly sp oke n of a s pl a nt f ood a re c a rb on ,

di oxid (C O ) a ga s w a te r (H O ) a liquid a nd mi ne ra l
Z , , Z , ,

s a lts s olids Ca rb on di oxid o ccu rs i n th e a i r i n ve ry

,
.

dilute p rop orti ons two h u nd redth s of one p er ce nt

,

o n th e a ve ra ge In spite of its ext ra ordi n

. a ry

diluti on th e t ot a l a m ou nt i n th e e a rth s a tm osph e re is ’

e norm ous Bec a u se of th e c onst a nt c o t ributi ons to

. n
th e a tm osphe re from a v a riety of soUrces of wh i ch ,

pe rh a p s ordi na ry c ombusti on is th e most e a sily u nde r

st ood th is a m ou nt a nd p rop orti on a re m a i nt a i ned
, .

37
38 M AN UFA C TU R E O F FO O D S AND FUE L S
W ATER AN D M I N ERA L S A LTS — W a te r which a ls o ,

o ccu rs a s v a p or i n the e a rth s a tm osphe re is t a ke n by

’

pl a nts f rom the s oil i n wh ich it is a lw a ys p rese nt I t .

diss olves th e s oluble s o il i ng redie nts a nd c ove rs th e sur

fa ce o f th e s oil p a rticles wi th fi lms o f diff e re nt thic kness ,

v a rying with th e wet ness o f th e s oil The c omp ositi on .

an d signific a n ce o f th ese fil ms wil l be discuss ed a g a in

( C h a p
. V I ) .The s o i l i ng redie n ts a re mi ne ra l s a l ts the ,

nit ra tes sulph a tes , a nd p h os p h a tes of c a lcium p ot a ssium

, , ,

an d m a g nesium
T H E CO M P OS I T I O N O F F OOD M ATER I A LS — Th e f ood
.

m a te ri a ls a re a ll c omp a ra tively simp le c omp o u nds st a ble ,

an d s a tu ra ted ; th ey do not re a d ily ch a nge ch emic a lly .

Th ey a re c omp o sed of ten chemic a l eleme nts only ou t ,

o f th e m a n y (a b o u t 1 00 ) know n na mely c a rb on (C ) , ,
,

hyd roge n (H ) oxyge n ,

nit roge n (N ) c a lcium (Ca ) , ,

p o t a ssium (K ) m a gnesium (M g) sulph u r (S ) ph os

, , ,

p h oru s (P ) a n ,
d i ron (Fe ) Th e c omp ou nds i n wh ich
.

th ey occu r a s food m a te ri a ls a re a l l s oluble i nw a te r m ost ,

of th em f reely so He nce th ey a re of p ra ctic a lly u nive rs a l

.

o ccu rre n ce .

T H E A B S OR PT I O N O F F OO D M AT ER I A LS to be discussed ,

in Ch a p te r V I t a kes pl a ce th ro ugh roo ts f rom s o il a n

,
d
th rough le a ves f rom a ir i ns oluti on L a nd pl a nts with
,
.
,

roo ts i nth e s oil a n d stems i nth e a ir live i ntwo sets of ,

ci rcumst a nces Thei r unde rgrou nd p a rts i n d a rkness

.
, ,

i n c on t a ct with h a rd h e a vy s olid p a rticles t a ke up

, , ,

s oluble s oil c onstitue nts by thei r roots a nd m o ve them to

th e org a ns wh e re th ey will be c onve rted int o f ood Th ei r .

le a ves sp re a d i nligh t a nd a ir a nd t a ke up f rom th e l a tte r

such g a ses a s will e nte r th eir cells .

T H E W OR K I N G OVE R O F F OOD M AT ER I A L S W a te r a nd .

sa l ts ta ke n up by th e roots f ro m th e s oil a nd c a rb on ,

di oxid t a ke n by th e le a ves f rom th e a i r a re c onve rted ,

f ro m ra w m a te ri a ls into fi nish ed p rod ucts food Th e ,

.

ch a nges t a king pl a ce i n th e m a nu fa c tu re o f f oo ds f rom

ra w m a te ri a ls i n v olve ch a nges i ne ne rgy Ene rgy must .
 R ADIAN T E NE R GY 39

be a p p lied t o the ra w m a te ri a ls to c onve rt them i nt o

f oods This must obvi ously c ome from outside th e
.

p l a nt if the re is to be a ny a ccumul a ti on of f ood The .

ene rgy thus empl oyed re a ch es th e pl a nt i n the f orm of

ligh t .

R ADI A N T E N ERGY The re re a ch es th e su rf a ce of the

.

e a rth from the su nd a ily a na lm ost i nc onceiv a ble a m ou nt

of en e rgy A na lysis S h ows th a t this c onsists of ra ys of
.

di ff e re nt w a ve length s c a p a ble o f d oi ng di ffe re nt kinds

-
,

o f work Thus w e know th a t th e w a rmth of th e su ns

.
’

ra ys ra ises th e tempe ra tu re o f the a i r a n d s o il a nd a ll

othe r obj ects i n th ei r p a th
.

O the r ra ys di rectly a ff ect .

o ur org a n s of visi on a nd stil l othe rs a re used i nm a ki ng

,

p h ot og ra ph s S u nligh t ca nbe divided by a p rism a s by

.

d rop s of w a te r i nt o a spect rum (see Fig 9) or ra i nb ow .

,

Red Ora n
ge I Y e l ow l G reen I
2 Lea ves

FI G 9 . . D ia gra ms to ill ustra te a na lysis of ligh t by th e Spec

troscop e . a ,
sp ectru m of pu re sunligh t ; b, spectru m of sunligh t
p a sse d th rough chl orophyll .

m a de up of seve nc ol ors A the rm omete r pl a ced bey ond .

the visible c ol or a t th e red end of th e spect rum will Sh ow

th a t th e re a re i nvisible h e a t o r th e m l ys
r a ra i n a dditi o n
to the visible on es which a re c lled ligh t
a o r n
lumi us ys o r a .

S imil a rly a ph o t o gr a ph ic pl a te exp o sed bey o n d th e V i o let

en d of the spect rum w ill u nde g ch emic l ch ge th e e
r o a a n r ,

by dem onst ra ting th a t i nvisible but ch emic a lly a c tive

ra ys cc
a o mp a n y th e in visible h e a t r a ys a n d th e visible
luminou s ones Th us we see o ly p t n .
a a r
,
o f n
th e e e gy r

n
which th e su ns ra ys c ont a i but we k w th t he t ls
’
n o a a a
,
o

em a na tes f rom the sun a nd th a t ce rt a in o the r i nvisible ,

ra ys do ch emic a l w ork .

P H OT O S Y N T H E S S
I Th e c o mbi n a ti o
.
n o f c a r b on di o xid
4-
0 M AN U FA C T UR E OF F O O D S A N D FU E L S

(C 0 2 ) and w a te r (H O ) to f orm sug a r (C l O ) is a c

z e e

c omplished only by me a ns o f ligh t e ne rgy a bs orbed by

the g ree n p a rts of pl a nts a nd i s the refore c a lled ph o to
synthesis I t is the m ost imp ort a nt chemic a l re a c ti on
.

in n a tu re fo r i t u n
,
de rlies th e m a nu f a c tu re o f a ll foods
an d fuels Th e ra ys a bs orb ed a re m a i nly ne a r th e red
.

an d the blue i n the spect rum ; th e a bs orbi ng a ge nts a re

g ree n pigme nts in cells ne a r th e su rf a ce of th e b odies
O f pl a n ts .

P Y
C H LO R O H L L — Th e g ree n c ol or of le a ves or o the r
gree n p a rts o f pl a nts is due to the p rese nce of ce rt a i n
subst a nces th e chl orophyll pigme nts or chl orophyll con
, , ,

fi ned i nm o st i nst a nces to sm a ll b od ies c a lled pl a stids or

ch rom a t oph ores Chl orophyll a bs orbs ligh t of ce rt a in
.

c ol ors or w a ve le ngth s only Th e c ol ors wh ich p a ss

-
.

th rough give th e ch a ra cte ristic g ree nto le a ves a nd o the r

chl orophyll c ont a ining p a rts or org a ns C hl orophyll
-
.
,

s oluble in a lc oh ol chl orof orm eth e r a nd othe r f a t sol

, , ,

vents a ppe a rs to be c omp osed of f o u r c omp o u nds o f

,

disti nct c ol or qu a lities These h a ve bee nsep a ra ted wi th

.

g re a t S kill a nd thei r c omp osi ti on d ete rm ined T wo of

,
.

th em a re gree n two a re yell ow Th ey a re c ryst a ll i ne

,
.

subst a nces c omplex a nd u nst a ble th ou gh no t equ a lly so

, , ,

a s sh ow nby th e f a c t th a t th e yel lowi n g o f le a ves I s d ue

to th e pe rsiste n ce o f one or b o th o f th e yell o pigme nts w
while th e gree n ones a re u nde rg o ing withd ra w a l or
dec omp ositi on .

O CC UR RE N CE O F C H LORO P H Y LL Th e chl orophyll pig .

me nts do no t o ccu r i na ll pl a nts a nd th ey occu r only or ,

ma i nly i nth o se p a rts o f g ree npl a nts wh ich a re exp osed

to ligh t . Th e pigme nts occu r only i n th e p l a stids or
ch rom a t oph ores o f cells situ a ted so ne a r th e su rf a ce o f
th e pl a nt b ody th a t a n a dequ a te supply o f ligh t w ill
pe net ra te th em C hl orophyll will be f ormed h oweve r
.
, ,

on ly u nde r suit a ble c onditi ons na mely wh e n th e pl a nt , ,

a bs o rbs suit a ble a nd sufficie nt i rons a lts f rom th e so il a nd

the re is e nough light Al th ough i ronis not a c onstitue nt
.
 W HAT LI M IT S P H O T OS Y NTH E S I S 41

of chl orophyll pl a nts will not be he a lthy with ou t i t f or

, ,

they will not be g ree n E xcep t for pi ne S eedli ngs ordi

.
,

na ry pl a nts W il l not f orm chl orophyl l i nd a rkness This .

f a ct is utilized i nth e bl a nch i ng o f cele ry a nd a sp a ra gus ,

a s well a s bei n g a m a tte r of c omm onobse rv a ti on .

W H AT L I M I T S P H OT OS Y N T H S S E I .
— Ce rt a i n defi nite
fa ct ors ca n be rec ognized a s limiti ng a nd c ont roll ing
p h ot osynthesis These a re :
.

( )
1 T h e a m o u n t o f c h l or o h
p y ll exp osed to the li gh t .

This will v a ry not only with th e qu a ntity of p igment

,

a ctu a lly c on t a i ned in th e ch rom a t oph ores but with th e

number a nd th e exp osu re of the chl orophyll c ont a ining -

cell s i n le a ves a n d stems Pl a nts wi th sc a nty le a fa ge

.

Offe r less chl orophyll exp osed to ligh t th a n th ose with

n
m ore a bu d a nt f oli a ge .

(2) The a mou nt of li gh t tha t becomes effecti ve .

Pl a nts or pl a nt p a rts ma y be sh a ded ou t of existe nce

-
.

This is cle a r i n f orest a nd Ch a p a rra l for th e f oli a ge ,

an d o fte nth e l owe r b ra nch es thus die or neve r devel op

inthe sh a d ow of b ra nches a nd f oli a ge a b ove them . Com
p a ra ti v ely few species of pl a n ts a re a ble to th rive i nthe
sh a d ow of o the r pl a nts or of op a que obj ects Sh a ded .

pl a nts usu a lly m a ke up for deficient ligh t by gre a te r ex

p os u re o f chl o r o phyll eithe
,
r by f o r m in g chl o r ophyll i n
the epide rmis c ove ring th e le a ves or by l a rge r a nd ,

b roa de r le a ves
—
.

( )
3 T h e q u a li ty o f gli h t Th e ligh t r e a ch i ng th e
fo rest fl oor ma y be a s u nsuit a ble i n qu a lity a s it is i n
sufficie nt in q u a nti ty for i t ma y l a ck the p a rticul a r ra ys
‘
,

used i n ph ot o synth esis by re a son o f th ei r bei ng a l re a dy

a bs orbed by th e f oli a ge a b ove .

( )
4 T h e a m o u n t of wa rm t h .D iff e re nces i n tempe ra

tu re a re f ou nd to a ffect ph ot osynthesis i n na tu re m ore

th a n a ny o the r S i ngle f a ct or Ph ot osynthesis will not
.

t a ke pl a ce i n m ost pl a nts a t tempe ra tu res cl ose to th e

freezing p oint Th e ra te of ph o t osynthesis rises m ore
.

ra p idly th a n th e tempe ra tu re h oweve r up to a ce rt a in

, ,
42 M AN U FA C TU R E OF FO O D S AN D FU E L S

p o int a nd the nf a lls ra pidly if th e tempe ra tu re c onti nues

,

to rise E xcessively high tempe ra tu res a re rec orded i n

.

c onnecti onwi th expe rime nts o nd ese rt pl a nts ; bu t ord i

ma rily o u t o f d oors a s well inth e t ropics a s i ntempe ra te
, ,

regi on s the tempe ra tu res to wh ich pl a nts a re subj ected

,

a re ,
a f te r a ll c omp a ra tively m ode ra te Th e e ff ect o f
,
.

tempe ra tu re on ph o to sy nthesis must be c onceived a s a

chemic a l p rocess i nfl ue nced no t only by the a cti on o f
h e a t up on th e ra w m a te ri a ls bu t by th e a c ti ono f h e a t
up on th e living cell s in wh ich th e m o st imp ort a nt
chemic a l re a cti oni nna tu re t a kes pl a ce Th e tempe ra .

tu res which a re m ost f a v ora ble for o the r p ro t opl a smic

a ctivities a re f ortu n a tely a ls o th ose highly f a v o ra ble for

ph o t osy nthesis This is th e re a s on for th e l uxu ri a nt

.

g rowth o f th e t rop ics ra th e r th a n bec a use the re is a ny

,

c onside ra bly g re a te r a m ou nt of ligh t or a ny supe ri or

qu a li ty of th e ligh t i n t ropic a l regi ons .

( )
5 V a ri a ti o ns i n ca rb o n di oxid a re so sligh t a n d so

in f reque nt th a t th ey a re o f no p ra ctic a l imp ort a nce i n

a g ricultu re ; but th e ve ry g re a t d iluti ono f c a rb ondi oxid

i nth e a i r i nspite o f i ts e normo us v olume fixes th e ra te

, ,

a t wh ich f oo d ca nbe m a d e Th is ca nbe sh ow nby l a b ora

.

t ory expe rime nt for inc re a si ng th e p rop orti ono f c a rb on

,

di oxid i nth e a i r i nwhich pl a nts a re g row nwill i nc re a se

th e yield o f sug a r especi a lly if th e ligh t is a ls o i n
,
c re a sed ;
but th e ide a o f a ddi ng to th e c a rb o n d i oxid c onte nt of
th e a i r o na c omm e rci a l sc a le a s h a s bee nsuggested m ore
,

th a no nce seems o f no p ra ctic a l imp ort a nce a t p rese nt

,
.

T H E P H OTO C H E M I CA L R EA CT I O N S I N P H OT OS Y N T H ES I S .

Th e ge ne ra l na tu re o f the p rocess c a lled ph ot o sy nthesis

ma y be exp ressed i nth e f oll owi ng equ a ti on:

6 CO2 6 H2 0 CGHnO6 6 02
Ca rb o n D i oxid W a te r S u ga r O xyg en

Th is eq u a ti onexp resses th e f a ct th a t ou t of six m olecules

o f c a rb ondi oxid a nd six m olecules o f w a te r on e m olecule ,
 VO N B A YE R ’
S H Y P OT H E S I S 43

of sug a r (gluc ose ) a nd six m olecules o f oxygen a re

formed The oxyge n wh ich equ a ls the v olume O f c a rb on
.
,

di oxid t a ke n i n is give n Off to the a i r The re is g ood

,
.

re a s on to thi n k h oweve r th a t this equ a ti on me rely ex

, ,

p resses th e fi nal result O f a se ries o f che mic a l re a cti ons ,

th e det a ils of wh ich a re not now k now n The m ost .

pl a usible hyp othesis p rop osed reg a rdi ng th e c ombi na ti on

is th a t of th e em i nent E u rope a nchemist v on B a ye r ,
.

’
VO N B A Y ER S HY P OT H E S I S O F P H OTOS Y N T H ES I S .

Acc ordi ng to this hyp o thesis fi rst one m olecule of c a rb on , ,

di oxid ma y be c ombined with one m olecule of w a te r to

p roduce formic a cid with th e libe ra ti onof one m olecule
,

o f oxyge n; sec on d this f ormic a cid is reduced to f ormic

,

a ldehyd e with th e libe ra ti ono f m ore o xyge n; a n d thi rd ,

the f ormic a ldehyde m olecules ma y c ombi ne with one

an o th e r ma y p olyme rize so th a t a m olecule o f sug a r

c ont a ini ng six a t oms of c a rb on is p roduced Th e equ a .

ti ons c once rned a re a s foll ows

CO2 H ZO H2 0 0 3 (c a rb onic a cid )
H2 C 0 3 HC OOH O (formic a cid a nd oxyge n)
HCOOH H OOH O (f or mic a ldehyde a nd o xyge n)
6 HCOH (36 l 0 6 (gluc ose sug a r )
No chemist or physicist h a s thus fa r p roved by expe riment
the a ccu ra cy of th is ve ry suggestive hyp o th esis a nd the ,

field is still open for i nvestig a ti on .

T H E P R OD U CT S O F P H OTOS Y N T H ES I S Foods — I t is .

ce rt a i n th a t a sug a r c ont a i ni ng S i x a t oms of c a rb on

(C H 0 ) or a multiple of six (C H OH ) i nthe m ole
6 1 2 6 , I Z ZZ ,

cule is m a de i n d a yligh t i n chl orophyll c ont a i ni ng pl a nt -

cells a nd th a t fro m this sug a r s oone r or l a te r a ll o the r

,

foods a nd m o st of the oth e r p roducts of pl a nts a re f ormed

by a dditi on of one or a nothe r of th e che mic a l eleme nts
fo u nd i n th e di ffe re nt f ood m a te ri a ls The f ood ma y .

a ccumul a te a s sug a r or it ma y be st ored i n i n

,
s oluble
form temp ora rily or for l onge r times i nthe f orm of
st a rch or cellul ose fa t or oil Nit roge na nd ph osph orus
, ,
.
44 M AN U FA C T U R E O F FOO D S AN D FU E L S
'

ma y be a dded f orm ing still m ore c omplex c a rb on com

,

p o u nds a s we sh a ll p rese ntly see Alth ough the em i ne nt

,
.

pl a nt physi ol ogist S a ch s l ong a go sp oke o f st a rch a s the

”
fi rst visible p rod uct of a ssimil a ti on we know now th a t ,

h is f a mous dictum wa s only a pp rox im a tely c orrect .

S u ga r is the fu nda menta l p rodu ct of p ho tosynth esis a nd ,

beca u se of thi s fa c t ph o t osy nthesis i s the m ost imp ort

an t ch emic a l re a cti oninn a tu re
” for a l l f ood is built up
,

f rom sug a r .

Fu els — T h e fuels no less th a n th e foods a re a ls o O f

veget a ble origi n a nd a re built up f ro m th e sug a r m a d e
i n ph o t o synth esis .W ood for ex a mple is obvi ously so
, ,
.

Coa l is well know n to be th e ch a n ged rem a i ns O f w ood

an d o the r p a rts o f pl a nts I t is supp osed th a t oi l a nd
.

na tu ra l ga s h a ve a ls o bee nde rived from the cell dep osits -

O f a c cumul a ted pl a n t rem a i ns i n s ome c a ses f rom the

-
,

oil p rod uced by di a to ms Alc oh ol is obt a ined (see C h a p

. .

XX I I I ) by th e fe rme nt a ti ono f sug a r or of sug a r de riv a ,

ti v es rec on ve rted i nt o sug a r Th e study the ref ore o f

.
, ,

ph o t osynth esis is o f the utm ost imp ort a nce f rom th e

ec onomic p o int o f view S i nce m a ns c omfort a s wel l a s
,
’

h is no u rishme nt a re a bs olutely depe nde nt up on this re

a cti o n
. E ve ry thi ng th a t ma nu ses wh ich is no t mi ne ra l
c omes di rectly or i ndi rectly f ro m th e c ombina ti ono f ca r
b o n d i oxid a nd w a te r i nt o sug a r i n the g ree n p a rts o f
pl a nts i nw a rmth a nd ligh t With th e excepti ono f wi nd
.

an d w a te r i n m o ti on o r th e p owe r Obt a i n
,
ed the refrom ,

ma nu ses these s a me fuels a s th e s o u rces o f a ll the e ne rgy

whi ch h e requi res I ndeed u nde r ce rt a i nm a rket c ondi
.
,

ti ons i t is m ore p rofit a ble for th e f a rme r to bu rn c orn

th a n c oa l .
 C HAPTER V

THE B UILDI NG O F T HE LI VIN G M A C HI N E

TH E L I VI N G M A CH I N E th e livi ng b ody of pl a nt or
,

an im a l c onsists of p ro t opl a sm th e org a nized u nits of

,

which a re c a lled cells (see C h a p I I ) . .

T H E C O M P OS I TI O N OF P R OTO P LA S M — M ic rosc opic .

an a lysis a n d th e meth ods of the physic a l chemist sh ow

th a t p ro t opl a sm c onsists o f s ome subst a nces too c oa rse
to p a ss th ro ugh a p orcel a in filte r th ough g o i n g th rough ,

o the rs less fi n e S uch subst a nces a re c a lled c oll o ids or

.
,

a re s a id to be i n the c oll o id a l c on diti on or st a te Th e .

an a lytic a l ch emist sh o ws th a t p ro t opl a sm is m a de up ,

i na dditi onto w a te r o f p rotei ns o f v a ri o us s orts a nd th a t

, ,

it ma y encl ose a g re a t v a riety of subst a nces i n s oluti on

or o the rwis e c a rb o hyd ra tes f a ts a n
,
d o ils s a lts a nd o the r
, , ,

elect rolytes These simple a nd c omplex subst a nces

.
,

w a te r s a lts oils a nd p ro tei ns a re built i nt o a c oll o id a l

, , , ,

system or systems F ood is u sed th e ref ore to build a nd

.
, ,

to m a i n t a in this c omplex system B u t th e food must .

fi rst be m a de .

N I TR OGE N Th is eleme nt f ormi ng f ou r fi fth s of th e

.
,
-

e a rth s a tm osphe re is used by m ost pl a nts a nd by a l l

’

an im a ls only i n c omp o u nds M o s t pl a nts a bs orb these .

as n it ra tes from the s oil Ince rt a i ni nst a nces a mm onium

.

c omp o u nds a nd cy a na mid ca nbe used a s s ou rces o f nit ro

g e n,
a n d a few species o f c a rn iv o ro us pl a nts a re kn ow n
which c a ptu re or e nt ra p sm a ll i nsects a nd dev ou r or con
sume them to secu re nit roge n N it ra tes of c a lcium a nd .

p ot a ssium a re the c omm ons a lts used by pl a nts a s s o u rces

of n it rogen Nit roge n is a c onstitue nt eleme nt of p ro
.

tei n c omb ined with c a rb on hyd roge n a nd oxyge n to

, , ,

45
46 B UI LDING OF T HE LI VING M A C HI N E
form m olecules O f ve ry c onside ra ble size a nd c omplexity .

The sy nthesis of p ro te i ns ca nm o st na tu ra lly be c onceived

a s c on stitu ti ng a se ries o f i nc re a si ngly el a b ora te com
p ou nds st a rting f rom a sug a r a nd a nit ra te a s the s o u rces
,

o f th e a t o ms to be c ombi n ed .

P H Y S I CA L P R O P ERT I ES O F P ROT EI N S — These sub .

s ta nces a re difficult to d efi ne b oth bec a use th ei r c omp osi

,

ti onis only inc ompletely k now n a nd a ls o bec a use they ,

a re so dive rse . They h a ve h oweve r ce rt a i n p rope rties

, , ,

ch emic a l a nd physic a l i nc omm on; they a re c oa gul a ted

,

by he a t a nd by a lc oh ol ; they a re simil a rly s oluble i n

w a te r ; they ma y be p recipit a ted f rom a que ous s oluti ons
by v a ri ous s a lts Th ese c omp ou nds c onsisti ng of ve ry
.
,

l a rge a nd ve ry c omplex m olecules c ont a i ni ng i nv a ri a bly

c a rb on hyd roge n o xyge n a nd nit roge n a nd m a ny o f
, , , ,

th em sulph u r a nd ph osph orus a s well eithe r f orm con ,

sti tu en t p a rts o f th a t livi ng st ructu re c a lled p rot opl a sm ,

o r a re p rese n t i ni t a s p roducts or b y p roducts We ma y -

.

pe rh a ps the bette r c onceive h ow p ro t opl a sm is buil t if

we a ttempt th rough a cl a ssific a ti ono f th e org a nic nit ro
,

ge nous c omp ou nds o f the se ries le a di ng to th e p rotei ns ,

to g a i n a n ide a o f th ei r rel a tive c omp ositi on a n d co m

p l ex i ty
. Thus the simplest c omp ou nds i n this se ries ,

th e a mi n -
o a cid s a n ,d the nucle o p ro tei n s th e m ost co m
-
,

plex a re a pp a re ntly c onnected by c omp ou nds o f i nte r

,

medi a te c omplexity a nd c omm on p rope rties .

T H E A M I N O AcI Ds — These subst a n

-
. ces ch a ra cte ri zed ,

by c onsisti ng o f o ne or m ore a mi no (N H ) g roups a nd of Z

a n org a n ic a cid ra dic a l (c a rb oxy l gro up GOO H ) a re ,

much bette r k now na s th e p roduc ts O f p rotei ndec omp osi

ti on th a n a s p a rts of a n a sce ndi ng sy nthetic se ries of
o rg a n ic nit roge n c omp ou nds N eve rth eless thei r Oom.
,

p o siti ona nd st ructu re s uggest me a ns by which th ey ma y

c ome int o exis te nce Th us i t is k now n th a t f rom th e
.
, ,

nit ra tes t a ke n up f rom th e s oil nit rites a nd a mm oni a ,

ma y be de rived ; f rom the l a tte r hyd rocy a nic a cid

—
,

c omm o n i n n
pl a t tissues is O bt a i ned F rom th e a o .
 T HE S Y N TH E S I S OF P R O T E IN S 47

ti onof a mm oni a a nd hyd rocy a nic a cid up ona na ldehyde

a simple a cid migh t be f ormed thus ,

C H3 C OH H ON .
N H . OH
a cet a l d hyd
e e a mmoni a

H —
O 3 OH— N H 2
— COOH N H,
a mmoni a
a la n
In

S uch synthesis ma y t a ke pl a ce i n le a ves D u ring th e

a .

h ou rs of d a yligh t a ldehyde f orm a ti onma y occu r a s a p a rt

o f th e p rocess o f ph ot osyn thesis (see Ch a p I V ) At the . .

s a me time a p a rt o f th e a mm oni a de rived f rom the ni

,

t ra te a nd nit ri te ma y be c onve rted i nt o hyd rocy a nic a cid .

Th is ma y c omb ine with the a ldehyde wh ich is being

f ormed a t th e s a me time While this c onj ectu re a s to .

the na tu re o f the p rocess ma y not p rove to be c orrect i n

a ll its det ai ls it suggests a ve ry p romising li ne for
,

investig a ti on .

T H E S Y N T H E S I S O F P R OT E I N S F R O M A M I N O A C I D S -
.

P roteins h a ve a c tu a lly bee n synthesized f rom a mi no

a cids a n d a mi no a cids a re a lw a ys f o u d i nf o od s oluti ons
-
n
on thei r wa y to p a rts wh e re p ro tei n is bei n g st ored or
p rot opl a sm is being c onst ructed yet nothi ng or a lm ost ,

nothing is k now n a b o u t th e ch a nge wh ich t a kes pl a ce

i n the b ody of th e pl a nt Th e a mi no— a cids dis a ppe a r
.

an d th e c omplex p ro tei ns a ppe a r i nste a d I t is p rob a ble .

th a t a ny norm a l pl a nt cell ca n m a ke th e p rotei ns which

it requi res i f it receives suit a ble c a rb ohyd ra tes a nd
,

a mi n o a cids f or c on
-
st ructi on The re is no evide nce to .

S h ow th a t p ro tei nsy n thesis t a kes pl a ce m ore ra pidly i n

-

on e p a rt o f th e pl a n t th a n a nothe r Th e a mi no a cids — .

ma y be m a de ch iefly i nor ne a r th e chl orophyll c ont a i ni ng -

cells i nwh ich the c a rb ohyd ra tes a re f ormed i n d a yligh t ,

th ough th e f orm a ti on of p rotei n a ppe a rs to be quite

independe nt of ligh t .

We ma y c onceive th e p rocess of p rotei nsynthesis from -

a mi n o a cids t a ki n
-
g pl a ce a l ong a se ries which st a rts with
glyce rose (CgH O ) l a ctic a cid (a ls o C H O ) a nd one
s s , a fi s ,
48 B UILDING OF THE LIVING M A C HIN E
o
” r
m o re a mi no a cids O f these
-
subst a nces .
g l y ce ro se is ,

on e o f th e ve ry simplest sug a rs k n ow n a n d la c ti c a cid ,

is a co mm o n o rg a nic a cid f o rmed for ex a mple i n the ,

s o u ring o f milk (see C h a p XX I I I ) Th e a mi no a cid . .

-
,

a la n i n (C H N 0 ) Comm on
3 7 ly f ormed i n th e hyd rolysis
2 ,

o f p ro tei n s se rin (C H N O ) a s omewh a t less simple

, 3 7 S ,

c omp ou nd a nd c ys tei n (C H N 0 8) p a rticul a rly i nte r

, 3 7 2 ,

es ti ng as c ont a i ni ng sulph u r ma y be a rra nged i na se ries , ,

(wi th glyce rose a nd l a c tic a cid ) a s p rop osed by Abde r ,

h a lde n th us
,

gl ycero se l a ctic a cid a a l n

in serin cys tein p rotein
C H0 C H3 C H3 C HzOH C HgS H

C H OH C H OH C HN H2 C HN H2 C HN H2

C HzOH COO H COOH COOH COOH

I f ph osph orus is a dded to the p rotei nm olecule th e re re

s ul ts a n ucle o p ro tein Th e nucle o p ro tei ns a re th e m ost
-
.
-

c omplex subst a nces k now n .

T H E ORGA N I Z AT I O N O F P R OT O P LA S M The p ro t opl a sm .

o f the cell c o n sists Of two di ff e re nt p a rts cyt opl a s m a nd ,

nucleus Th e l a tte r will be discussed m ore fully i n a

.

l a te r ch a pte r (XX V ) a nd so ma y be dismissed h e re

with a b a re me nti on Th e cyt opl a sm exists i n two d if
.

ferent c o n diti ons a cc o rdi ng to th e rel a ti ons o f its v a ri ous

,

c omp o ne nts to one a no th e r Inone c onditi oni t is fluid .

an d c a p a ble o f fl o wi ng m oveme nts ; i nth e o the r it fo rms

a mo re o r less fi rm j elly like st ruc tu re ,
I n th e liquid
-
.

c onditi on w a te r c ont a i ni ng s a lts i n t rue s olu ti onc ons ti

,

tu tes a c on ti nu ou s ph a se i nwh ich th e c oll oid a l p a rticles

o f p ro tei n o il , an d o the r subst a n
,
ces a re i n suspe nsi on
o r i n c oll o id a l s oluti o n In th e fi rme r c on diti on the
.

p rotei n p a rticles c oa lesce to form a c onti nu ous ph a se

which e ncl o ses m i nute d rops o f w a te r a nd diss olved s a l ts
a s well a s u l tra mi croscO p i c dro ps o f O i l a n d f a ts Which .

o f these two c on diti ons will exi st a t a ny give n time de

 T HE FUN C TI ONING O F P R O T O P LA S M 49

pe nds inp a rt a t le a st a nd pe rh a p s wh olly on th e elec

, , ,

tri ca l ch a rges of the diss olved s a lts a n d th e p rotei nmesh

w ork Cyt opl a sm is th us see n to be a n org a nized a nd
.

c omplic a ted c oll oid a l system i n which p owe rful bu t i m

perfectly unde rst ood forces a re a t w ork a t th e b ou nd a ries
betwee n the p rotei n gel a nd th e liquid oi l a nd w a te r
s oluti ons .

TH E F U N CTI O N I N G OF P R OT OP LA S M — We ma y thu s .

c onceive p rot opl a sm to be m a de a nd m a i nt a i ned a s a

self ope ra ting mech a ni sm Inresp onse to th e a ffinities o f
-

the p rotei ns which a ssemble or a rra nge themselves with

w a ter i nt o c oll oid a l systems a nd by me a ns of th e a cti ons
,

an d i ntera cti ons on th e f a ces of these systems whe re a ,

gre a t p a rt of the e ne rgy requi reme nts of th e livi ng or

g a n i sm a re devel oped a n d s a tisfied Th e ope ra t i ng o
. r

fu ncti oni ng of the system th us desc ribed c onstitutes liv

i ng or life
,
.Living p ro t opl a sm is th e n a st ructu re i n , , .

which ch emic a l p rocesses go oni nresp onse to a t omic a nd

m olecul a r a ffi nities physic a l a s well a s chemic a l ch a nge
,

c ont inu a lly t a king pl a ce with th e bi ndi ng rele a se a nd

, , ,

a pplic a ti on o f p owe rful f orces which a ct up on th e p a r

ti cl es of m a tte r h oldin g th em i npl a ce a s p a rts O f systems

, ,

pushi ng them ou t or pulling th em i n Th us th e fu ncti on .

in g o f p ro t opl a sm livi n g,
is c onti nu a l
,
ch a nge physic a l ,

an d chemic a l .

T H E G R OWT H OF P ROTOP LA S M Th e g rowth of p rot o

pl a sm is one of th e fu ncti ons of this self Ope ra ting mecha -

nism a nd c onsists i ni nc re a si ng or a ddi ng to the c oll oid a l

,

systems which c onstitute th e livi ng subst a nce Thus .

b oth w a te r a nd c oll oid a l subst a nce i nc re a se a s p ro

top l a sm g rows Th e p rop orti ons v a ry indiffe re nt st a ges
.

o f the p rocess of g ro wth bu t th e systems a re c on

,
ti nued
with thei r physic o ch emic a l c a p a cities a nd ch a ra cte r
-
.

TH E D EAT H OF P R OT OP LA S M Whe nch a nge bec omes

.

imp ossible or whe n it bec omes excessive i n th ose col

, ,

l oi da l systems of w a te r p ro tei ns etc which c on

, ,
stitute .
,

livi ng pr ot opl a sm de a th e nsues D e a th ma y be p ro

,
.
50 B UILDING OF T HE LI VI NG M A CHIN E
duced by a lm o st a ny inte rrupti on or i nte rfe re nce with
the ope ra ti onof the systems which c onstitu te p ro topl a sm .

Thus c oa gul a ti on p ro duced by h e a t a lc oh ol or o the r

, , , ,

ag en ,
t st oppi ng a nd p reve nting chemic a l ch a nge is f a t a l , .

Th e s oluti ono f c oll oid a l gels by a cid a lk a li or elect ricity , ,

wil l c a use de a th Chemic a l ch a nges which result i np re

.

ci p i ta ti on o r s oluti on o f p ro tei n o r othe r esse nti a l con

s ti tu en ts of th e system will st op ope ra ti on a n d this
c onstitutes de a th .

T H E M I N ERA L C O N S T ITU E N T S Wh a t the b o t a nist .

h a s c a lled th e a sh c on
“
sti tue nts a m ong th e food ma
teri a l s o f pl a n ts na mely c a lcium m a gnesium a nd p o
, , , ,

ta ssi u m a nd the sm a ll a m ou n
,
ts of i ron c orresp ond to '

wh a t the a nima l physi ol ogist spe a ks o f a s diet a ry a cces

sori es. These eleme nts a re a ll indispe nsa ble to h e a lth
an d norm a l a ctivity but th ey do not a ll a pp e a r a s con
,

sti tu en ts o f p ro t opl a sm They a re t a ke n u p f rom the

.

s o il inth e fo rm o f nit ra tes sulph a tes ph osph a tes While , ,

.

sulph u r a nd ph o sph orus a re c onstitue nts of p rot opl a sm

with ni t roge n c a rb on hyd roge n a nd oxyge n a nd wh ile
, , , ,

m a g nesium is a c onstitue nt of chl orophyll c a lcium p o , ,

ta ssi um a n
,
d i ro nh a ve neve r bee n sh ow nby a n a lysis to

be c onstitue nts eith e r of th e livi ng p rot opl a sm or o f a ny

esse nti a l p rod uct S pecul a ti on a nd eve n a sse rti on h a ve
.
, ,

bee n f reely i ndulged i n; but bey ond rec o g nizi ng ce rt a i n

a pp a re n t uses i nce rt a i npl a nts a nd s ome of th e physi ,

ca l a n d ch emic a l p rope rties o f th ese eleme nts whe reve r

they o ccu r o ne c a nno t now go Ince rt a i npl a nts c a lcium
, .

neut ra lizes th e ox a lic a cid which migh t o th e rwise a ccu mu

l a te i nh a rm ful q u a ntity b ri ngi ng it d ow na s a c ryst a lli ne
,
'

p recipit a te o f ca l cium ox a l a te Th a t this is i nv a ri a bly .

i ts fu n cti on is d isp ro ved fo r ex a mple by th e f ree o x a lic

, ,

a cid i n rh ub a rb i n w ood s o rrel e tc

,
N or d oes c a lci um ,
.

neu t ra lize th e o the r a cid s which ch a ra cte ristic a lly a ccu

mul a te i n th e tissues o f pl a nts — cit ric a cid i n lem ons ,

m a l ic a cid i na pples s a licylic a cid i n c ra nbe rries etc

, ,
.

P OTA S S I U M sh ow nto be feebly ra d i o a ctive ma y owe

,
-
,
 I R ON 51

its m a in useful ness to this p rope rty I t is a p owe rful

.
-

a lk a li its i on
,
s a re ve ry a ctive Fu rthe rm ore expe riment
.
,

i ndic a tes th a t c a lcium a nd p o t a ssium i ons a re b oth

needed to neut ra lize e a ch o th e r Th e c omp ou nds of
.

p o t a ssium a re freely s oluble a nd he nce th e m ore a ble to

displ a y th ei r m olecul a r a ctivity .

I RO N is equ a lly i ndispe ns a ble Ins ome wa y now quite

.

u nguessed a nd u np roved it t a kes its p a rt i n th e met a

,

b oli c p rocesse s of th e org a nism bei ng i n pl a nts a s i n

,

an im a ls i ntim a tely if not di rectly c onnected with the

f orm a ti on of c ol oring m a tte rs Th us i ron is used i n
.
,

medici ne to a ssist i nthe a llevi a ti onof a naemic c onditi ons

i n hum a n bei ngs rest ora ti on to h e a lth bei ng i ndic a ted
,

by heighte ned c ol or ; a nd i n pl a nts bl a nch ing ma y often

be cu red by a dministe ri ng i ron .
 C HAPTER VI

T HE I N CO M E M O VE M E NT S A ND O UTG O
, ,

O F M AT E R I AL S I N PLAN T S

CELLS AR E B AGS O F W ATER an — Cells

d o rg a nisms ,

th emselves a re esse nti a lly b a gs of w a te r Th e physic o

,
.

chemic a l systems cells c omp osed O f w a te r a nd c oll oids

, ,

a rra n ging themselves i nt o self ope ra ti ng mech a nisms a re

-
,

eithe r sa c like th e c onte nts o f th e s a cs or b a gs bei ng

-
,

a que o us s oluti on s e ncl osed i nw a lls of livi ng p rot opl a sm ,

or th e p ro t opl a sm is disp o sed in a sp on ge like m a ss en -

cl osi ng d rops o f w a te r In eith e r c a se th e beh a vi or of

.
,

living cells is th a t O f b a gs o f w a te r so fa r a s i nc ome , ,

m oveme nts a nd o utg o o f m a te ri a l s a re c once rned In

,
.

pl a nts th e s a cs or b a gs a re mech a nic a lly supp orted a nd

st re ngth e ned by e ncl o si ng cell w a lls O f cellul ose D is .

s olved i n the cell sa p a re m a ny subst a nces so fi nely ,

d ivided th a t m olecul a r m o veme nts a nd ch emic a l ch a nges

ma y ta ke pl a ce wh ich w o uld be imp o ssible i n th e s olid
st a te Th u s we fi nd di ffu sio n o f a disso lved sub st a nce
.

t a ki ng pl a ce th rough o u t a v olume o f liquid a nd a p er

fectly u n ifo rm dist ributi onresul ti ng ; but th e m oveme nt
o f diss o lved subst a n ces i nt o a nd o u t o f cells implies th e
a bili ty o f th e m olecules o f th e d i sso lved subst a n ce to
p a ss th ro ugh such memb ra nes as th e c oll oid a l system
wh ich we h a ve desc ribed p osses ses .

P ER M EA B I LI TY — On th e su rf a ce of c ont a ct betwee n
.

two d i ff e re n t m a te ri a ls a s for ex a mple betwee n p rot o

,

pl a s m a nd cell w a ll betwee nnucleus a nd cyt opl a sm a n

, ,

a c tu a l memb ra n e is f ormed by th e livi ng cyt opl a sm .

This memb ra ne ma y be too th i n to be see n u nde r th e

mic rosc ope bu t its existe nce ma y be o the rwise p roved
,
.

Thu s th e c ol oring m a tte r i nroo ts a nd othe r p a rts of red

52
 O S M O TI C P R E S S UR E 53

beets a nd th e sug a r i n the roo ts of sug a r beets a re re

, ,

ta i n ed withi nth e cells ; they do n ot le a k o u t i n t o the s oi l

Onth e o the r h a nd w a te r is t a ke nup by the roots He n
.

,
ce .

we see th a t cell memb ra nes ma y be pe rme a ble to w a te r

-
,

bu t not to a ll subst a nces eve n th ough they ma y be dis

s olved in w a te r C ell memb ra nes a re the ref ore sp oken
.
-

of a s p a rti a lly pe rme a ble or semi pe rme a ble memb ra n -

es .

OS M OTI C P R S S U R E
E — M olecules o f w a te r m ove with
.

inc re a sing f reed om a s w a te r is t ra nsformed f rom the s olid

to the li quid or to th e g a s e ou s c on diti on S imil a rly .
,

the m olecules of s a lt for ex a mple m ove with i nc re a sing

, ,

sp ee d a nd a mplitude a s th e s a lt diss olves i n w a te r In .

c onseque nce th e m oveme nts of th e m olecules of sub

st a nces be a ting up on the w a lls of th e e ncl osi ng or dis
s olvi ng medium will exe rt p ressu re This p ressure is .

know na s osm o tic p ressu re a nd i t will obvi ously be p ro

,

p orti ona l to th e c once nt ra ti on tempe ra tu re a nd othe r

, ,

c onditi ons p rev a ili ng If equ a l p rop orti ons of th e s a me

.

subst a nce a re i n s oluti on on opp osite sides o f a mem

b ra ne or i n di ffe re nt p a rts of a s oluti on the osm otic
, ,

p ressu re will be equ a l ; bu t if th e p rop orti ons a re u n

equ a l th e osm otic p ressu re i nth e m ore c oncent ra ted p a rt
of th e s oluti onwill be g re a te r ; a n d if the e ncl osi n g mem
b ra ne is pe rme a ble th e m oveme nts th ro ugh i t will result
,

i nequ a lizi ng the p ressu re by th e e nt ra nce o f m ore of th e

diss olved subst a nce i nt o th e les s c once nt ra ted a nd m ore
o f the w a te r i n t o the m ore c once nt ra ted p a rts Osm otic .

p ressu re , th e refore dete rmi nes th e di recti on a nd ra te of

,

m ovement of diss olved subst a nces a nd of the s olve nt

itself if the i nte rve ni ng memb ra nes a re pe rme a ble .

LI V I N G CELL S AR E OS M OTI C S Y S TE M S t a ki ng i n a n d ,

givi ng Off i n s oluti on i n w a te r s olids liquids a nd g a ses , ,

.

I nt a ke we th ink of a s occu rri ng p ri ncip a lly i nroots a nd

le a ves i n l a nd pl a nts but th e outg o is not less ce rt a i n
,
.

R oo ts a bs orb w a te r o xy ge n a nd mi n
,
e ra l s a lts from the
,

s oil ; le a ves a bs orb CO a nd 0 from th e a i r

2 2 .

T H E STR U CTU RE OF R OOT S Th e root of the ordina ry

.
54 M O V E M E N T O F M AT ER IAL S I N P LAN T S

l a nd pl a nt se rving the d ouble pu rp ose of a bs orp ti ona nd

,

o f mech a n ic a l supp ort a nd livi ng a nd g rowi ng i na m ore

,

or less c omp a c t a n d resist a nt medium p ossesses p eculi a ri ,

ties of st ructu re cl osely rel a ted to the c onditi ons i nwhich

it lives A roo t is a cyli nd ric a l org a n b ra nched it ma y
.
, ,

be with a c onic a l ra the r th a na blu nt tip which is c a pp ed

, ,

or c o ve red wi th cell s m ore o r less c on ti nu a lly w orn off .

This thimble sh a ped ca p (Fig 1 0 ) p ro tecti ng the te nde r

-
.

tissues formi ng th e g rowing p oi nt of the tip is me rely a

, ,

me a ns of meet ing th e mec ha nic a l we a r i nesc a p a ble whe n

a p oi n t is pushed f orw a rd th rough a b ra sive m a te ri a l .

Immedi a tely behi nd th e thimble sh a ped ca p a nd the -

p oint a t which new cells a re f ormed to repl a ce th ose

FI G 1 0
. Lo ngi tu di n
. a l se cti o no f R o o t- ti p , S h o wi ng ca p , encl osi ng
an d p ro te c ti ng th e growi ng poi nt, be hi nd whi ch is th e re gi onof grow th
i nl e ng th Inthis regi o nthe tiss u es d iff e renti a te
. .

which a re w ornO ff by a b ra s i on the re is wh a t is k now na s

,

th e growi ng regi on only two or th e re millimete rs i n

,

le ngth i nwh ich g rowth i nle ngth t a kes pl a ce F rom this

,
.

a rra n gem e nt it is evide nt th a t only a ve ry sh ort segme nt

o f th e root is pushed th rough th e resist a n t s oil Fu rthe r .

b a ck f rom th e tip whe re th e cells h a ve ce a sed to g row i n

,

le ng th th e epide rm a l ce lls b ra nch o ut f orm i ng a bs orbi ng

, ,

h a i rs (Fig which a pply themselves o fte n ve ry

.
,

cl osely to th e su rf a ce of the p a rticles o f s o il with which

,
 R OO T S T R U C TU R E 55

they c ome i nc ont a ct These h a i rs with thi nw a lls li ned

.
,

with p rot opl a sm a nd c ont a ini ng c onside ra bl e v olumes o f

cell Sa p b ri ng th e s oil s oluti ona nd th e cell sa p i nto di rect
-
,
-

c ont a ct thus est a blishi ng c onditi ons up onwh ich the i n

,

t a ke or a bs orpti onof w a te r a nd of mi ne ra l s a lts f rom the

s oil a re a bsolutely depe nde nt S till fu rthe r b a ck f rom
.

th e tip of th e root is the p a rt i nwhich g rowth i ndi a mete r

ma y t a ke pl a ce a p a rt which exte nds to th e b a se of
,

th e stem .

FI G .
— C ross s ecti on o f Y o ung R oo t i n th e re gio n
-
, roo t

h a irs wh ere grow th i nlength h a s cea se d a nd a bs orp ti onis m os t ra pid

,
.

t du ct i n th e xyl em or w oo d ; p h si ev e t ub e i n phl oem o r b a s t ; p

, , ,

p eri cy cl e whi ch b o u nds th e v a scul a r tissu es ; en end o dermis whi ch

, , ,

b ou nds cortex 0 ; e epid ermis m a ny of th e cells of whi ch p u t ou t

, , ,

e x ten si ons i n h a irs h ,

.

M E CH A N I CA L R E S U LTS OF R OOT STR U CTU R E Whe na .

p a rt O f a roo t g rows i ndi a mete r th e root h a i rs c ove ri ng

,
-

its su rf a ce a re c rushed a g a i nst th e su rrou ndi ng s oil I n .

c re a si ng di a mete r is ordi na rily a cc omp a nied by i nc re a si ng

st re ngth Thus the mech a nic a l supp ort whi ch th e pl a nt
.

req ui res is supplied The p ositi on o f th e z one of root

.
56 M O V E M E N T O F M AT ER IAL S I N PL AN T S
h a i rs only a sh ort dist a nce b a ck o f th e tip is a lm ost
, ,

d i rec tly u nde r wh a t ma y be c a lled th e e a ves of th e d ome

o f fo li a ge .The re results f ro m this p o siti o n th e c onta ct
o f th e a bs o rbi n g root h a i rs wi th th o se p a rticles of s oil
-

m o st likely to h old w a te r on th ei r su rf a ces Th e a b .

so rp ti on O f w a te r a nd o f m a tte r d iss olved in th e s oil

w a ter t a ki ng pl a ce th ro ugh th e h a i rs i n c onta c t wi th
,

s oil p a rticles must be equ a lled by th e t ra nsfe r o f these

,
.

subs ta nces f ro m cell to cell u ntil they a re u sed .

M OVE M EN T O F W ATER AN D D I S S O LV ED S U B S TA N CES .

Th e m oveme nt O f w a t e r a nd o f s olutes f rom th e roo t

h a i rs to a nd th ro ugh a dj a ce nt cells fi na lly c a rries th em
t o th e v a s cul a r or c on duc ti ng ti ssues Th ese with th e .
,

mech a nic a l ly st re ng the ni ng fib ro us tissues f orm the c ore ,

o f th e ro o t I nto th e cells a nd vessels th e se c a ll ed

.
,
-

t ra cheid s a nd ducts o f th e w ood th e a bs orbed w a te r a nd

,

s olu tes a re disch a rged by th e a dj oi ni ng cells Th ro ugh .

th e vessels o f th e w ood o f th e root s th e s oluti onis c a rried

to th e simil a r t issues f orm in g th e w ood o f stem b ra nches ,

an d le a ves Th us th e re is a c ond uc ti ng syst em for th e

.

tra n sfe r o f w a te r a nd ra w m a te ri a ls wh ich is c ont inu ous

f ro m ti p to ti p o f th e h ighe r l a nd pl a n t Th rough th is .

th e s olu t i onis m o ved by me a n s not yet pe rfec tly u nde r

s tood but wi th a speed wh ich except i nseve re c onditions
, , ,

insu res th e pl a nt a g a i nst wilti ng .

T H E S TR U CTURE O F S T E M S The stem of th e ordi na ry

.

l a nd pl a nt gives mech a nic a l supp ort a n d fu rn ish es th e

p a th s a l o ng wh ich th e subs ta nces a bsorbed by th e roo t
re a ch th e le a ves i n wh ich f ood m a n uf a ct u re i s m a inly
c a rried o n S t re ng the ni ng a nd c ond uc ti ng tiss ues the re
.
,

fore m a ke up th e bulk o f th e stems b ra nch es twigs a nd

. .
,

eve nst a lks o f le a ves fl o we rs a nd f ruit s S tems li vi ng

, . .
,

an d g rowi n g i nth e a i r a re only feebly supp orted by it

,

an d he n ce devel op mech a n ic a l tissues q uite d i ffe re nt from

th ose o f roo ts Th e pl a ce a nd m ode of growth of s te ms
.

is a l so c orrespondi ngly d i ff e re nt S tems a re f o rmed o f .

se gme nts c a ll ed j o i n ts o r n odes a n d in, tem odes a n d ,

 T HE A S CE NT O F S AP 57

growth i nle ngth which t a kes pl a ce only wh e nth e j o ints

,

a re ve ry y o ung occu rs i nseve ra l j oi nts a nd not a t the tip

,

a l one a s i nroots The le a ves a nd b ra nch es sp ri ng f rom

,
.

th e nodes Th e i nte rna l st ructu re (Fig 1 2) c orresp onds

. .

with this exte rna l a rra ngement the re bei ng g rowi ng con
, ,

ducting a nd mech a nic a lly st re ngthe ni ng tissues i n the

,

stem which pe rsist f rom ye a r to ye a r Th ese a re u nited .

i nt o a cylinde r su rrou ndi ng the ce nt ra l pith a nd e nci rcled

by the p rotective tissues epide rmis c ork etc c oll ectively
, , ,
.
,

termed the ou te r b a rk or c ortex Th e g re a te r p a rt O f

.

FI G 1 2
.
ss
Cro s ec
. t i o n - o f a Tulip T re e B ra n ch f o ur y,e a rs O l d ,

sh owi ng th e w oo d to cons is t of b o th condu cti ng tissu es (du cts etc ) ,

.

an d m h i
ec a nca lly s u pp o r t i ng t issu e s (fib e rs e t
,
c ) su r r
.o u nded by
th e b a rk .

th e st re ngth e ni ng a nd v a scul a r tissues a re w ood wh ich ,

is c omp osed of fibe rs a nd o f tubes th ugh which w te

ro a r

an d diss olved ra w m a te ri a ls m ove upw ds f m ts

a r ro r o o

to le a ves .

A
T H E S CE N T OF S AP — The na me
.
s a p is give n to th e

w a te r a nd ra w m a te ri a ls c a r ied up th e stem th ugh

r r o

the w ood The mech a ni cs of th e m oveme nt a re not yet

.

completely u n ,
n
de rst ood bu t it is evide t th t if w te
a a r

is l ost by ev a p ora ti on from th e le vesa a n d is c mbi ed

o n ,
58 MOV E M E N T OF M AT E R I AL S I N P LANT S

i n th e chl orophy ll c onta i ni ng cell s to m a ke food a nd if

-
,

th e n it ra tes a nd o the r diss olved subst a nces a re a ls o used

i n food m a nuf a c tu re they will be p ulled upw a rds by a
,

ce rt a i n a m o unt o f f orce In th e ducts a nd t ra ch eids

.

themselves c on ,
t a ini ng l onge r or sh orte r c olum ns o f w a te r ,

te n si ons a rise or exist wh ich li ft the c olum ns o f w a te r .

W a te r a nd diss olved subst a nces ma y be p ressed f rom a d

j oi ni ng cells i nt o th e d ucts a nd t ra cheids i n th e roo ts .

Thus seve ra l or m a ny forces a re c ombi ned i n b ri nging

a b o u t the lif ti n g of g re a t qu a ntities o f w a te r U nde r .

ordi n a ry c on diti ons o f na tu re e nough w a te r is lifted to

the top of the t a llest t rees to supply th ei r t opm ost le a ves .

In the c a se o f l a rge t rees th e q u a ntity of w a te r ma y

a m o un t to seve ra l t ons p er ye a r .

T H E STR U CTU RE OF L EA V ES The le a ves of th e usu a l

.

l a nd pl a nt a re broa d fi a t th in exp a nses of g ree n tissue

'

, ,
.

Th e s tructu re of le a ves (Fig 1 3 ) is by no me a ns so simple

.

FIG 1 3
. . C ross- se c ti o no f Le a f
(ca s te r be a n) sh o wi ng th e u pper
an d l o w e r e pid ermis pe rf ora te d by gu a rd e d O pe n
, i ngs (s to m a ta ) a nd
en cl o si ng the m es o phyll co nsis ti ng of a p a lis a d e l a y er of co mp a ct
,

ce lls a n d th e sp o n gy po rti on be lo w .

as thei r form They a re c ompletely c ove red by a si ngle

.

l a ye r of epide rm a l cells wh ich a re fla t ra the r th i n with , ,

o ute r w a lls wh ich a re w a te r p roo fed th ough th e i nne r

-
,

an d side w a lls a re f reely pe rm e a ble to w a te r Th is single .

 L E AV E S AS A B S O R B ING O R GAN S 59

l a ye r of cells f ormi ng the epide rmis is i nte rrupted a t i n

terv a l s with p ores k n o w n a s s to ma ta which h a ve
gu a rd
ce lls th a t ope n a n d cl ose G e ne ra lly spe a ki ng th e re a re
.
,

m ore st om a t a i nthe epide rmis th a t c ove rs the u nde r side

o f le a ves th ough pe nd a nt le a ves or ve rtic a lly e rect
, ,

le a ves h a vi ng equ a l exp osu re to ligh t on b oth sides

, ,

h a ve a m ore ne a rly equ a l numbe r of st om a t a onth e two

sides The fla t thi n exp a nse which we c a ll a le a f h a s
”
.
,

a f ra mew ork o f vei ns or ne rves which a re the “

supp orting a nd c onducti ng tissues Th ese a re the con .

ti n u a ti on a n d extensi on with out i nte rrupti on i n the

le a ves of the supp orti ng a nd c onducti ng ti ssues o f th e
stem b ra nch es a nd twigs
, , .

s
B etween the epide rmis a n d the fra mew ork is th e
mesop hy ll th e cells of which c ont a i n chl orophyll Th e
, .

uppe rm ost l a ye r of mes ophyll cells ma y be m ore or less

b rick sh a ped with r el a tively sm a ll sp a ces betwee n them
-
.

This l a ye r is c a lled the p a lis a de l a ye r Betwee n it a nd .

th e epide rmis c ove ring the l owe r su rf a ce the re is a m ore

or less sp on gy tissue th e cells O f wh ich c ont a i n m a ny
,

chl orophyll g ra ins a nd betwee n th e cells a re a i r sp a ces

,
-

which a re Conti nu ou s th rough ou t th e m a ss of th e le a f .

I nt o a nd th rough a nd ou t of these sp a ces c a rb ondi oxid ,

o xyge n a n d w a te r v a p or diff use

,
I t is f rom these i nte r .

cellul a r a i r— sp a ces th a t c a rb on di oxid a nd oxyge n e nte r

th e cell diss olvi ng i n th e w a te r o f the cell w a lls a nd
,
-

diffusi ng a t ra tes v a ryi ng with the diff e re nce i n c onee n

tra ti on of these g a ses i nside a nd o utside the cell .

LEA V ES As A B S OR B I N G OR GA N S — Le a ves the n a re , ,

th i n a bs orbing org a ns sp re a d ou t i n th e a i r They a b .

s orb b oth ligh t a nd ce rt a i n g a se o us c onstitue nts o f the

a tm o sphe re Bec a use O f the pigme nts wh i ch they con
.

t a i nth ey a bs orb only ce rt a i nrays o f ligh t They a bs orb .

on ly ce rt a i n c onstitue nts Of the a tm osphe re na mely , ,

c a rb on di oxid a nd oxyge n bec a use th ey use only these ,

two . They a bs orb CO only du ri ng th e h ou rs of ph ot o

2

synth etic a ctivity du ri ng the h ou rs o f w a rm d a ylight

,
.
60 M O V E M E N T O F M AT E R I AL S I N PL AN T S

They a bs orb O cont i n u o usly for it is i nv olved i n th ose

2 ,

met a b olic ch a nges o f v a ri o us f oods which rele a se the

e ne rgy st ored i n them d u ring thei r m a nuf a ctu re The .

a bs orpti o no f C O a n d O by cell s f rom the i n

2 2 te rcellul a r
sp a ces inthe mes ophyll o f le a ves t a kes pl a ce inth e s a me
m a nne r a s the a bs orpti on of w a te r a nd mi ne ra l m a tte r
by ro o t h a i rs f rom the s o il The a i r of the inte rcellul a r
-
.

sp a ces is c onti nu o us th ro ugh th e ope n st om a t a with

, ,

th e u nc o nfi ned a i r o utside Th rough out th is c onti nu o us

.

m a ss di ff usi onw i ll te nd to m a int a i n u niform c once nt ra

ti on o f th e c onstitue nts Th rough the wet memb ra nes
.

b o u nd ing cells c ont a i ni ng chl orophyll a nd a bs orbi ng ligh t

CO will e nte r diss olvi ng i nth e w a te r O f the memb ra nes
2 , ,

an d will be c ombi ned i n ph o t o sy nthesis to f orm sug a r .

Th us du ri ng the h ou rs o f d a yligh t C 0 will a lw a ys te nd 2

to be c ombi n ed i n chl orophyll c ont a i ning cell s wh ich -

,

the refore will c onti nue to a bs orb i t f rom a dj a ce nt i nte r

,

cellul a r sp a ces S imil a rly 0 w ill be t a ke ni na s ra pidly

.
, 2

a s it is used .

L EA V E S As ORGA N S O F GAS E X CH A N GE —C orresp ond .

in g to the i n t a ke of CO a nd O by th e livi ng cells Of

2 2

le a ves is the disch a rge of O a nd CO f rom livi ng cells 2 2

int o th e inte rcellul a r sp a ces wh e neve r th e p ressu res o f

these diss olved g a ses with i n th e cells exceed th ei r p res
su res i n a i r wh ich fills the i nte rcellul a r sp a ces Thus .

du ri ng the h ou rs o f ph ot o sy nthetic a ctivi ty th e v olume

o f O give no ff f ro m g ree nle a ves is equ a l to th e v olume
2

of CO a bs orbed
2 Bu t d u ri ng the h o u rs wh e n ph ot o
.

s ynth esis is no t t a ki ng pl a ce the livi ng cells a bs orb O

usi ng it i n dest ructive met a b olism (see C h a p V I I I ) .
,

an d give o ff C 0 He nce le a ves a re c a lled org a ns of

2.

g a s exch a nge .

EVAP OR AT I O N will t a ke pl a ce i n evi t a bly a nd its ra te ,

will depe nd up on th e d ry ness o f th e a i r The l oss o f .

w a te r f rom th e b ody o f th e pl a nt h a s u nf ortu na tely bee n

c a lled tra nspi ra ti o n bu t th e re is no di ffe re nce betwee n
,

t ra nspi ra ti ona nd ev a p ora ti oni npl a nts excep t th a t pl a nts

 R OO T S AS E X CR E TIN G O R GAN S 61

r egul a te thei r t ra nspi ra ti onwithi nce rt a i nlimits They .

do so by v a ri ous me a ns the ch ief O f wh ich a re th e w a te r

,

p roofi ng Of th e o ute r w a lls of th e cells O f the epide rmi s ,

an d the cl o si ng of thei r st om a t a by th e gu a rd cells .

R OOT S AS EX CR T N G
E I O R GA S
N — From th e f oreg oi ng ,

on e will c orrectly i n fe r th a t roo ts not only a b sorb but

th a t they a ls o get ri d of wh a teve r di sssol ved subst a nces
th ey ma y h old i nexcess a s c omp a red with the s oil s olu
,

ti on p rovided th e cell memb ra nes a re pe rme a ble to

,

these subst a nces Th e roots di sch a rge c a rb on di oxid

.
,

ce rt a i n org a nic a cids a nd v a ri ou s oth e r subst a nces

'

.
,

Some of these a ff ect th e s olubility o f s ome of the s oil

c onstitue nts othe r exc ret a a re p oi sono us a nd if a ll owed
,

to a ccumul a t e by n eglect of suit a ble a lte rna ti on of

c rops will imp a i r th e p roductiveness of the s oil
, .

Th e s a me p h y si ca l p rin
‘

SE CR ETI ON AN D EX CRETI O N .

ci p l es a pply to the sec reti on o f sug a r s th e exc reti on of ,

w a ste subst a nces a nd th e disch a rge o f liquid w a te r f rom

,

pl a nts a s u nde rlie th e a bs orp ti onO f diss olved subst a nces .

Th us we h a ve gl a nds receiving f ro m th e o the r p a rts of

th e pl a nt b ody by diff usi on a nd o sm osis ce rt a i n sub
st a nces wh ich they ma y so t ra nsform th a t they ca na ccu
mul a te a nd r et a i nth em Th us it is c onceiv a ble th a t the
.

gl a nds onth e su rf a ce o f a ge ra nium le a f receive f rom the

le a f sug a rs of one or m ore ki nds from which they m a ke
an d sec rete the v ol a til e oi l which gives th e ch a ra cte ristic
fra gra nce to the foli a ge of this pl a nt Th e re a re gl a ndu .

l a r h a i rs which exc rete w a te r p resum a bly bec a use the,

pe rme a bility O f th ei r o ute r cell memb ra nes to w a te r ex -

ceeds the pe rme a bility o f a dj a ce n t memb ra nes so th a t ,

the w a ter will esc a p e th rough the w a lls which a re m ost

pe rme a ble to it S uch h a i rs o ccu r on th e le a ves of
.

be a ns .

GUTTATI ON AN D B LEEDI N G — Whe n the p ressu re O f

w a te r i n cells exceeds th e a bility of th e cell memb ra nes
to h old it w a te r will be squeezed ou t
,
This p ressu re of .

w a te r or cell sa p is know na s tu rgor A t ce rt a intimes

,
-
,
.
62 M O V E M E N T O F M AT E R IAL S I N P LAN T S

a nd u nde r ce rt a in c onditi ons it ma y be ve ry high I t .

tends to be so a t nigh t whe nev a p ora ti onf rom the le a ves

sl ows d ow nor ce a ses wh ile the a bs o rpti on O f w a te r by
,

the roots c onti nues Th us w a te r a c cumul a tes i nthe tis

.

sues oi the pl a nt M uch of wh a t we c a l l dew on th e

.

g ra ss is a c onseque nce o f the squeezing ou t of w a te r i n

-

this wa y Th is is te rmed gu tta ti on Bleedi ng f rom

. .

w ounds is delibe ra tely b rough t a b o ut by ma ninhis ta p

ping of m a ple t rees i n th e sp ri ng to m a ke sy rup
an d sug a r ; a nd of ce rt a i n c a cti a nd simil a r dese rt
pl a nts f rom which w a te r a nd s a ps a re c ollected wh ich a re
used eithe r with or with ou t fe rme nt a ti on Th us we see
.

th a t the p rocesses of a bs orp ti on sec reti on a nd exc reti on

, ,

a re d ue to th e k in etic e ne rgy o f the m olecules of dis

s olved subst a nces a nd of their c omm on s olve nt w a te r , .
 C HAPTER VII

T H E S T O R AG E A ND DIG E S TI O N OF F O OD S
I N P LANT S

S TORA GE . We h a ve see nth a t the f oods m a de i ng ree n

le a ves a nd i n o the r p a rts of the pl a nt ma y be used a t
on ce for th e c onst ructi on a nd ope ra ti on O f th e pl a nt
b ody or they ma y b e st ored This st ora ge ma y occu r
,
.

wh e re they a re m a de or it ma y t a ke pl a ce i nothe r p a rts

,

o f the pl a n t b ody Inth e l a tte r c a se th e f oods must be

.
,

rem oved from th e pl a ces o f m a nufa ctu re a nd this ,

m oveme nt like th e a bs orpti on of ra w m a te ri a l a nd the

,

sec reti on a nd exc reti on of el a b ora ted c omp ou nds ca n ,

t a ke pl a ce only i n soluti on i n w a te r .

T H E CO M M O N P L A CE S OF ST OR AGE I f ph ot osynth esis

.

is m ore ra pid th a n the di ffusi on o f sug a r f rom the cells

i nwhich i t is m a d e i nth e g ree nle a f th e sug a r will a ccu
,

mu l a te i n th e cells i nwhich it is f ormed I t will ge ner .

a lly b e c on ve rted i nt o s t a rch whi ch I s I ns oluble i nw a te r

,

an d occu rs i n th e f orm o f g ra nules f ormed i n th e g ree n

ch rom a t oph ores Obse rv a ti on S h ows th a t du ring the
.
,

h ou rs of d a yligh t such a ccumul a ti on ordi na rily o ccu rs

n
,

i nmild or w a rm we a th e r a nd th a t du ri g the nigh t the

,

cell s a re emp tied a g a i n Th us wh ile th e le a ves of ora nge

.
,

t rees ma y be ve ry full of st a rch a t su nset they a re gene r ,

a lly spe a ki n g c ompletely emp tied O f it by su nrise a nd

‘

, ,

the sug a r wh ich h a s bee n rem oved from the le a ves ma y

be st ored elsewh e re a s st a rch or i no the r f orms or it ma y ,

be used i nthe m a nu fa ctu re of m ore c omplex c omp ou nds

in o the r p a rts O f th e pl a nt .

T H E P L A CE S OF M OR E P ER M A N E N T STOR AGE v a ry with

the le ngth of life of th e pl a nt Thus we find inpl a nts .
,

63
64 S T O R AG E AN D DIG E S TI O N OF F OOD S
which a re a nnu a ls th a t th e f ood a ccumul a ted i none pl a nt
is beque a thed i nm ore o r less pe rm a ne nt f o rm to its o ff
S p ri ng being s tored i n seed In th e c a se o f pl a nts
,
.

which g row one se a s o n a nd seed th e nex t a s do ,

beets oni ons etc the f ood m a d e d u ri ng the fi rst g rowing

, ,
.
,

se a s onis st ored i nth e roo ts a nd o the r u nde rg ro u nd p a rts ,

an d is used the f oll o wi ng ye a r inthe p roducti ono f fl owe rs

an d f rui t Inthe c a se o f pe re n
. ni a ls th e st ora ge O f f ood
ma y t a ke pl a ce i nth e b a rk i nth e roots i nthe seeds a nd , ,

f ruits inthe pith a nd i nv a ri ous o th e r p a rts of th e pl a nt

, ,

b ody Th us we fi nd sug a r st ored i n sug a r beets a nd

.

on i ons The sug a r wh ich is fou nd i nth e sa p o f m a ples

.
,

g ra pe vi nes a nd o th e r pe re nni a ls c omes f rom the dige sted

, ,

s to res of st a rch dep osited i n v a ri ous p a rts of th e pl a nt

b ody du ri ng the p recedi ng se a son S t a rch a ccumul a tes .

i nthe tube rs o f th e p ot a t o u nde rg rou n d a nd is p rese nt , ,

i nc orna nd i nthe o th e r g ra i ns .

STAR CH ma y be c onceived a s a di rec t de riv a tive o f

sug a r obt a i ned by withd ra wi ng a m olecule O f w a te r f ro m
,

the m olecule o f sug a r Bei ng i ns oluble it a ccumul a tes.

‘
,

o r is dep o sited i n speci a l cells a n

,
d i n speci a l o rg a ns o f
the cell know n a s a my lop la s ts a nd c onstitutes one O f ,

the m ost c omp a c t a nd ec onomic a l form s i nwh ich no n

nit roge no us food is a ccumul a ted a nd st ored .

OT H ER S TO R ED F O ODs I n th e tube rs of th e d a h l i a
.
-

the c a rb ohyd ra te f ood a ccumul a tes i n s oluble f o rm but ,

no t a s sug a r rem a i ni ng i ns oluti oni nth e f orm of i nu li n

, ,

a subst a nce of the s a me p rop orti ona l c o mp ositi on a s

s ta rch a nd cellul ose I t ma y be p recipit a ted by d roppi ng
.

pieces O f d a hli a tube r i nt o st ro ng a lc oh ol or st rong glyc

e ri ne i n wh ich c a se i t w ill c ome d ow n i n th e fo rm o f
,

sp h aaro c ryst a ls
-
In iv ory p a lm n . uts f rom wh ich th e -
,

bu tt ons O f me ns cl o th es a re ve ry co mm o nly m a d e th e
’
,

c a rb ohyd ra te food o f the pl a nt is st ored i n the form o f

cellul ose which is a pplied to the cell w a lls u ntil they -

bec ome e norm o usly thi ck ; a t the s a me time th e cellul o se

is so c omp a ct a nd fi rm th a t it is ve ry h a rd Inv a ri ous .
 M O VING M AN U FA C T U R E D F O O D S 65

o the r nu ts a nd ing ra ins Olives c a st or be a ns one fi nds

, , , ,

o ils st ored in g r e a te r or less qu a ntities Th ese a re non .

nit roge nous foods wh ich h oweve r si nce they a re i n , ,

s oluble i nw a te r a re j ust a s secu rely st ored a nd a re j ust

,

a s imm o v a ble a s st a rch or cellul o se S imil a rly p ro tei ns .

,

o f v a ri o us s orts i ns oluble i nw a te r a re dep osited i nbe a ns ,

pe a s a nd m a ny o th e r seeds a nd i n the stems a nd roo ts

, , ,

from which they ma y be rem oved only by digesti on .

M EA N S O F M OV I N G M A N UFA CTU RED F OODS — F oods ,

like th e f ood m a te ri a ls ca n be m oved f rom cell to cell

,

on ly i ns oluti on; but i ns oluti onthe nit roge nous a nd non

nit rogenous f oods m ove re a dily th ro ugh o ut th e cell a nd ,

f rom cell to cell th rough p e rme a ble memb ra nes i n

, ,

a cc ord a n ce with th e p ri nciples discussed i nthe p reced i ng

ch a pter (V I ) .

T H E P AT H S OF M OV E M E N T — Th e f oods a re m oved .

(t ra nsl oc a ted ) m a i nly a l ong ce rt a i n p a ths These p a th s .

ma y v a ry f rom se a s on to se a s on a s for ex a mple i n th e ,

sug a r m a ple In this t ree th e sug a r m oves d ow n from

.

the le a ves to th e pl a ces of st ora ge th rough the i nne r b a rk

th rou gh ou t th e summe r ; but i n th e sp ri ng it is m a i n ly
th rough th e w ood th a t th e sug a r c ont a i ni ng sa p is m oved -

f rom th e pl a ces o f st ora ge upw a rds to the swelli ng buds .

One ma y O fte nfi nd th a t th e i nne r b a rk O f t rees or sh rubs

t a stes sweet whe re a s the re is no such fl a v or i nth e w ood
,
.

One ma y s ometimes no tice on pulling a p a rt a st a lk of ,

g rowi ng whe a t oa ts or b a rley i n th e a fte rnoon o f a

, , ,

w a rm su nny da y i ne a rly summe r th a t th e j oi nts a re b oth ,

sweet a nd te nde r In the e a rly m orni ng before th e

.
,

le a ves h a ve m a de a ny f ood a nd whe n the g ra i n is ,

ne a rly ripe the re w ould be no sweet ne ss i n the st a lks

, ,

bec a use the re w o uld be no sug a r i n t ra nsit Th e i nne r .

b a rk or phl oem of th e v a scul a r bu ndles is the c onduct

, , ,

in g tissue th ro ugh which m o st o f th e sug a r is m oved f rom

pl a ces O f m a nuf a ctu re to pl a ces o f st ora ge ; whe re a s th e

wood ma y be the p a th of m oveme nt of sug a r etc from ,
.
,

th e pl a ces o f wi nte r st ora ge to th e regi ons of

sp r ing g rowth .
66 S T O R AG E AN D DIG E S TI O N OF F OO D S
D IGES T IO N The rem ov a l of s olid f ood f rom the pl a ce
.

o f temp o ra ry o r pe rm a n e nt st o ra ge for ex a mple th e re , ,

m ov a l o f a ccumul a ted st a rch f rom a p o ta t o le a f to a

p o ta to tube r o r f ro m the tube r to the g rowi ng sp rout
, ,

imp lies th e c onve rsi o n o f st a rch which is i ns oluble i n ,

w a te r i nt o s o me o the r subst a nce wh ich diss olvi ng i n

, ,

w a te r the reby bec omes m ov a ble S uch chemic a l ch a nge

, .

f rom i ns oluble to s oluble m a te ri a l is k now na s digesti on .

In s ome c a ses digesti on me rely i nv olves th e c on ve rsi on

of on e s oluble s ubst a n ce i nt o a noth e r .

T H E M EA N S O F D I GES T I O N S t a rch ma y be c on
. ve rted
i nto sug a r i n the l a b ora t ory by t re a tme nt wi th d ilute
a cid dilu te a lk a li a n
, d e nzyms,
Bu t th e a cti onof d ilute
.

a cids a n d dilut e a lk a lis if it is to be ra pid must t a ke

'

, ,

pl a ce a t tempe ra tu res h igh e r th a n th ose wh ich p rev a il

i nthe livi n g b ody Th e p rocess o f digesti on th e refore
.
, ,

mu st be ca rried on m ore ra pidly th a n dilute a cids a nd

a lk a lis a cti ng a t b ody tempe ra tu res m a ke p ossible Th e
, ,
.

m e a ns empl oyed by a nim a ls a nd pl a nts i ndigesti ng thei r

f o od a re ce rt a i nsubst a nces k now na s enz y ms Th ese a re .

c omplex c o mp ou nds h a vi ng v a ri ou s c omm on ch a ra cte rs

th e m ost sig nifi ca nt O f wh ich is th ei r c a p a city to ch a nge
th e ra te O f ch emic a l re a cti ons wh i ch migh t o th e rwise go
o n with o ut th em Th ey a re the ref ore c a t a lyze rs
.

CATA LYZ ER S Ch a rc oa l pl a ti num sp onge a n

.
,
d v a ri ous ,

o the r subst a n ces h a ve th e rem a rk a ble q u a lity o f ch a ng

ing th e ra te o f ch emi ca l re a cti o ns t a ki ng pl a ce i n thei r
p rese nce S ome c a t a lyze rs a re empl oyed i n indust ri a l
.

p ro cesses fo r ex a mple th e u se o f pl a ti num sp onge i nth e

,

t re a tme nt of a mm o ni a i nmu niti onm a nu f a c tu re a nd o f ,

fi nely d ivid ed nickel i n c onve rti ng o ils i nt o s olid f a ts .

Bu t the c a t a lyze rs O f m o st imp ort a nce to ma na re th ose

en zg/ m s c o nce rn ed i nthe physi o l o gic a l p ro cesses i n livi ng
o rg a n isms C a t a lyze rs p rodu ce e ff ects o u t o f p rop orti on
.

to thei r o wnq u a n tities O fte na ve ry sm a ll q u a ntity o f

.

a suit a ble e n zym di a st a se fo r ex a mple will c onve rt a

, , ,

ve ry l a rge a m ou nt o f st a rch i nt o sug a r i f th e tempe ra tu re

 E N ZY M S 7

a nd the r c onditi ons a re f a v ora ble a nd time be give n

o

f or th e ope ra ti on Th e st a teme nt ma y s a fely be m a de

.

th a t we do not now k now of a ny chemic a l ch a nge t a ki ng

pl a ce i n pl a nt cells a t le a st i n which e nzyms a re not
, ,

i nv olved Th e one excep ti on to this ge ne ra l rule is a n

.

excep ti on due to ou r own ig nora nce ra the r th a n to a ny

knowledge to th e c ont ra ry na mely the se ries O f chemic a l , ,

re a cti on s t a ki ng pl ace i nph ot osynthesis .

E N ZY M S G e ne ra lly spe a ki ng these subst a nces occu r

.
,

insuch sm a ll qu a ntities th a t th ey esc a p e ch emic a l a n a ly

sis I t is believed h oweve r th a t they a re c oll oid a l a nd

'

.
, ,

p o ssibly p rotei ns Th ey a re s oluble a nd h e nce th ey a re

'
.
,

diss olved i n th e cells p resum a bly i n c oll oid a l s oluti on ,

.

Th ey a ct a t ordi na ry tempe ra tu res th ose a t wh ich cell ,

a ctivities a re p o ssibl e M ost of them a re dec omp os a ble .

by he a t .

CLA S S I F I CAT I O N O F E N ZY M S Th e e nzyms a re so .

-

nume rous th a t cl a ssific a ti on is necessa ry Th ey a re di .

v i ded a cc ordi n g to th ei r physi ol ogic a l or chemic a l a c

ti v i ti es thus : ,

I The hydrolyz i ng enz yms (Hydrola s es ) : th ose which

.

a cc omplish th e a dditi on o f w a te r with a subseque nt

splitting of the m olecules a cted up on Of th ese we ma y .

me nti on (a ) the li p a s es e nzyms wh ich a ct up onthe f a ts ,

an d o ils st ored i n c a st or b e a ns B ra zil nu ts c orn hemp , , , ,

etc a nd S plit them i nt o glyce ri ne a nd f a tty a cids a s the

.
, ,

seeds sp ro u t ; (b ) the a my la ses e nzyms wh ich c onve rt ,

st a rch i nt o sug a r ; (c) o the r c a rb ohyd ra ses such a s cel ,

lu la se wh ich digests th e cellul ose p a rts o f cell w a lls ;

n
,

p ecti a s e which digests th e ceme n ,

ti ng m a te ri a l i n or .

betweenth e cell w a lls p a rticul a rly of f ruits which f orms , ,

th e jelly m a de from f ruits ; i nverta s e which f orms fer ,

menta bl e sug a rs e g gluc o se f rom c a ne sug a r ; (d )

,
. .
,

emu lsi na n d th e o the r e nzyms wh ich digest th e gluc osides

fou nd i na lm onds will ow b a rk etc to form gluc ose a nd , ,
.
,

v a ri ou s o the r p roducts ; (e ) the p ro tea s es wh ich a ct up on ,

p rotei na nd nucle o p ro tei ns c onve rting them i nt o a mi no

-
,
68 S T O R AG E AN D DIG E S TI O N OF F OO D S
a cids a nd othe r c om p ou nds s oluble a nd the refore us a ble .

I I Th e oxi di z i ng enz yms (oxi da ses ) which a re con

.

ce rned in th e i nt ra cellul a r oxid a ti ons which rele a se th e

e ne rgy req ui red for w ork by or i n livi ng org a nisms .

I I I The de a mi na ses which b re a k d ow n a mi no a cids

- -

n
.
,

a d simil a r subst a n ces to a mm oni a a nd c a rb ohyd ra tes .

I V Urea s es c a p a ble of b re a king u re a d ow n i nt o a m

.
,

m oni a (N H OH) a nd c a rb ondi oxid (C0 )

i (See Ch a p 2 . .

XX I V ) .

V Coa gu la ti ng enz y ms such a s c a use m ilk to cl o t

.
,
.

V I The fermenta ti on enz yms which c onve rt suga r

.
, ,

for ex a mple i nt o a lc o h ol a nd c a rb ondi oxid wh ich s o u r

, ,

milk sp oil butte r ra ise b re a d etc

, , ,
.

This cl a ssific a ti on i ncludes e nzyms not i n a ny wa y

c onnected with digesti oneithe r i na nim a ls (see Ch a p X ) .

or i npl a n ts but which c a rry onfe rme nt a ti ons a nd pu t re

,

f a c ti ons (see Ch a p XX I I I ) or a re c once rned i n m a i n

,

.
,

t a ini ng th e fe rtility O f s o il (see Ch a p XXI V ) or a re .

,

inv olved i n the met a b olic p rocesses wh ich supply th e

li vi ng cell s with e ne rgy We ma y me nti ons ome o f these .

e nzyms h e re i na dditi onto the desc rip ti ons i nth e pl a ces

,

refe rred to .

The oxi das es Th ese a re e nz yms o f v a ri ou s s orts

wh ich a ccele ra te th e oxid a ti o ns g oi ng on i n livi ng cells
an d fu rn ish i ng the cells with requi red e ne rgy Th ey a re .

c once rned i n th e physi ol ogic a l u tiliz a ti on a s s ou rces o f ,

e ne rgy of the c a rb ohyd ra tes espe ci a lly th e su g a rs They

, ,
.

ma y a ls o be c once rned i n fe rme nt a ti ons a nd they a re ,

u nd oubtedly c once rned i ns ome of the oxid a ti ons i nv olved

i n m a i nt a in i ng s oil fe rtility .

The de a mi n a s es t a ke p a rt i n b re a ki n
-
g d ow n a mi no
a cids a n d simil a r c omplex ni t ro ge n o u s c omp o u nd s to th e
a mm on i a a nd c a rb ohyd ra te ou t of whi ch th ey we re
f ormed m a ki ng th e c a rb ohyd ra te a v a il a ble for oxid a ti on
,

in th e cell by wh ich e ne rgy is rele a s ed for u se

,
.

The c oa gu la ti ng e n z yms a re rep rese n ted by re nnet Ob

t a in a bl e in c omme rce i n th e f orm o f j u nket t a blets
” ,

,
 G E R M INATI O N AND S P R ING G R OWT H 69

the c omm onuse o f which is th e cl o tti ng o f milk P ecta se .

is a n e nzym f ou nd i n pl a nts wh ich h a s th e p rope rty o f

,

c a usi ng the pectic subst a nces which ceme nt the cells

t ogeth e r to c oa gul a te a nd form a fi rm j elly l i ke sub sta nce
'

-
.

I t a cts i nc onj u ncti onwith th e s a lts of c a lcium .

The fermenta ti on en z yms empl o yed i n ra isi n

, g b re a d
d ough i n m a ki ng wi ne bee r a nd ch eese etc effect
chemic a l ch a nges i n c om
, , , , .
,

plex nonnit roge nous org a nic -

c omp ou nds resulti ng i nth e p roducti onof ga s a lc oh ol or , ,

o the r desi red subst a n ces I t is p rob a ble th a t fe rment a

.

ti one nzyms a re c once rned i n dise a se due to b a cteri a or

fu ngi p a ra sitic i n hum a n bei ngs .

G ER M I N ATI O N AN D S PR I N G GR O WT H Th e a cti on of .

e nzyms is illust ra ted i nrem a rk a ble deg ree i nth e ge rmina

ti onO f seeds a nd i nth e r esump ti onof g rowth which h a s
bee ni nte rrupted by c old or d rough t Th e sp routi ng a nd .

growi ng emb ry o a nd the devel op i ng seedli ng a re nou r

i sh ed by the f ood st ored in the seed wh ich is digested ,

an d m a de a v a il a ble by th e e nzyms wh ich w ork onst a rch ,

cellul ose f a ts a nd oil s a nd th e p ro tei ns M e a ntime

, ,
.
,

o the r e n zyms th e oxid a ses a re a ctive i n b ri ngi ng a b out

, ,

th e oxid a ti ons which supply th e y ou ng a nd g rowi ng pl a nt

with necess a ry e ne rgy S imil a rly sp ri ng g rowth con
.
, ,

sisti ng I n th e p roducti on of new f oli a ge i n el ong a ti on ,

an d thicke ni ng p e rh a ps i nbl oomi ng a nd fruiti ng a s well

, ,

is m a de p ossible by th e digesti on a nd u se by me a ns of ,

enzyms O f f ood st ored in i ns oluble f orm i np a rts of the

,

pl a nt O fte n rem o te from th e se a t of g rowth E nz ym .

a cti on the ref ore u n de rlies th e p roducti on st ora ge m ove , ,

me nt a nd use o f f ood i n pl a nts a nd a nim a ls Enzyms

,
.

a re p roduced by an d a re a lw a ys p rese nt i n the livi ng

, ,

p rot opl a sm which uses them in e ff ecti ng a nd regul a ti ng

,

chemic a l ch a nge .
 S E C T I ON 3
THE M A I N TEN A N C E OF LIFE
 C HAPTER VIII

T HE UTILI Z ATI O N OF F OO D

L I V I N G T H I N GS R E Q U I RE F O OD for seve ra l re a s ons In .

th e fi rst pl a ce thei r own b odies which a re m a d e up i n

, ,

p a rt Of livi ng p ro t opl a sm a nd i n p a rt of m a te ri a l which

th e livi ng p ro t opl a sm m a nuf a ctu res a nd l a ys d ow n must ,

h a ve a s o u rce of supply for m a ki ng p ro t opl a sm a nd these

othe r p ro ducts ; this s o u rce is i nth e f ood Inth e sec ond .

pl a ce livi ng thi ngs a re like m a ch i nes i n th a t they use

,

en ergy a n, d e ne rgy c a nno t c ome from nothi ng but must

be devel oped from s ome s o u rce which inth is c a se is a ls o ,

the f oo d Inrespect to its e ne rgy requi reme nt a livi ng

.

org a n ism is i nex a ctl y th e s a me situ a ti ona s a ny f a mili a r

non— livi ng m a chi ne whi ch is ope ra ted th ro ugh the bu rn
i ng o f fuel By wa y of illust ra ti on recu r to th e a u tomo
.
,

bile to wh ich r efe re nce wa s m a de i nCh a p te r I I I Aut o

, .

m obiles a re d rive nby th e expl o si ono f a na i r ga s mixtu re -

i nthe cyli nde rs Th e re a s on g a s oli ne ca n be used a s a

.

s ou rce of p owe r for a ut om obil es is th a t it h a s high e ne rgy

v a lue wh ich e ne rgy is rele a sed wh en th e g a s oli ne is
bu rned An a ut om obile is a device for c onve rti ng this
.

chemic a l e ne rgy O f bu rni ng i nt o th e mech a nic a l e ne rgy

of m oti on An y livi
. ng thi ng which d o es a ny ki nd o f
w ork mu st expe nd e ne rgy obt a i ned from f ood i n d o ing
its w ork bei ng i nthis respect i nex a ctly th e s a me situ a
,

ti ona s th e a ut om obile .

For th e m a nuf a ctu re of m a te ri a ls the n a nd a s a s ou rce , ,

of e n e rgy f oods a re necess a ry Wh ich eve r o f th ese e nds

,
.

they ma y se rve th ey h a ve to u nde rgo chemic a l ch a nges

,

i nthe pro cess Th ese ch a nges a re O f v a ri ous kinds s ome

.
,

of which will be de sc rib ed p rese n tly Th e te rm meta b .

73
7 41 T HE UTILI Z ATI O N O F FO O D

olis m is u sed by th e bi ol ogist a s a c onve nie nt ge ne ra l

te rm to i nclude a ll these chemic a l ch a nges of f oods a nd
o the r subs ta n ces m a d e o u t of them .

M ETAB O LI S M I N G EN ERA L Chemic a l p rocesses i nliv

.

in g thi ngs i nv o lve fi rst ,

a p,
rel im in a ry st a ge vi z the , .
,

t a ki ng i n o f the f ood m a te ri a ls ; sec ond th e met a b olism ,

i tself or th e t ra nsform a ti ons by wh ich th ese a re w ork ed

,

o ve r i n t o cell subst a nce or p rod ucts onth e one h a nd or

-
,

used up a s s ou rces of e ne rgy onth e o th e r ; a nd th i rd the ,

getting ri d o f such subst a nces or p ro ducts resulti ng f rom

th e chemic a l a cti ona s a re no t to be ret a i ned withi n th e
p ro t opl a s m M a te ri a ls to be g o tte n ri d o f a re eith e r
.

m a nu f a ctu red thi ngs like e nzyms wh ich b o th pl a nt a nd ,

an im a l cells form a nd sec rete or w a ste p roducts like th e

, ,

c a rbondi oxid ga s wh ich is p roduced whe neve r f ood is

-

o xidized . Th e ch emist ry O f living p rot opl a sm is exceed

in gyl c o mplic a ted f a r,
t o o c omplic a ted f or m a nk i nd t o

h ope to u n de rst a nd it c ompletely with i na ny re a s ona bly

sh ort futu re t ime Enough is k now na b ou t it h oweve r
.
, ,

th a t th e di ff e re nt pu rp oses se rved by th e met a b olic

p rocesses ca n be set d ow n a s is d one i n succe ed i ng
,

p a ra g ra phs .

B A S I C M ETAB OLI S M —Livi ng p rot opl a sm di ff e rs f rom

.

d ead p ro t opl a sm i n th a t i n livi ng p rot opl a sm the re is

g oi ng onc onti nu ously wh a t we ma y th i nk o f a s th e life “

”
p rocess wh ich is m a de up o f ve ry c omplic a ted chemic a l
,

a ct ivi ties wh o se st opp a ge m e a n s de a th I t is t rue th a t

.

i n m ost ki n ds O f pl a nts a t s ome time i n th ei r hist ory ,

an d i n s ome ki n ds of a ni m a ls li fe is tided ove r f ro m

,

se a s onto se a s onby a d ryi ng ou t of th e p rot opl a sm with

a c orresp on ding c utti ng d ow n of th e ch emic a l a ctivity
g oi ng on withi n it ; so seeds a nd sp ores m a na ge to live
o ve r f ro m on e se a s o nto th e n ext or for seve ra l se as ons
, ,

bec a u se the ch emic a l p rocesses fu nd a me nt a l to life a re

cut d ow nto th e a bs olu te mi nimum a nd not m uch ma
teri a l is used up B u t i n a c tively livi n
. g p ro t opl a s m
th ose chemic a l p rocesses wh ich m a ke up th e b a sis o f life
 G R OWTH M E TA B O LI S M 75

go o na t a f a i rly ra pid ra te a nd i nc onnecti onwith them

,

th e re is a usi ng up o f m a te ri a ls a nd so of necessi ty , , ,

the re must be a repl a ceme nt of this m a te ri a l Livi ng .

thi ngs requi re f ood th e ref ore simply to e na ble them

, ,

to go onlivi n g eve n th ough they ma y not do a nythi ng

,

else .

G R O WT H M ETAB O LI S M Livi ng th in
. gs g row by ma k
in g n e w p ro t o pl a sm which is a dded to th a t which they
a l re a dy p o ssess an d by m a nuf a ctu ri ng such nonlivi ng
,
-

m a te ri a ls a s a re necess a ry to growth For ex a mple the .

,

cellul ose w a lls o f pl a nt cells a nd the h a rd p a rts of the

,

bones i na nim a ls a re nonlivi ng m a te ri al s which a re m a de

-

by livi ng p rot opl a sm in c on

,

necti on with growth In .

pl a nts g rowth met a b olism g oes onm ore or less c onti nu

,

ou sl y th ro ugh o ut life ; i nm a n y O f th e l owe r a nim a ls th e

sa me thing is t rue ; but i na ll th e high e r a nima ls includ ,

in g m a n,
g ro wth met a b o lism ( excep t i n the ski n a nd in
the rep roductive a nd bl oo d formi ng tissues ) g oes ononly
-

du ri ng the e a rly p a rt o f life Whe nth e a nim a l or h um a n

.

bei ng h a s a s we sa y g o tte nits g rowth this met a b olism

, , ,

for the m ost p a r t c omes to a n en d .

T H E M A N UFA CTUR E OF M ATER I A L — In a dditi on to

th ose m a nu fa ctu ri ng p rocesses c onnected with growth ,

which h a ve j ust bee n sp oke n of m a ny ki nds of cells ,

b oth pl a nt a nd a nim a l ma ke sp eci a l ki nds of subst a nces

l

, ,

s ome o f wh ich se rve th e org a nisms which m a ke them ,

a s th e s a liv a m a de by the cells of th e sa liv a ry gl a nds

, ,

se rves i n the digesti on of f ood ; o th e rs like th e opium ,

which th e p oppy pl a nt m a kes h a ve no fu ncti on th a t ,

we k now o f i n th e l ife O f th e orga nisms wh ich p ro

’

duce them Whe n one rec a lls the l ong lists o f a nim a l
.

an d pl a nt p roducts th a t a re used i n i n dust ry a nd , ,

a s d rugs i n medici ne the imp ort a nce of th is s ort of

, ,

met a b olism is cle a r .

T H E M ETAB OLI S M OF A CTI VE FU N CT I O N P ra ctic a lly .

a l l ki n ds o f a nim a ls a nd a good m a ny ki nds of simple

pl a nts p e rf orm a ctive m o ti ons o f one ki nd or a no the r .
76 T HE UTILI Z ATI O N OF FO O D

N ow, a ctive m o ti on wheth e r a ch ieved by a na ut om ob ile

,

o r by a n a n im a l depe nds on ch emic a l t ra nsf orm a ti ons

,

which t a ke pl a ce i n the fuel subst a nce or subst a nces .

I n livi n g org a nisms th ese t ra nsfo rm a ti ons c ome u nde r

the h e a d of met a b olism a nd to d ist ingui sh them f rom
,

th e ch emic a l p rocesses which a re c once rned p rim a rily

wi th the fund a me nt a l life p rocesses or wi th g rowth they , ,

a re cl a ssified a s fu n cti ona l met a b oli sm .

N ERV O U S A CT I V I TY —In p ra ctic a lly a ll a nim a ls the re

.

is still a noth e r ki nd o f fu ncti ona l a ctivi ty th e a c tivity ,

O f th e n e rv o us sys tem Whe n we re a li ze th a t a l l our

.

me nt a l p rocesses as well a s o u r feeli ngs the th i ngs th a t

m a ke up th e re a lly w orth while p a rts o f life — depe nd
up onp rocesses g o ing oni no u r ne rv o us systems we Ca n ,

see th a t this is a ve ry imp ort a nt p a rt of ou r b odily fu nc

ti on in g; an d a s we sh ould expect i t depe nd s up ont ra ns
, ,

f orm a ti ons with i np rot opl a sm Alth o ugh of u tm o st im .

p ort a nce t his met a b olism c onsumes little fuel i n

,

c omp a ris on with th e a m ou nt necess a ry for c a rryi ng on

m oti on .

M ETA B OLI S M As A S OU R CE O F E N ERGY ca n be disti n

i h d f m met b lism wh ich gives ise m i l
'

g u s e ro a o r to a te r a s .

For ex a mple i n growth or i n th e sec reti on o f s a liv a or

,

ga st ric j uice o r i n th e dep ositi on o f f a ts or O ils i n th e

,

b odies O f pl a nts or a nim a ls th e re a re ch emic a l t ra ns

m
,

form a ti ons a s th e resul t o f wh ich f ood a te ri a ls a re

w orked ove r i nt o speci a l ki nds o f subst a nces wh ich ma y

be eith e r d ep osited i npl a ce a s i nthe h a rd p a rts O f b ones
,

or i nth e cellul o se w a lls o f pl a n t cells or ma y be sec re ted , ,

a s i n s a liv a or g a s t ric j uice For th e fu nd a me nt a l life

.

p rocess th a t is for b a sic met a b olism a nd for th e m a ki ng

, , ,

o f m o ti on s onth e p a rt o f a nim a ls th e pu rp ose of met a b ,

ol i sm is a s a s ou rce of e n e rgy for keepi ng th e m a ch i ne

g o i ng a nd d oi ng its w ork This ki nd O f met a b olism
.

th e met a b olism fo r e ne rgy — c onsists a lm o st a lt ogeth e r ,

in pl a n ts a nd a nim a ls O f a p a rt icul a r ki nd O f ch emic a l

,

p rocess bu rni ng or a s it is k now nto th e ch emist oxida

, , ,
 FO O D U S E D I N M E TA B O LI S M 77

ti on,
si nce it c onsists o f a u ni on of th e m a te ri a l th a t is
bei ng bu rned with oxyge n For a f ood subst a nce t o be
.

useful a s a s o u rce o f e ne rgy i t must be of a ki nd th a t will

,

oxidize with in th e p ro t opl a sm a n d so yield th e e ne rgy

which the p ro t opl a sm must h a ve if it is to live a nd fu nc
ti on F oodstu ffs b o th i npl a nts a nd a n
.
,
im a ls must the n , , ,

be m a te ri a ls th a t ca n be m a de ove r i nt o livi ng p rot o

‘

pl a sm or i nt o subst a nces which th e p ro t opl a sm m a nu

,

f a ctu res for one pu rp ose or a nothe r or th a t ca nbe bu rned ,

an d so yield th e e ne rgy whi ch th e p rot opl a sm needs .

I t is w orth while to be a r i n mi nd th a t one imp ort a nt

di ffe re nce betwee n pl a nts a nd a nim a ls is th a t i n pl a nts
the ki nd O f met a b olism th a t results i n the f orm a ti on of
m a te ri a ls p red omi na tes ove r th a t which c onsumes them ,

wh ile i na nim a ls th e reverse is th e c a se S i nce m ost O f .

th e m a te ri a l th a t is st ored i n pl a nts h a s high e ne rgy

v a lue the result O f th is di ffe rence is th a t pl a nts i nge ne ra l
,

g a i n In weigh t a nd p o tenti a l e ne rgy supply th rough ou t

thei r lifetime wh ile a nim a ls a s th e a dult st a ge is re a ched
, , ,

c a rry on a met a b olism th a t is a lm ost e nti rely c once rned

with the c onsump ti on of e ne rgy ; so th a t the met a b olic
p rocesses of a nim a ls te nd to b a l a nce th ose O f pl a nts ,

l ea v i ng na tu re a s a wh ole little if a n y the g a i n er i nt ot a l

'

a m ou n t of stored u p ene rgy -

.

K I N D S O F F O OD U S ED I N
‘

M E TA B O I S M
L — I t h a s j ust
bee n st a ted th a t a ny subst a nce th a t ca n be o xidized
withi np rot opl a sm ca nse rve a s a b a sis for e ne rgy yieldi ng -

met a b olism Th e th ree gre a t cl a sses of f oodstuffs c a rb o

.
,

hyd ra tes f a ts a nd p rotei ns ca n a ll do th is a nd so a re

, , , ,

a l l useful a s s ou rces o f e n e rgy For the met a b olism .

which gives rise to m a te ri a ls only such subst a nces a re

u seful a s ca n be c onve rted i nto th e desi red m a te ri a ls .

Thus for the m a nuf a ctu re of p rot opl a sm i n growth or ,

i n th e rep a i r o f a nim a l w a st a ge th e p ro tei nf orme rs ,

-
,

na mely th e a mi no a cids a re a bs olutely esse nti a l so th a t

,
-
, ,

u nless th ey a re a v a il a ble i nsu fli ci ent a m o u nts no p rot o

pl a sm ca n be m a de eve n th ough othe r ki nds of f ood
,
78 THE UTILI Z ATI O N OF FO O D

stu ff s a re p rese nt i n a bu nd a nce The digestive e nzyms .

an d o the r m an auf c t u r ed p ro ducts o f livi n g p r o t o pl a sm

a re for th e m o st p a r t o f p r o tei n c o mp o siti o n a
,
n d s o r e

qui re a mi no a cid s fo r their p roducti on One c o uld if

- .
,

n ecess a ry n o u r
,
ish h imself i n defi n itely o n a diet c o n sist
in g of p r o t ei n s a s th
,
e E ski m o s f reque n tly h a ve to d o
,

bec a use p rotei ns ca nse rve b o th a s tissue f orme rs a nd a s -

e ne rgy yielde rs Ca rb ohyd ra tes a nd f a ts a re ext remely

-
.

v a lu a ble foods bec a use they fu rnish a bu nd a nt e ne rgy in

e
r a i d ly us a ble f o r m but,
by themselves th ey a re i n
s ufli ci e tn bec a use t h ey h a ve no tissue bu -
i ldi n g c a p a city .

M ETA B O LI S M I N A N I M A LS I S O FTE N W A S T EFU L . I t is

a cu i
r o us an d imp o r t a n t f a ct a n,
d o n e i n wh ich a n im a ls
seem to di ffe r m a rkedly f ro m pl a nts th t ,
a i n the f me o r r

much O f th e met a b olism th a t o ccu rs c onsumes a gr e a t

de a l m o e m a te i a l th a nis th e oretic lly ecess y
r r a n a r . F o r

ex a mple we ca nc a lcul a te th e e ne rgy necess a ry for d oi ng

,

a ce rt a i na m ou n t o f w ork one O f th e simplest c a lcul a ti ons

,

the e nginee r h a s to m a ke We ca n me a su re ve ry a ccu

.

ra tel y the e n e rgy yielded by th e bu rni ng o f a g ive n

,

a m ou n t of food m a te ri a l Bu t whe nth a t f ood m a te ri a l

.

is bu rned i n th e b ody i n c onnecti on with the d oing o f

a give n a m o u n t o f w ork we fi n ,
d th a t much m ore f ood
is bu rned th a nis necess a ry to a c cou nt for th e w ork th a t
is d one I t is a f a mili a r p ri nciple o f e ngi nee ri ng th a t
.

th e e n e rgy consumed i nd oi ng a p a rticul a r job is g re a te r

th a nth e e ne rgy th a t ca nbe rec ove red i nth e d oi ng O f th e
w ork S i nce we h a ve th e l a w k no w n to physics a s th e
.

la w o f th e co n se rv a ti on O f e ne rgy by wh i ch we me a n
,

th a t th e to ta l a mou nt o f e ne rgy nei the r i nc re a s es or de

c re a ses — a l th ough it ma y be ch a nged f ro m one f orm
i nto a noth e r this e n e rgy wh ich fa ils to a ppe a r i n th e
d oi ng o f the w o rk must a ppe a r i ns ome oth e r f o rm ; a nd
t h e f orm i n whi ch i t a ppe a rs i s a lw a ys h e a t . This l a w
a pplies a s well to th e e n e rgy t ra ns form a t i ons O f livi ng
th i ngs a s to th e e ne rgy tra nsform a ti o ns g oi ng o ni nde a d
m a chi nes S ince we fi nd th a t we do not get a s mu ch
.
 —
WA R M B L O O D E D AN I MAL S 79

e ne rgy ou t i n th e f orm of w ork a s i s c onsumed i n th e

wa y O f f oodstu ff s we sh o uld expect to fi d the b a l a nce
,
n
p utti ng i na n a ppe a ra nce i n th e f orm o f h e a t a nd th a t ,

is ex a ctly wh a t h a ppe ns ; wh e reve r met a b olism g o es on

which is a t a ll w a steful he a t is p roduced S ome times
, .

this h e a t se rves a u seful pu rp ose a s it d oes i nth e w a rm,

bl ooded a nim a ls .

W AR M B LOODED A N I M A L S m a ke up the two h igh est

-
,

an im a l g ro up s bi rds a nd m a mm a ls These di ff e r f rom

, .

a l l o th e r livi n g thi ngs i nth a t th ei r b odies a re m a i nta i ned

a t subst a n ti a lly th e s a me tempe ra tu re da y i na nd da y o u t ,

th rough summe r a nd th rough wi nte r S i nce th is me a ns .

th a t thei r b odies d u ri ng m ost O f this time a re w a rme r

th a nth ei r su rro u ndi ngs th ey must p roduce a gre a t de a l
,

of h e a t . Th e a nim a l th a t is w a rme r th a nth e a i r a rou nd

it will te nd to c ool off j ust a s w o uld a st ove or a ny othe r
In an Ima te obj ect i nth e s a me situ a ti on This c ooli ng off
.

wh ich is g oi ng onne a rly a l l th e time is c o u nte rb a l a nced

by th e p roducti onO f l a rge a m o u nts of h e a t which f rom ,

th e st a ndp o i nt O f efficie nt met a b olism w ould be consi d

e red w a steful si nce it rep rese nt s a l a rge c onsumpti on of
f oo d for wh ich no useful w ork i nth e e nginee ri ng se nse
is a cc omplish ed W a rm bl ooded a nim a ls p rofit by th is
.
-

l a rge he a t p ro d ucti on b ec a use it e na bles th ei r b odies to

-
,

st a y a t a f a v ora ble tempe ra tu re f or a ctivity a t a l l times .

Onth e c oldest d a ys o f wi nte r a s well a s onth e ple a s a ntest

d a ys o f summe r th e t emp e ra tu re with i n th ei r b odies is
,

f a v ora ble to met a b ol i c a ctivity .

C O LD — B LOOD ED A N I M A L S on th e othe r h a nd p roduce

, ,

much less h e a t a nd i n c onseque nce th ei r b odie s a re a t

,

a ll times ve r y n e a rly a t th e tempe ra tu r e o f th e su rro u n d

i ngs It f oll ows th a t whe neve r th e su rrou ndi ng tem
.

p e r a t u re f a l l s t o a p o i n t u n f a v o ra ble f o r met a b olism th e ,

c old bl ooded a nim a l mu st suff e r a gre a t d imi nuti on or

-

a n a mo l st c o mplete suspe n si o n o f a ctivity D u r i ng .a

l a rge p a rt of th e ye a r i n c old clim a tes th e c old bl ooded

, ,
-

an a im ls a re hibe rn a ti n g livi n g
,
o ve r t o th e n ext se a s o n
80 T HE UTILI Z ATI O N O F FO O D

by vi rtue o f a ve ry sm a ll a mou nt O f met a b ol is m wh ich

suffices to keep the p ro t opl a sm a live bu t d oes no t pe rmi t
i t a ny re a l a ctivity Th e a ctive life O f a w a rm bl oo ded
.
-

an im a l is l onge r th a nthe e ff ective life Of th e c old bl ood ed -

an im a l wh o se ch ronol ogic a l a ge is th e s a me by j ust the ,

numbe r o f d a ys th a t th e c old bl ooded a nim a l h a s bee n

-

i n hibe rna ti o n .

W A S TE M ATER IA LS ARE P R ODU CED I N CO N N E CT I O N W I T H

M TA O LI S M
E B — This is m ore st riki ng i n a nima ls th a n
.

i npl a nts a lth o ugh the g re a te r e a se wi th which we rec og

,
-

nize the p roducti ono f w a stes i n a nim a ls ma y be d ue to

th e highe r key on which a ll th e m et a b olism of a nim a ls
is pitched At a ny ra te we re a lize th a t i na nim a ls w a ste
.
,

p roducts a re p roduced wh ich must be g o tte n ri d of I t .

is a cu ri ous f a c t a b o ut th e met a b olism of rest or b a sic ,

m et a b olism a s it O ccu rs i na nim a ls th a t it is a cc omp a nied

, ,

by ste a dy w a ste o f th e ve ry c omplex subst a nces of which

l ivi ng p rot opl a sm itself is l a rgely c omp o sed Th u s p ro .

tei ns wh ich h a ve bee ndesc ribed i na ne a rlie r ch a pte r a s

,

th e m ost imp ort a nt c onstitue nt s of livi ng p rot opl a sm ,

a re bei n g c onti nu a ll y l o st f rom th e p rot opl a sm a nd h a ve

to be a s c on ti nu a lly repl a ced Alth o ugh w e do not u nde r
.

st a nd p recisely why th is h a ppe ns th e f a ct th a t it d oes

,

h a ppe nis of a good de a l O f imp ort a nce For one th ing .

,

it me a ns th a t a l l a nim a ls i ncludi ng o u rselves must a o

, ,

qui re i n th e diet th e necess a ry m a te ri a l s by wh ich th e

p ro tei nth a t is l ost i nb a sic met a b oli sm ca nbe repl a ced .

S i nce a n im a ls a re so c ons tituted th a t they ca n only re

pl a ce p ro tei ns with p rotei ns th is me a ns th a t the diet
,

must a lw a ys i nclude p ro tei ns or th e re will be a w a st ing

a w a y o f th e life subst a n ce with its ul tim a te de a th if th e
,

l oss is all owed to go too fa r .

W A S T E I N G R O WT H M ETAB O LI S M is l es s no tice a ble .

Inf a ct i t w o uld seem a s th ough a n

, im a ls sh ould be a ble
to pu t t o gethe r th e con stitue nts o f which p ro to pl a sm is
c omp o sed wi th o u t much w a ste I t is t rue h oweve r th a t
.
, ,

if a s usu a lly h a ppe n s the subst a nces which they get

 WA S T E IN FUN C TI O NAL M E TA B O LI S M 81

inthe food do not c orresp ond ex a ctly with th e subst a nces .

of which th ey a re to m a ke th ei r p ro t opl a sm they will ,

h a ve to p ick a nd ch oose the thi ngs which they ca n use

an d rej ec t th e b a l a nce This gives rise to a na ccumul a
.

ti on of u nused m a te ri a ls i n c onnecti on with g rowth .

S Ome O f thes e ma y be se rvice a ble i n c on necti on with

b a sic or fu ncti ona l met a b olism o the rs ma y be wh olly ,

w a sted G rowi ng b oys ofte nea t a s much a s twe nty fi ve

.
-

p ou nds of food a week a l th ough du ri ng th a t time they

,

ma y not g a i n m ore th a n a p ou nd i n weigh t a nd th ei r ,

c ombi ned b a sic a nd fu ncti ona l met a b olism will not ne a rly
a cc o u n t for th e oth e r t we nty f o u r p ou nds The bes t wa y
-
.

t o expl a i n th e g re a t su rplus Of f ood which g rowi n g chil

d re nregul a rly ea t is by supp osi ng th a t th e g rowth met a b
ol i sm is w a steful so th a t i n p u tt in
,
g t ogethe r a p o u nd
of b o dy subst a n ce a g ood de a l m ore th a na p ou nd of food
is requi red .

W A S TE N F U N CTI O N A L M ETAB O LI S M
I — AS we h a ve .

a l rea dy tri ed to m a ke cle a r th e esse n

’

ti a l di ffe re nce b e
,

twee nfu ncti ona l met a b olism a nd g rowth met a b olism i s

th a t i nthe f orme r th e libe ra ti onof e ne rgy is th e imp ort
an t fe a tu re ; a fe a tu re which is secu red by the bu rni n g Of
fuel We a re a l l f a mi l i a r with th e f a c t th a t whe reve r
.

fuel is bu rned w a ste p roducts a re f ormed S i nce bu rning .

is re a lly the chemic a l p ro cess of oxid a ti on na mely th e , ,

c ombi ni ng of oxyge n with food the re a re b ou nd to be ,

p roduced a s a result o f this c ombi na ti on subst a nces o f

which th e ene rgy v a lue h a s bee n exh a usted a nd wh ich
“

a re th e ref ore f ro m th e st a n
, ,
dp oint of the tissues w a ste ,

p roducts Chief a m ong these is c a rb ondi oxid one of

.
-
,

the m a te ri a ls which a s we sa w i n a n e a rlie r ch a p te r

, ,

se rves a s the ra w m a t e ri a l for the m a nuf a ctu re of sug a r .

Besides c a rb ondi oxid w a te r a nd v a ri ous oth er less Well

, ,

know n w a st e p roducts a re f ormed i n c onnecti on with

fu ncti ona l met a b olism .

WA S TE M U S T B E G OTTEN R I D OF I F META B OLI S M I s To

CON TI N U E Th e fu rna ce fi re c a nno t be kep t g oing if
-
82 THE UTILI Z ATI O N OF FOO D

th e p rod ucts of c ombus ti on a re no t rem oved Th e re .

must be sufficie nt d ra ugh t to c a rry th e flue g a ses up the

-

ch im ney a nd th e a shes must be cle a red o ut f rom t ime

,

to time . This necessi ty O f get ti ng ri d of th e p rod ucts

o f chemic a l a ctivity a pplies to b odily met a b olism It .

f oll ows th a t u nless th e w a ste p roducts a re c a rried O ff a s

they a re f ormed the me ta b olism itself is ce rt a i n to be
h a mpe red Some me a ns of get ti ng ri d o f w a ste p roducts
.

the nis a necess a ry p a rt of th e equipme nt of livi ng tissues .

M ETA B OLI S M G O ES O N I N EV ERY CE LL A p oi nt th a t

.

m ust be kept in mi nd is th a t a ll liv ing p ro t opl a sm h a s

th ese v a ri ous ch emic a l a ctivities g oi ng onwi thi nit This .

me a ns th a t eve ry livi ng cell must h a ve a ccess to f ood .

Fu rthe rm ore si nce a l a rge p a rt o f met a b olism is o xid a

,

ti on supplies o f o xyge nmust be a v a il a ble to eve ry cell

, ,

an d eve ry cell m ust h a ve s ome me a ns o f riddi n g itself

o f w a ste p roducts .

T H E S ERV I CES O F S UPP L Y — I n th e highe r a n

. im a l
f orms i ncludi ng ma n m ost o f th e c ells o f th e b ody a re
, ,

so situ a ted th a t they c a n no t p ossibly h a ve di rect a ccess

to th e s o u rce o f supply o f th ei r f ood or oxyge no r to a n y
o utside regi oni n t o which th ey ca nd isch a rge th ei r w a ste .

This me a ns th a t i na ll high e r forms s ome speci a l me a ns

mus t be p rovided for i nsu ri ng to eve ry cell th e necess a ry
f ood a nd oxyge na nd th e m e a ns of get ti ng ri d O f its w a s te
p rod u cts Th ese we ma y c a ll th e servi ces of su pp ly
. .

Th ey a re f ou r i nnumbe r : (1 ) a ge ne ra l t ra nsp ort a ti on

system ; (2) a se rvice for th e p rep a ri ng a nd delive ry o f
food ; (3 ) a se rvice for th e delive ry of o xyge n; (4 ) a
se rvice for the rem ov a l O f w a stes .
 C H APTER Ix

T HE T R AN S P OR TATI O N S Y S T EM I N ANI MAL S

CO N D I T I O N S F A V OR I N G CEL L A CT I V I TY .
— E ve ry livi ng
cell resembles a ti ny l a b ora t ory wh ose st ructu re dete r
'

mines th e na tu re o f th e p ro cesses it c a rries on Th e w ork .

d one by a ne rve cell is di ffe re nt from th a t pe rf ormed by

a muscle fibe r a n d b o th di ffe r from the op e ra ti on s Of a n
a m oeb a . Y et h oweve r g re a t th ei r st ructu ra l diffe re nces
,

an d thei r m odes of ope ra ti on a ll cells a g ree i nrequi ri ng

,

cert a i n c onditi ons for thei r pe rfect ope ra ti on In the .

fi rst pl a ce th e temp e ra tu re mus t be a b ove f reezing a nd

,

bel ow 1 20 F i n m ost i nst a nces And wh a t is of the

°
. .
,

utm ost imp ort a nce i nc onnecti onwith the subj ect m a tte r
,

o f th e p rese n t ch a p te r eve ry a ctive cell must be b a thed

,

to a gre a te r o r less exte nt by a liquid This w a te r front .

a ge se rves two pu rp o ses I t m a i nt a i ns the p rope r deg ree

.

O f fluidity O f the p ro t opl a sm a n d c ont a i nin,

g a s it d oes
, ,

f ood subst a nces it a cts a s a st oreh ouse from wh ich the

,

cell d ra ws its supply a nd i nt o wh ich it disch a rges its

w a stes.

N EED OF TRA N S P ORTATI O N N ot on ly must eve ry cell

.

h a ve th is w a te r front a ge from which it withd ra ws its

necess a ry food but it is evide nt th a t this fl uid must
,

be re newed or shifted i norde r to p rovide for new supplies .

Am ong th e sm a lle r a q u a tic a nim a ls wh ich c onsist o f a ,

si ngle cell or o f m a ny cells l o c a ted a t or ne a r the su rf a ce

o f th e b ody the m o veme n
,
t of the w a te r or of th e b ody
in the w a te r is a l l th a t is necess a ry in this respect .

Am ong th e h ighe r a nim a ls on the o the r h a nd m a ny of ,

the cel ls a re bu ried f a r be n e a th the b ody su rfa ce a nd a re

thus rem ote from th e immedi a te s ou rce of supplies This .

83
84 T R AN S P O R TATI O N S YS TEM I N ANI M AL S
h a ndic a p is ove rc ome by s ma ll flui d c ont a ining sp a ces -

a mon g th e tissues up onwhich eve ry cell h a s a f ront a ge .

M ore ove r these sp a ces usu a l ly f o rm a c ontinu o us sys

,

tem th a t i s supplied a t defini te p oi nts wi th f ood

m a te ri a ls a nd kep t in a cti on by a definite p ump ,

the h e a rt so t ha t the w a nts of eve ry cell a re a dequ a tely

,

p ro vided for .

CELL D I FFU S I O N . Th e diss olved f ood m a te ri a ls p a ss

int o the cell by th e p ro cess o f diffusi on a s desc ribed i n ,

a p revi ous ch a pte r . He nce i t foll ows th a t the cell mem

b ra ne must not only be th i na nd pe rme a ble to p e rmit of
a ra pid m o veme n ,
t bu t it must be O f rel a tively g re a t ex
tent wh e n c omp a red to th e e ncl o sed p ro topl a smic m a ss .

Th is st a te of a ff a i rs exists only whe n th e m as s is sm a ll ,

a s will be see n f ro m the f oll o wi n g illust ra ti on:

CE LL S U R FA CE AN D B OD Y S U R FA CE — I f we t a ke a
cubic f oo t bl ock of w ood a nd sa w it i nt o cubic i nch bl ocks
t h e exte rn a l su rf a ce of these sm a lle r m a sses is twelve

times g re a te r th a nth a t o f th e origi na l one If we red uce .

it a ll to s a wdust th e result a nt su rf a ce is i nc re a sed to a

much gre a te r deg ree Th e fi ne r the subd ivisi on the
.

g re a te r th e e nti re su rf a c e .

Applyi ng this p ri nciple to th e b ody O f a ma n f or ex ,

a mple we fi n
, d th a t th e o ute r su rf a ce me a su res a p p rox i
m a tely sixtee n squ a re feet a rel a tively sm a ll a m ou nt
,

wh e nc omp a red to th e m a s s weighi ng 1 50 p ou nds But .

i nre a lity this b ody excludi n,

g much o f th e b ones b ody ,

fl uids a nd ce rt a inothe r st ructu res is subdivided i nt o a n

,

inc onceiv a bly g re a t numbe r o f cells Inth e bl ood a l one.

it is estim a ted th a t the re a re cells a nd ,

i t is s a fe to sa y th a t th e e nti re b ody c omp rises a t le a st

five times th is numbe r If the refore th e su rf a ce of the
.
, ,

bl ood cells h a s a supe rfici a l a re a o f squ a re y a rds ,

as h a s bee nestim a ted it f oll ows th a t th e su rf a ces O f a ll

,

the cells o f th e b ody me a su re not less th a n sq u a re

fee t or m ore th a n
,
times g re a te r th a n the b ody
su rf a ce .
 WAT E R FR O NTAG E 85

W ATER FRO N TA L W
GE I N A A O — Am ong the
ER N IM LS

on e celled a n
-
im a ls no speci a l t ra nsp orta ti onsystem exists ,

for re a s ons a l re a dy desc ribed Ins omewh a t highe r a ni

.

m a ls such a s hyd ra it will be remembe red the c omp one nt

, , ,

cell s a re a rra nged to f orm a n o u te r p ro tective a nd a n

inne r digestive l a ye r su rrou ndi ng the ce nt ra l digestive
c a vi ty F ood m a te ri a l ente ri ng th e sa c is ci rcul a ted by
n
.
, ,

me a ns O f fl a gell a b or e on s ome of th e cells a nd is thus ,

b rough t i nt o i ntim a te rel a ti onwith th e e nti re i nne r l a yer ,

by which it is t a ke nup Th e excess of nut ritive m a te ri a l

.

di ffuses to th e oute r cells N o speci a l t ra nsp ort a ti on

.

system exis ts a nd i n th is respec t hyd ra resembles its

,

rel a tives th e sea a nem ones j elly fi sh es and th e m ore

, ,
-

dist a ntly rel a ted sp onges Whe re the di ffusi onst re a m a ls o

supplies th e needs of sc a tte red cells betwee n the oute r
an d in ner b ody l a ye rs .

FI G 1 4 . D i a gra m of th e D iges tiv e S ys te m

.
of a Fl a tw orm .

M onth i n center of figure .

TH E F LATW OR M T R A N S P ORTATIO N SY S TE M The fla t .

w orms a re sm a ll a nim a ls usu a lly no t o ve r a n i nch in

,

le ngth freque ntly be a utifully c ol ored a nd a bu nd a ntly

,

rep rese n ted a m ong th e rocks a l ong the se a c oa st a nd to a

less exte nt i nfresh Wa te r Th e b ody is fla t a nd le a f like
.
-
,

a s thei r na me suggests a n d onits u nde r su rf a ce be a rs the

,

m outh ope ning This le a ds i nt o a ce nt ra l c a vity f rom

.

wh ich i ntesti na l b ra nch es exte nd th rough th e a nim a l to

its outer m a rgi ns whe re e a ch e nds bli ndly (Fig .
86 T R AN S P O R TATI O N S YS TEM I N ANI M AL S
With inthis system the f oo d is not only diges ted but i s ,

circul a ted th rough ou t its e nti re extent by me a ns o f mus

cul a r c o n t ra cti ons O f th e e ncl osing w a ll th us e na bli ng ,

eve ry cell of the diges tive t ra ct to sec u re a p orti ono f the

f ood supply The excess diff uses th rough the i ntest ina l
.

w a lls supplies th e cells o f the i mm edi a te neighb orh ood

,

an d still f a rthe r o ne nte rs nume ro us liquid fil led c a vities -

betwee nthe o utlyi ng b ody cell s th a t a re th us supplied ,

.

In these a nim a ls th e refore th e t ra nsp ort o f f ood ma

, ,

teri a l s f a lls up on th e digestive t ra c t a n d to a less exte nt

up onth e b ody fluid th a t is kept i nm oti onme rely by th e
m oveme nts o f th e b ody .

FI G 1 5 . . Co mm o n W a te r Fl ea (D a ph nia ) sh owing h ea rt (h) in

th e mids t Of a l a rge bl oo d spa ce tra v ers e d by th e i nte s tine (d)
.

M ODER ATELY CO M P LEX TR A N S P ORTAT I O N S Y S TE M S .

Ab o ve this p oi nt the re a re seve ra l inte resti ng types o f

t ra nsp ort a ti on systems a nd of these we sh a ll ex a mi ne
,

on e rep rese n ti ng a h a l fw a y st a ge betwee nth e fl a tw orms

an d th e highe r cl a sses O f a nim a ls (Fig I t exists i n .

th e c ra bs i nsects spide rs a nd th eir a llies a nd a cc ordingly

, , ,

is ch a ra cte ristic o f two thi rd s o f a ll th e know nspecies o f

‘
-

an im a ls In th ese c re a tu res th e re i s a n ex te rna l b ody

.

w a ll e ncl osing a c a vity sp a nned by th e digestive t ra ct a nd

,

pe net ra ted by nume rou s muscles th a t neve rth el ess le a ve

a c on tinu ous th ough i rregul a r sp a ce o f rel a tively g re a te r
m
v olu e th a ni nth e fl a tw orms I t likewise h olds a fluid .
,

bu t a s speci al cells or c orp uscles exist i nit a nd since it ,

h a s a f a i rly definite ch emi c a l c omp osi ti on ,

this fl uid is
 C I R C ULAT O R Y S Y S TE M

now known a s bl ood Fu rth e rm ore the b ody w a ll is

.

rigidly held i np ositi onby a fi rm he a v y shell or c a se a n d ,

thus me re b odily m oveme nts a re no l onge r sufficie nt to

shift th e b ody fl uid ; he nce a muscul a r pump the h e a rt , ,

is p resent to d rive i t a l ong a f a i rly defi nite route In .

m a ny of the l a rge r species the re a re vessels le a ding from

the he a rt to v a ri ous p a rts of th e b ody which a re thus ,

m ore di rectly sup p lied with ma

teri a l s f rom the digestive t ra ct a n d
respi ra t ory org a n s bu t s oon
,
e r or
l a te r th e bl ood p ou rs th rough the
ope n e n ds O f these a rte ries i nt o
the m a i n b ody c a vity a nd is thus
b rought i nt o di rect c ont a ct with
the v a ri ous tissue cells .

CI R C ULAT OR Y S Y S TE M OF H I GH ER
A N I M A LS — In th e h ighest g roup
of an im a ls th e ve rteb ra tes or
,
.

th ose with b a ckb ones th e t ra ns ,

p o rta ti on o r ci rcul a t ory system is

a n exte n sive a nd c omplic a ted set
o f vessels c ommu n ic a ti ng with a
c omp a ct a nd p owe rful pum p th e ,

he a rt (Fig . Th e ex a ct c o u rse
o f the bl oo d th ro ugh these ch a n

nels is not a n imp ort a nt m a tte r

i n this c onn ecti on bu t it ma y be
,

s a id th a t the one or two gre a t

—
, ,

2 gmm
vessels or a rte ries le a vi ng th e
;
F 1 6 D i
ft 3
1

g
‘
° 0

zjjna l A 2mm
e l ’ f S 1
y
h e a rt b ra nch repe a tedly a s they a s

m a ke thei wa y i nt o th e tissues a nd
r H hea t L 1 1
m rV e i
m S t p pl d
n
,
S
.
,
Y

fi na lly c ommu nic a te with th e ca p 3&2 ?

t ) l e

i lla ri es
. Th ese a re exceedi ngly
sle nde r a nd delic a tely w a lled vessels which it is i m , ,

p ort a nt to not e a re so nume rous a nd widely dist ributed

,

th a t th ey a re i n th e vici nity of p ra ctic a lly eve ry cell of

th e b od y Onth e oth e r h a nd th e c a pill a ries u nite with
.
,
88 T R AN S P O R TATI O N S YS T E M I N AN I M AL S
l a rge r ves sels th e vei ns wh ose u ni o n fi na lly results i n
, ,

on e or t wo m a in vessels e n
, ,
te ri ng th e h e a rt .

I t will be see nf rom this b rief desc ripti ono f th e ci rcul a

ti onth a t the bl ood neve r le a ves these vessels a nd a cc ord , ,

in gyl a t no p o int d o es it c ome in di rec t c ont a c t with th e

ce lls of th e b ody excepting th ose of the bl ood i tself a nd
th ose li ni ng the ci rcul a t ory system I t is a cl osed system .

an d th e t ra nsfe r of nut ritive m a te ri a ls to the tissues a n d

th e rem ov a l o f w a stes to th e bl oo d st re a m is p rovid ed
for by a speci a l p ro cess th a t w i ll be m ore cle a rly u n de r
st ood a fte r we h a ve ex a m ined s ome o f th e peculi a rities
o f bl ood .

T H E B LOOD The f a mili a r red fl uid known a s bl ood

.
,

whe n ex a mi ned u nde r the mic rosc ope is see n to c onsist ,

o f a fl uid th e pl a sm a i n which the re a re fl oa ti ng cells

, ,

o f two types th e red a n ,

d white c orpuscles Th e pl a sm a .

c onsists l a rgely of w a te r i n wh ich f ood a nd w a ste sub

st a nces a re diss olved t ogethe r with ce rt a i n o the r ma
,

teri a l s s ome o f which a i d i n th e dest ruc ti o n o f d ise a se

c a rryi ng ge rms wh ile o th e rs regul a te v a ri o us a ctivities
,

o f th e b ody cells Th a t i t is m ore th a n w a te r i s a ls o

.

indic a ted by its a bility to c oa gul a te or form a cl o t ina n

i nj u red vessel th a t thu s plugged up i s i nc a p a ble o f
, ,

fu rthe r bleedi ng Th e wh ite c orpuscles a re likewise a o

.

tive a ge nt s i ne ngulfi ng a nd digesti ng ge rms i nth e bl o od

st re a m Ins ome inst a n
. ces they ma y eve nc a rry on th is
w ork outsid e o f th e c a p il l a ries th ro ugh wh ose w a lls t h ey
h a ve b ored a wa y ; or they ma y c a rry off a nd diges t f ra g
me nts o f i nj u red cells wh ose rem ov a l p a ves th e wa y for
rep a i r p ro cesses Th e red c orpuscles owe th ei r na me to
.

th e p rese nce of a pi gme nt th e hem ogl obi n c a rried i n , ,

thei r subst a nce As this m a te ri a l is O f th e u tm ost i m

.

p ort a nce i nthe p rocess o f respi ra ti on i ts ge ne ra l ch a ra c ,

teri sti cs will be c on sid e red i nth a t c onnecti on .

LY M P H . Th e bl ood c a pill a ries a re not only widely

-

dist ributed bu t th ro ugh o u t th e g re a te r p a rt O f thei r ex

,

te nt they a re su rrou nded by sli t like sp a ces so th a t p ra c -

,
 L Y M P HATI C OR A B S OR B E NT V E S S E L S 89

tic a lly eve ry b ody cell bei ng i n th e neighb orh ood of a

,

c a pill a ry b orde rs up on one of these c a vities (Fig

, .

Fu rthe rm ore the w a lls of th e c a pill a ries a re ve ry thi na nd

,

pe rmit sm a ll qu a ntities o f liquid f rom the bl ood c onti nu

a lly to di ffuse i nt o these sp a ces a nd th u s b a th e th e tissue
,

cells This fl uid k now na s lymph is of th e high est i m

.
, ,

p ort a nce to the existe nce of the cells O f h ighe r a nim a ls .

V e ry much a s a fish lives i nw a te r a cell lives i nlymph .

It is the s ou rce from which it de rives the f ood th a t h a s

bee n b rough t f rom th e digestive t ra ct a nd th e oxyge n ,

th a t h a s c ome from the gills or lu ngs a nd is the medium ,

i nt o which it disch a rges its c a rb ondi oxid s a lts a nd othe r , ,

w a ste p roducts Lymph the ref ore a cts i nth e na tu re of

.
, ,

FI G 1 7 . . D i a gra m I llus tra t i ng B l oo d C a pill a ri es (c) surro unded

by a lymph sp a ce (l ) whi ch is dra ine d by a na bs orb ent or lymph a ti c
ca pill a ry (a ) .

a r et a il st a ti oni nte rmedi a te betwee nth e wh oles a le supply

i nthe bl ood a nd the c onsume r or the cell
, ,
.

LY M P H AT I C OR A B S OR BE N T V ES S E L S Lymph is con .

tin u a ll y di ffusi n g th rough the c a pill a ry w a ll s a s h a s ,

bee ndesc ribed a nd the excess usu a lly esc a pes by wa y of

,

a speci a l system k n ow n a s the lymph a tic or a bs orbe n t

vessels This c omp rises a v a st numbe r of lymph ca p il
.

l a ri es a,
e ch Of which e nds i n o ne o f th e ci rcum c a pill a ry -

tissue sp a ces a nd onthe othe r h a nd u nites with sim il a r

,
90 T R A N S P O R TATI O N SYS TEM I N ANI MAL S
b ra nch es wh ose subseque nt u ni on fi na lly results in one
or two l a rge vessels ope n i ng di rectly i nto a l a rge vei ni n
th e regi ono f the Sh o ulde r . Th e lymph st re a m di ffusing
,

f rom th e bl ood a nd fl ow i ng p a st the cells e nte rs these

,

lymph c a p ill a ries a nd is fi na l ly retu rned to the bl ood .

S MM R
U A. Y — A t ra nsp ort a ti on system in th e f orm of
,

vessels or ex te nsive bl ood sp a ces c a rries nu t ritive ma

,

teri a l s to a n d rem o ves w a ste p rodu cts f rom eve ry b ody

cell in a ll bu t th e simplest species o f a nim a l s In the .

h ighe r types th e bl ood d oes no t c ome int o d i rect c ont a ct

with the tissues bu t a c onst a nt st re a m o f fl uid the lymph
, , ,

di ffuses f rom th e c a p ill a ries i nt o a dj oi ni ng sp a ces f rom

which th e cells a bs orb their food m a te ri a ls a nd i nt o wh ich
they disch a rge th eIr w a s tes Bey ond th is p o int th e
.

st re a m e nte rs c omm u nic a ti ng lymph c a pill a ries a nd ,

sl owly j o u rneys th rough th ese i nt o l a rge r vessels th a t

fina lly p ou r th ei r c onte nts i nt o the bl ood st re a m We .

sh a ll now c onside r th e meth od whe reby food subst a nces

a re i n t rod uced i nt o th e t ra nsp ort a ti on system a nd the
w a stes a re rem oved .
 C HAPTER X

T HE F OO D P R E P A R ING
-
S Y S TEM

D I GES T I O N . Ina ne a rlie r

ch a p te r (V I I ) we sa w th a t
the c omplic a ted m a te ri a ls which a re bu ilt up by pl a nts
an d st ored withi nthem h a ve to u nde rg o a b re a ki ng d ow n -

i nto sirnp l er c omp ou nds i norde r th a t th ey ma y be m oved

a b out f rom pl a ce to pl a ce or i norde r to m a ke them use
,

ful for the p u rp oses of th e pl a nt This b re a king d ow n

.
-

of the c omplic a ted in t o the sim p ler subst a nces we h a ve

le a rned to c a l l di ges ti on Th e f oo d of a nim a ls c onsis ts
.

eithe r of th e p rot opl a smi c b ody subst a nce O f a nim a ls or

pl a nts or of th e m a te ri a ls which th ey h a ve m a nuf a ctu red
an d st ored a w a y withi n themselves M o st of th ese ma
.

teri a l s a re too c omplic a ted to be used by th e cells of th e

an im a l b ody i n p recisely th e form i nw hich th ey a re
t a ke n i n a nd so must unde rg o digesti on before th ey
,

ca nbe u tilized .

CLA S S E S OF F OOD S 4 As we h a ve a l re a dy le a rned th e

.
,

m a ny ki nds O f subst a nces c a lle d f oods ca n be gro uped

int o a few cl a sses Inth e fi rst pl a ce we Set off th e foods
.
,

p rop er which ca n y i eld e ne rgy or be built i nt o p ro t o

,

pl a sm fro m th ose which c a nno t ; the f orme r a re O ften

,

c a lled nu tri ents wh ile th e foods which a re O f use for

,

o th er p u rp o ses a re c a lled food a ccessori es In th e cl a ss

.

of f ood a ccess ories f a ll i n o rg a nic m a te ri a ls like ordi na ry

t a ble s a lt a nd the lime s a lts wh ich h a ve to be t a ke nby
,

y ou ng child re ni norde r th a t thei r b ones ma y be p rope rly

.

f ormed a n ,
d th e series of org a nic subst a nces wh ose i m
p ort a nce we h a ve only l a tely c omme nced to re a li z e th e ,

V it a mi ns These a re c omplex org a nic subst a nces which

.

a re n ot n ut riti o us i nth e se nse th a t they fu rnish us with

92 T HE FOO D—P R E P A R I NG S Y S TEM

a ny e ne rgy nor so fa r a s we know a re built i nt o livi ng

, , ,

p ro t opl a s m yet if they a re left ou t of th e diet we do not ,

th rive I t is evide nt th a t s ome esse nti a l b od ily p rocesses

.

c a nnot go onwell u nless these vi t a mi ns a re p rese nt bu t ,

j ust wh a t th e na tu re o f these p rocesses ma y be is

not a t p rese nt a t a ll cle a r .

T H E F OODS P RO P ER f a ll i n to
t h r e e c l a s s e s : c a rb ohydr a tes
(sug a rs a nd st a rch es ) f a ts a nd , ,

p rotei ns Th e a ccess ories do not .

h a ve to be digested they a re util ,

i z ed by th e b ody i n th e f orm i n
wh ich th ey a re t a ke n i n All o f .

th e nut rie nts except s ome o f th e

simple r suga rs h a ve to be digested ,

eith e r bec a use i nth e f orm i nwhich

th ey a re t a ke n i n they a re in
s oluble i nw a te r a nd so ca nnot be
t a ke n up by th e b ody fl uids or ,

bec a use th ey a re no t i n such

chemic a l form th a t the cells ca n
F G 18 D i gra m of use th em
“

I . . a .

A lim nt y C n l i n
e a r
T H E A LI M E N T ARY T R A CT i nth e
a a a
High V te b te (m n)
er er ra a

s t m h ; high e r a nim a ls c onsists fi rst of a

.

6 eS p h gus ;
, O a a, o ac , ,

i sm ll i n te s t ine; 0 l g
, a
tube i nt o which is i nse rted a t one
, ar e

i nt s t i n ; g s liv y
e e a a r
,

en
,

gl nd ; p p n a s F
a , ad th e m ixtu res O f c o mplic a ted
cre . or

th s k O f s i mpli i ty th
e a e
m a te ri a ls whi ch th e a nim a l e a ts
c e

liv er nd t h p ss g s
a e a a e
O pe ni ng f o m th li m n
r
a n d with i n wh ich
e a
th e p ro cess o
e
f
t y c n l (n s t ils
a r a and p rep a ri ng th e food f or the use of
o r a
‘

wi ndpipe ) a mi tte d
t h e cells g oes on a n d sec ond o f
re O .

, ,

cell ma sses c a lled gl a nd s which m a nu f a ctu re th e v a ri ou s

-
,

d igestive sec reti ons a nd p o u r th em o u t i nt o th e a l i men

t a ry t ube Th e th ree f ood cl a s ses the c a rb ohyd ra tes
.
, ,

f a ts a nd p ro tei ns h a ve to be t re a ted i ndivid u a lly ; th a t

, ,

is to sa y the p ro cesses by wh i ch c a rb ohyd ra tes a re di

,

ges ted do no t se rve fo r f a ts no r is the digesti ono f fa ts ,

ex a ctly like th e digesti on o f p rotei ns S o we fi nd the .

 T HE M AN U FA C T U R E OF E NZY MS 93

a lime nt a ry t ra ct org a nized to de a l p rope rly with e a ch

cl a ss o f f ood .

T H E M A N UFA CTU R E O F E N ZY M S — The meth od of

diges ti on a s st a ted p revi ously (see Ch a p V I I ) is to
,
.
,

b ri ng i n c ont a ct with th e f ood a p a rticul a r ki nd o f sub

st a nce k now na s a nenz ym which h a s the speci a l p rope rty
,

o f h a s te n i ng th e dec omp ositi on o f th e c omplex f ood

m a teri a l i nt o the simple r subst a nces which we c a ll th e
di ges ti on p rodu cts D i ff e re nt kinds O f e nzyms must be
.

p rovided for the di ffe re nt f ood c onstitue nts I n th e .

h ighe r a nim a ls the e nzym s a re m a nu f a ctu red a s p re ,

vi o u sly st a ted by g roups of livi n

, g cells which h a ve th a t
a s thei r speci a l fu n cti on a nd which we c a ll secre ti ng

FI G 1 9
. .
— D i a gra m S impl e Gl a nd
of a . T h e s ecreti ng cell s (a )
m a k e up a t ub e , cl os e d a t t h e b o ttom , o p eni ng a t th e top into th e
a lim en ta ry ca nal S u rro u ndi ng t his t ub e is a netw ork O f fi ne bl oo d
.

v essels I n th e sp a ces b etw een th e bl o o d v ess els a n d th e gl a nd

th ere is tissu e fluid .

gla nds (Fig T hese. gl a nds with d ra w f rom th e tissue

fluid which su rrou nds th em v a ri o us f ood c ons tituents

thei r fu ncti ona l met a b olism c onsist s i n m a nuf a ctu ri ng
ou t o f this m a te ri a l the speci a l subst a n ces which se rve
a s en zyms These l a tte r a re p ou red ou t a l ong with a
.

st re a m of w a te r f rom th e gl a nd i nt o th e a lime nt a ry t ra ct ,

whe re the e nzym bec omes mixed with the f ood Th e .

m ost f a mi li a r ex a mple of this is th e s a liv a which is ,

m a nuf a ctu red by the s a liv a ry gl a nds p ou red ou t i nto th e ,

m outh a nd th e re mixed with th e f ood a s it is bei ng

,

chewed .
94 THE FO O D—P R E P A R ING S Y S TEM

SA LI V A .
— Th e fi rstthe di gestive e nzyms is th a t in
of

s a l iv a
. I t is c a lled p tya li n a s ort O f a myl as e a nd i ts , ,

a cti onis to dec omp ose st a rch in t o a f orm Of sug a r which

is f a m ili a r to the chemist but not especi a lly f a mili a r i n
o rd in a ry life I t is the so c a lled m a lt sug a r being ex
.
- -
,

a c tly th e s a m e s o rt o f sug a r th a t is p rod uced i ngermin at

ing b a rley i n c onnecti on with th e m a nu f a ctu re O f m a lt .

This m a l t sug a r h a s to u nde rg o fu rth e r digesti onin th e

-

sm a l l i ntest ine bef ore it is re a d y to se rve a s fuel for

livi ng cells .

G A S TR I C JU I CE Afte r the foo d is sw a ll owed i t e nte rs

.

th e st om a ch whe re i t c omes i n c ont a ct with th e sec ond

e nzym o f the se ries p ep si n a p a rt o f the g a st ric j uice
, , .

This p rote oly tic (i e p rotei nsplitting ) e nzym begi ns . .

-

the dec omp osi ti on O f th e p rotei ns reduci ng them f rom ,

th e highly c omplex f orm i n wh ich th ey a re e a te n to

simple r f orms k now n a s p rote o ses a nd pept ones Th e .

a cti o n o f pepsi n is like th e a cti on o f pty a li n i n th a t i t

is no t fi na l but p rep a ra t ory Fu rth e r a cti on must t a ke .

pl a ce to c omplete th e digesti onth a t is st a rted by pepsi n ,

a s well a s by pty a li n .

P A N CREAT I C JU CE I — A fte r th e p a rtly digested f ood

p a sses f rom th e st om a ch i nt o the sm a ll intesti ne it comes
i n c o nta c t wi th th e sec reti o n O f th e p a nc re a s to wh ich
we give the na me o f p a nc re a tic j ui ce I t cont a i ns th ree .

e nzyms Th e fi rst is a n e nzym wh ich h a s ex a ctly th e

.

s a me a cti ona s th e p ty a linO f s a liv a i e ca nc a use st a rch ,

. .
,

to t a ke up wa ter a n d split i nt o m a lt o se o r m a l t sug a r

'

an d a s fa r a s we k n ow is th e s a me e n zym Th e sec o nd .

i s lip a se which digests f a ts Th e thi rd is tryp si n which

, .
,

digests p rotei ns T ryp si n i s a m ore e ff ective e nzym

.

th a npepsi n i nth a t i t c a rries th e digesti on i e h y drol y

, ,
. .
,

tic splitti ng O f p roteins much fu rthe r In f a ct th e re

, .
,

sul t of digesti on by t ry p si n is to b re a k th e p roteins up

int o th e a mi no a cids o f wh ich thei r m olecules a re
-

c omp osed .

T H E CO M P OS IT I O N O F P ROTEI N S As h a s a l re a d y been .
 T HE I NT E S TINAL J UI CE 95

st a ted (Ch a p V I I ) the ve ry c omplex p rotei n m olecules

.
,

a re built up by c ombi n i ng th e m olecules of ce rt a i nnit ro

en c on t a i ni ng org a nic a cids to which a re give nth e n a me
g
-

o f a mi n o a cids-
The re a re twe nty di ff e re nt a mi no a cids
.
-

now know na s c ons titue nts of p rote i ns One p rotei nma y .

di ffe r from a nothe r i n the a ctu a l a mi no a cids wh ich a re -

p rese nt or i n the rel a tive p rop orti ons of th e di ffe rent

,

a min o a cids or in the wa y i n which the a min

-
o a cids a re -

m
,

pu t t ogeth er i n its m olecules A S imple a th em a tic a l

.

c a lcul a ti on will sh ow th a t by st a rti ng with a s m a ny a s

twe nty di ffe rent a mi no a cids a nd g ra nting th a t they ma y
-

be put t ogethe r in di ffe re nt w a ys or i nd iffe re nt p rop or

ti ons or th a t one or m ore ma y be left ou t a g re a t v a riety
, ,

o f p ro tei n s ca n exist S i nce we a re na tu ra lly i nclined

.

to believe th a t the di ff e re nces wh ich o ccu r a m on g a nim a ls

an d pl a nts a re i n m a ny c a ses a t le a st b o u nd up with
, ,

chemic a l diffe re nces i n thei r p roteins we ca n thu s con ,

cei v e h ow th e di ffe re n ces a re p ossible I nfa ct i t is h a rd .

,

for us to im a gi ne h ow livi ng p ro t opl a sm wh ich h a s m a n y ,

fe a tu res i nc omm on whe reve r it is fo u nd a nd yet which

ca n t a ke this i n finite v a riety of f orms Could exist we re ,

it not th a t i ts ch ief c onstitue nt is a subst a nce h a vi ng the

peculi a r c omplexity of p ro tein Th e re a s onwhy p rotei n .

must be digested i nto i ts c onstitue nt a mi no a cids bec omes -

cle a r from th e fa cts which h a ve j ust bee nset dowh ; for ,

while in one or a ll o f th e th ree respects o utli ned a b ove

th e p ro tei no f th e f ood is a lw a ys cert a into di ffe r from the
p rote i n withi n th e b ody o f th e org a nism which e a ts it ,

th e a mi no a ci ds i nth e f ood p rotei na re ex a ctly th e s a me

-

a s the a mi n o a cids i nthe b ody p ro tei n

-
He nce it f oll ows .
,

th a t if the f ood p ro tei ns a re b roke nup i nt o thei r c onsti

tuent a mi no a cids th ese ca n the n be rec ombi ned i n
“

m
-
,

p rope r p rop orti onto for th e b ody p roteins .

T H E I N TE S TI N A L JU I CE — Th e f ood which h a s been

a cted up onby th e p a n c re a tic j uice will h a ve a ll its st a rch
c onve rted int o m a lt sug a r a ll or m ost of its p ro tei ns
-

—
,

dec omp osed int o a mino a cids a nd its f a ts digested by ,

96 T HE —
FO O D P R EP A R ING S Y S TEM

the lip a se o f th e p a nc re a tic j uice Fu rthe r a l ong i n the

.

w a ll o f the intesti ne a re ti ny gl a nds which sec rete wh a t

is c a lled i ntes ti na l ju i ce Th is j uice c o nt a i ns seve ra l
.

e nzyms Am ong th em one suppleme nts the a c ti on of

.
,

t rypsi n so th a t i f a ny p ro tei ns esc a pe digesti on i n the

,

uppe r p a rt o f the sm a ll i ntesti ne they a re su re to be a c ted

up ona s the food re a ches the regi onwhe re the intesti na l
j uice is sec re ted Th e o the r e nzyms of the i ntesti na l
.

j uice a re th ose which c onve rt c a ne sug a r (ordi na ry t a ble

sug a r) mi lk sug a r (one o f the c onstitue nts of milk ) a nd
, ,

th e m a l t sug a r wh ich is p roduced f ro m st a rch by th e

a cti on O f p ty a lin in to the simple sug a r gluc o se wh ich
'

, , ,

is the a ctu a l c a rb ohyd ra te th a t is m a de u se of by th e cells .

M O VE M EN TS O F T H E A LI M E N TAR Y T R A CT —For th e .

digestive p rocess to go on p rope rly th e f ood must be

mixed th oroughly with th e digestive j uices , a nd i t must
be p ropelled sl owly a l ong th e a lime nt a ry t ra ct f rom re
gionto regi ona s well The p ro cess begi ns i nth e m outh
.
,

whe re by th e a ct O f chewi ng eve ry p a rticle o f f ood is

, , ,

o r sh ould be b ro ugh t i n t o c ont a c t with th e s a liv a Th en

, .

c omes the p rocess o f sw a ll owi ng wh e reby th e well ,

m oiste ned f ood is p a ssed f rom th e m o uth i nt o th e

st om a ch Inth e st om a ch th e food m a ss is subj ected to
.

a ch u rn ing by the muscul a r a c ti ono f th e st om a ch w a lls ,

wh i ch serves to mix it well with g a st ric j uice a nd is th e n ,

expelled little by little i nt o th e sm a ll i ntesti ne This .

p rocess is a g ra du a l one a nd is usu a lly not c ompleted fo r

two h ou rs or m o re a fte r th e me a l is e a te n Th e f ood th a t .

e nte rs th e sm a ll i ntestine f rom th e st om a ch is a g a i n

chu rned by muscul a r m o veme nts of th e i ntesti ne to i nsu re
th a t th e p a nc re a tic j u i ce h a s a ccess to eve ry p a rticle a nd
is p ropelled sl owly a l ong by oth e r muscul a r m oveme nts .

Th e n a fte r th e d igested m a te ri a l h a s bee n a bs orbed a c

,

c ordi ng to th e desc ripti on i n th e nex t p a ra g ra ph th e ,

resid ue whi ch is o f n
, o fu rthe r v a lue is p a ssed i n t o th e
,

l a rge i ntesti ne wh e nce i t is d is ch a rged f rom th e b ody

, .

A B S OR PT I O N Th e di gested f ood withi n the a l i men

.
-
 ALI M E NTA R Y T R A C T MO V E M E N T
t a ry t ra c t is re a dy for use by th e livi ng cells Bu t i n
.

o rde r to re a ch them it must p a ss f ro m th e a lime n t a ry

t ra ct i nt o th e bl ood to be c a rried by th e bl ood to a ll p a rts
,

o f the b ody . The p rocess by which i t gets f rom th e

a lime n t a ry t ra c t i nt o the bl ood is c a lled a bsorp ti on I t
.

c onsists i nthe p a ss a ge of the m a te ri a l th ro ugh th e ve ry

thi n memb ra ne c onsisting of one l a ye r of cells which
, ,

l ines th e a li me nt a ry t ra c t a nd th ro ugh a sec ond thi n

,

memb ra ne which m a kes up the w a ll O f th e c a p ill a ry .

This p rocess of a bs orpti on t a kes pl a ce a lm ost wh olly i n

th e sm a ll i ntesti ne In esse nce i t is th e s a me p rocess
.
,

desc ribed i nCh a pte r V I a s the me a ns by which subst a nces

p a ss th rough pl a nt memb ra nes Whe neve r th e i ntesti ne
.

c ont a ins digest ed f ood i na ny qu a ntity the c once nt ra ti on

o f these f oodstu ffs withi n it is h ighe r th a n i n th e bl ood ,

an d c onseque ntly f oodstu ff s di ffuse f rom the i ntesti ne

int o th e bl ood The m a te ri a ls which e nte r the bl oo d
.

i n th e i ntesti na l c a pill a ries le a ve it sh ortly i n th e ca p il

l a ri es of th e tissues which r equi re nou rishme n t so th e
,

bl ood d oes not h a ve a ch a nce to bec ome l oa ded with

f ood stuff s to a n exte nt which w o uld i nte rfere with
”

a b sorpti on.
 C HAPTER XI

R E S PI R ATI O N A ND T HE DI S P O S AL OF WA S T E S

R S P I RAT I O N
E — FO I the s a me re a s on th a t a n e ngine
.
‘

requir es fuel wh o se oxid a ti on or bu rn i ng supplies the

drivi ng p owe r or e ne rgy so the V it a l m a chi ne depe nds,

up onf oods a nd a ste a dy supply o f oxyge nto oxidize them .

Th e highe r a nim a ls wh e n dep rived o f oxyge n die i n a

few mi nutes ; l owe r f orms like th e a m oeb a ma y live for , ,

twe nty fou r h ou rs but s oone r or l a te r p ro t opl a sm for

-
,

eve r e nds i ts a c tivities wh e ndep rived of a supply of this

subst a nce Oxyge n th e ref ore is a s esse nti a l to the life
.
, ,

Of a n a n im a l or pl a nt a s w a te r s a lts of v a ri ous ki nds , ,

sug a r a nd h ighe r c omp ou nds usu a lly cl a ssed a s f oods

,
.

One o f th e results o f th e ox id a ti on O f cell subst a n ces

is the form a ti ono f c a rb ondi o xid a nd w a te r th a t a s w a ste
p roducts esc a p e to th e exte ri or Th is p roc ess emb ra cing .
,

the i nt roducti on of oxyge n i nt o th e cell a nd the givi ng

O ff O f c a rb o n di oxid ga s is k n o w n a s resp i ra ti on
,
In .

highe r forms h oweve r wh e re v a st numbe rs of cells a re

, ,

p a cked t ogeth e r a ccess ory respi ra t ory st ructu res to c a rry

,

o xyge nto th e cells a n d to rem o ve th e c a rb ond i oxid f rom

them a re usu a lly p rese nt Th ey c omp rise such org a ns .

as gills o r lu n gs fi tted to a bs orb th e o xyge n f rom the

,

su rrou ndi ng medium a nd delive r i t to th e bl ood th a t i n ,

tu rn t ra nsfe rs i t to the cells He nce i n a dditi on to the

'

.
,

cell or i nte rna l respi ra ti on the re is th is a ccess ory or , ,

ex te rna l respi ra ti on
, .

D I FFU S I O N O F G A S ES I neve ry lite r o f a i r th e re a re

.

21 0 cubic ce ntimete rs o f oxyge n 7 90 cubic ce ntime te rs ,

of n i troge n a nd ,
cubic ce ntimete rs o f c a rb on di oxid
ga s. Inf resh w a te r eve ry l ite r c ont a i ns diss olved i n it
,

98
 R E S P I R AT O R Y O R GAN S 99

a b o u t 7 cubic ce ntimete rs o f o xyge n 1 5 cubic ce ntimete rs ,

of n it roge n a nd v a ryi ng sm a ll qu a ntities o f c a rb ondi oxid

,
.

Insea w a te r th ese subst a nces a re slightly less i na m ou nt

-
.

In respi ra ti on the p a ss a ge o f Oxyge n f rom th e a i r or

w a te r i nt o the bl ood or th rough the su rf a ce of a cell is
not a peculi a rity o f living th ings bu t f oll ows the l a w o f
the d iff usi ono f g a ses th a t is esse nti a lly the s a me a s the
d iffusi on of liquids I f we t a ke a cl osed ja r divid ed by
.

a thi n m o ist memb ra n e such a s a sheet of p a rchme nt

, ,

int o two equ a l c omp a rtments i n one of which th e re is

twice a s much ga s a s i n th e o the r th e m olecules w ill , .

p a ss th rough the memb ra ne u ntil th e p ressu re on ea ch '

side is the s a me The net result is a p a ss a ge of ga s f rom

.

th e c omp a rtme nt of highe r p ressu re or g re a te r a m ou nt

int o th a t with less u ntil th e density i n b oth c omp a rt
,

me nts is equ a l
-
.

Inthe cell whe re oxyge nis c onti nu a lly be i ng used a nd

the p ressu re is a cc ord i ngly less th a n i n th e su rro u ndi ng
a ir or w a te r or i n th e lu n
, gs wh e re th e a m ou nt p er u nit
v olume of this ga s i n th e bl ood is less th a n in th e a i r ,

o utside th e th i nm o ist lu n g w a ll di ffusi onb ri ngs a f resh

“
-
,

supply In th e s a me wa y th e a bu nd a nce o f c a rb on di
.

oxid i na cell le a ds to its diffusi oni n t o th e w a ter or bl ood

whe re the a m ou nt is less .

R ES P I R AT OR Y ORGA N S In u nicellul a r org a nism s

.

whe re the b ody su rf a ce is rel a tively l a rge a nd c omes i n

immedi a te c ont a ct with th e su rrou ndi ng medium (a ir or
w a te r ) no speci a l respi ra t ory org a ns exist Eve n i n
, .

highe r types such a s th e hyd ra j elly fi sh sp onges a nd

, ,
-
, ,

m a ny w orms livi ng i nm ois t pl a ces respi ra ti ont a kes pl a ce,

th rough the gene ra l b ody su rf a ce a nd th e needs of th e ,

u nde rlyi ng cells a re mi niste red to by di ff usi on with or ,

with out th e a i d of a ci rcul a t ory system ; Eve ni na nim a ls

of l a rge size wh e re respi ra t ory org a ns a re p rese nt th e
, ,

ski nma y c onti nue to t a ke a n a ctive p a rt i n the p rocess .

Inthe f rog for ex a mple the ski nis a lw a ys a ctive i nthis

, ,

reg a rd a n d du ring hibe rna ti on or the wi nte r sleep is th e

, ,
1 00 R E S P I R A TI O N AN D DI S P O S AL OF WA S T E S
s ole o rg a no f respi ra ti on Onthe o the r h a nd the re a re
.
,

m a ny a nim a ls e ncl osed i nshells o r h orny c a ses or sc a les ,

fe a the rs o r h a ir a s i nth e highe r types th a t a re ei the r

, , ,

too thick o r too dry to pe rmit of th e d iffusi on o f g a ses

th ro ugh th e ski n a nd a ccess ory respi ra t ory o rg a ns con
, ,

st ructed on v a ri ous pl a ns pe rf orm th e duty o f ga s , ,

exch a nge .

G ILL S —. Th e c onditi ons su rro unding a n a nim a l th a t

spe nds its life i n the w a te r a re ve ry evide ntly d iff e re nt
f rom th ose o f a l a nd dwelle r ; h e nce i t is no t su rp risi ng
-

to fi n d th a t th e speci a l respi ra t ory org a ns o f a qu a tic

f orms a re c onst ructed o n a di ff e re nt pl a n f rom th ose
‘

b re a th ing a tm o sph e ric a i r They usu a lly t a ke th e f orm

.

o f delic a te exp a n si ons or outgrowths of the ski n k now n

a s gi l ls . I n m a ny o f the w orms th e y a ppe a r a s g re a t
plume l i ke st ructu res b orne on th e h e a d or a s f e a th e ry
-

o r b ro a d pl a te li ke o rg a n
-
s a l ong th e sid e of th e b ody .

In s ome O f th e s n a ils th e gill resembles a m a n y pet a led -

fl owe r c a rried onth e b a ck o r exists i nth e f orm of delic a te

,

an d sle nde r o u tg rowth s widely dist ributed o ve r th e b ody .

I n the fishes they a re c omb like org a ns situ a ted a l o n -

g
the sides o f th e th ro a t In eve ry inst a nce th ese org a ns
.

a re pe n et ra ted by a nexte nsive c a pill a ry netw ork or sys

tem of bl ood sp a ces th a t a re thu s m ost f a v ora bly situ a ted
fo r receivi ng oxyge na n d disch a rgi ng c a rb ondi oxid Eve n
n
.

m ore f a v ora ble c o ndi ti o s a re p rese nt whe re th e w a te r

is kep t i nm o ti onby m oveme nts o f th e gi lls a s h a ppe ns ,

in ma n y species of w orms sh rimps a nd s ome of th e , ,

s a l a m a nd e rs for ex a mple
, .

R ES P I RATOR Y ORGA N S O F L A N D A N I M A LS — I n m ost

te rrest ri a l a nim a ls th e su rf a ce o f th e b ody is m ore or
less e ncl osed in a p ro tective c oa t or oute r skelet on th a t
reduces th e ch a n ces o f i nj u ry by e nemies he a t o r ligh t ,
, ,

an d a t the s a me time p reve nts excessive ev a p o ra ti on of

w a ter f rom th e b ody Fu rth e rm ore this i nvesti ng she a th
.
,

is not only rel a tively th i ck bu t i nm o st i nst a nces it is fa r

too dry to p e rmit th e re a dy exch a n ge of g a ses th rough
 T R A CH E AL S Y S TE M OF I N S E CT S 1 01

the skin Th e respi ra t ory org a ns o f l a nd a nim a ls the re

.
,

f ore a re built on a di ffe rent pl a n from th ose of a qu a tic

,
'

S pecies a n d ge ne ra lly spe a ki ng a re i n

,
g rowths o f the b ody
w a ll c ommu nic a ti ng with the o utside by na rrow p a s
,

s a ges a nd bu ried deep a m ong th e tissues of th e b ody

, , ,

a re kep t sufficie n tly m oist to meet the requi reme nts .

B re a th ing org a ns o f this typ e c omp rise two cl a sses : the

t ra ch e a l system of i nsects a nd rel a ted f orms a nd lu ngs , ,

chiefly a ch a ra cte ristic o f high e r a nim a ls .

TH E T R A CH EA L S Y S TE M O F I N S E CT S Ina l l but a few .

sm a ll i nsects respi ra ti onis c a rried onby a u nique mech a

nism k now na s th e t ra che a l syst em (Fig I f the b ody .

FI G 20 . .
— T h e T ra ch ea l S ys te m of th e C o c k roa ch .

of a g a ssh opp er, ex a mple I s c a refully ex a mi ned sev

for
‘

r
, ,

e ra l sm a ll openi ngs ca n be see n a l ong the S ides O f th e

b ody a nd fu rt he r i nvestig a ti on sh ows th a t e a ch O f these
,

le a ds i nt o a delic a te h orny tube th a t p e net ra tes the tis

sues a nd b re a ks up i nt o a l a rge numbe r O f exceedi ngly fi ne
b ra nches A swelli ng m ovement of th e b ody w a ll lik e
.
-

th a t of the h um a n ch est i n b re a thi n g d ra ws a i r i nt o ,

these c a na ls a nd dist ributes it to th e cells The O xyge n .

is the na bs orbed by th e cells a nd c a rb on di oxid is liber

a ted by th em a n d is th en disch a rged by a reve rse
c ont ra cti on .

Ins ome of th e Wa te r bugs a n d beetles th a t hu nt thei r

f ood i n p onds a nd gently ru nni ng st re a ms pe ri odic a l ,

visits a re m a de to the su rf a ce whe re a sm a ll qu a ntity o f

a i r is i mp ris oned u nde r the wi ng c ove rs or a m ong h a i rs -
1 02 R E S P I R ATI O N AN D DI S P O S AL OF \
VA S T E S
o nth e b ody a nd is b re a thed i nd uri ng the time o f sub
me rge nce Th e y o u ng of m a ny flies a nd beetles a re a ls o
.

a qu a tic a n d i nth e i nte rv a ls O f feed ing c ome to the sur

,

fa ce a nd b re a th e i na supply o f a ir Whe re such species .

a re a pest a s in th e c a se o f the m osquit o a t hi nl a ye r o f

, ,

o i l sp re a d o ve r th e su rf a ce o f the w a te r cuts O ff the a ir

supply a nd d est roys th e i nsect .

L U N GS Lu ngs a re i ng ro wths o f th e b ody su rf a ce or

.

o f th e digestive t ra ct wh ich a m o u n ts to th e s a me thi ng

, ,

th a t c ommu n ic a te with th e o u tside by c omp a ra tively

na rrow Ope ni ngs a nd on th e i nte ri or exp a nd to f orm a
sa c O f v a ryi n g deg rees o f c o mplexity In s ome O f th e .

s a l a m a nde rs liz a rd like sc a leless a nim a ls rel a ted to the

,
-

f ro g th e lu ng is me rely a sm oo th w a lled b a g I n th e
,
-
.

f rog a nd t oa d its i nne r su rf a ce is i nc re a sed by th e d evel op

me nt o f a few folds In still highe r species th ese f olds .

bec ome ext ra ordi na rily nume rous a nd of c omplic a ted

a rra n geme nt exp osi ng a l a rge a m o u nt O f su rf a ce to th e
,

a ir. Inma n for ex a mple th e Su rf a ce o f the lu n

,
g me a s ,

u res a b o u t 2000 squ a re feet or 1 25 times th a t o f th e ,

ge ne ra l su rf a ce O f the b ody Wh e n i t is remembe red .

th a t th e lu ngs a re exceedi ngly th i nw a lled a re kep t con -

,

ti n u a lly m oist a n d a re pe net ra ted by a ve ry exte nsive

,

c a pill a ry ne t w ork i t will be see n th a t th e hi gh a b sorp

-
,

ti o nof oxyge nmeets with a re a dy expl a na ti on .

Lu ng o r pulm ona ry respi ra ti onis a ss oci a ted with rh y th

mic a l m oveme nts o f the chest o r th ora x D u ri ng i nspi ra .

ti onth e th ora cic c a vity in wh ich th e lu ngs a re l o c a ted , ,

is e nl a rged th e lu ngs exp a nd to fill th e i nc re a sed sp a ce

, ,

an d a supply o f o xyge nc ont a ini ng a i r is th us b re a th ed i n

-
.

In exp i ra ti on th e ch est c a vity d ec re a ses i n v olume a nd

p resses th e a i r to th e o u tside S uch m oveme nts a s h a s .
,

bee n no ted a re p u rely a ccess ory a nd do no t c onstitute

,

respi ra ti o n .

R ES P I RAT ORY P I GM EN T S Bl ood pl a sm a is c a p a ble o f .

a bso rb i n g a b ou t a s much o xyge n as w a te r a nd i ts ca ,

p a c i ty in thi s reg a rd is fa r less th a nth a t o f hem ogl obin

t ,
 E X CR E TI O N 1 03

a c omplex reddish c omp o u nd c ont a i ni ng i rona nd c a rried

i nthe red c orpuscles U nde r ordina ry a tm osphe ric p res
.

su re this subst a nce f orms with oxyge na feeble c omp o u nd

th a t u nde r red uced p ressu re a s i n a n a i r pump a g a in
, ,
-

b re a ks up i nt o hem ogl obi na nd free oxygen Inth e lu ngs .

the ref ore oxyge n c ombines with the hem ogl obin a nd in
,

th i s c ombi ned st a te is tra nsp orted to th e tissue c a pill a ries .

He re the su rrounding lymph is p ra ctic a lly with ou t oxy

g e n ,the p r essu r e o r te nsi on is ne a rly ze ro a nd a s i nth e ,

a i r pump the o xyge nsep a ra tes f ro m th e h em ogl obi n dif

-
,

fuses th ro ugh th e ca pill a ry w a ll i nt o the lymph a nd from

the nce i nt o the cells .

Ins ome o f th e l o we r a n im a ls such a s the cl a ms s na ils

, , ,

c ra bs a nd thei r rel a tives c omp o u nds o f c oppe r a ct like

,

h em ogl obin i n the respi ra t ory p ro cess The re is a ls o .

so me evide nce th a t subst a nces c a rryi ng z inc a nd ma n

g a n ese m a y pe rf o rm a simil a r fu n cti oni nce rt a inspecies ,

but th e subj ect is one of g re a t c omplexity a nd is ve ry

impe rfectly u nde rst ood .

Inthe rem ov a l o f c a rb ondi oxid from the b ody th e c a se

is quite simil a r to th a t O f o xyge n Up ondi ffusing f rom .

the cell this ga s e nte rs the lymph di ffuses th rough the ,

c a pill a ry w a ll a lw a ys t ra velli ng i nt o regi ons o f l ow ten

,

si on a nd i nth e bl oo d f orms a l oose u ni onwith the pl a sm a

,

th a t rele a ses it u nde r th e dimi nish ed c a rb ondi oxid p res

su re of the lu ngs th rough wh ose w a ll s it m a kes its esc a pe .

E X CR ET I O N — Th e oxid a ti on o f cell m a te ri a ls results

.
,

a s h a s bee n desc ribed i n th e f orm a ti on o f w orthless

!

p roducts k nowna s exc reti ons Ina dditi onto c a rb ondi .

o xid ga s these i n
,
clude mi n e ra l s a lts w a te r a nd a con , ,

si dera b l e n umbe r of simpl e nit roge n c omp ou nds In .

th e a m oeb a p a ra m oecium a nd m a ny rel a ted u nicellul a r

, ,

an im a ls these subst a nces a re believed to a ccumul a te i n

,

on e or two defi n ite sp ots i n th e b o dy a l ong wi th w a te r

a bs orbed fro m the o utside At freque nt i nte rv al s this
.

c ont ra ctile v a cu ole a s it is te rmed empties th e d roplet

, ,

to th e exte ri or a n d thus fu ncti ons a s a simple exc ret ory

 1 04 R E S P I R ATI O N AND DI S P O S AL OF W A S TES
or a g n In othe r u nicellul a r a nim a l s a nd eve n i n the
.
,

hyd ra like a nim a ls a nd th e sp onges th e exc reti ons a re

-
,

rem oved f ro m th e b ody with o u t th e a i d of a n y speci a l

org a n s E lsewh e re i nth e a nim a l ki ngd o m th e re usu al ly
.

a re defi n ite exc ret ory org a ns c a lled kid neys th a t i nm a ny ,

speci a l c a ses coOp era te with org a ns h a vi ng o the r fu ncti ons

a s well Th e digestive t ra ct of w orms the l ive r of nu m
.
,

ero u s s n a ils cl a ms a nd ve rteb ra tes th e w a lls of th e he a rt

, , ,

an d neighb ori ng bl ood vessels a ll pl a y a mi n or th o ugh

O fte na ve ry imp ort a n t p a rt i nth e exc ret ory p rocess .

I n a few g ro ups o f a nim a ls a p orti on o f th e w a ste

p roducts a re no t rem oved f rom th e b ody bu t i nth e form ,

o f v a ri o usly c ol ored p igme n ts a re dep osited i n th e ski n .

Th e wi ngs of m a ny bu tte rflies owe thei r c ol or to such

c omp ou nds a nd th e silve ry shee no f a fish s b ody is like
,
’

wise d ue to a no th e rwise u seless subst a nce embedded i n

th e s ca les .

E X CR ETO R Y OR GA N S O F LO W ER A N I M A L S Th e di ffe r .

en t types of exc ret ory org a n s a m ong th e l owe r a nima ls

a re m a n y a nd th ei r m odes O f ope ra ti on a re equ a lly di
,

ve rse In th e fl a tw orms th ey t a ke th e form o f m a ny

.

widely sc a tte red cells which i n f orm a ppe a r much l i ke

highly b ra nch ed a m oeb a e The b ra nch es exte nd int o the
.

sp a ces betwee n th e tissu e cells c ollect th ei r w a stes i nt o

,

a d ro plet th a t is pe ri od ic a lly disch a rged i n t o a delic a te

tube with wh ich e a ch cell is i n c ont a ct A fl a gellum a t
n
.

the j u ncti onO f cell a nd tube d rives th e w a stes a l o g i nt o

l a rge r vessels th a t fi na lly ope n th rough the b ody w a ll -

by one or m o re p ores .

Inth e segme n ted w orms such a s th e e a rthw orm i nthe

, ,

s na ils cl a ms a nd th ei r a llies th e kid neys (one or m ore

, , ,

i nn umbe r ) p rese nt th e f orm o f m ore or less v olumi nous

tubes one end o f whi ch c ommu nic a tes wi th th e exte ri or
,

wh ile th e opp osi te ex t remity ope ns i nt o th e b ody c a vity .

Th e m a i np orti o nO f this o rg a nis eithe r di rectly b a th ed

wi th bl ood o r is supplied wi th c a pil l a ries f rom which the
w a s tes a re ext ra cted by th e kid ney ce lls a nd a re the reu p on
 KIDN E Y S OF HIGH E R ANI M AL S 1 05

emptied int o the cent ra l c a vity In m ost i nst a nces th e

.

i nne r end o f th e kid ney tube is p ro vided with h e a vy cili a

wh ose c onti nued be a ti ng d rives a ste a dy cu rre nt o f b ody
c a vity fluid th ro ugh the c a na l a nd thus w a shes the w a stes
to the exte ri or .

K I D N EY S O F H I GH ER AN I
'

M A S
L — The kid neys i n th e
.
‘

Ve rteb ra tes a re rel a tively l a rge p a i red o rg a ns a tt a ched

withi nthe b ody c a vity one a ch side of the b a ckb one A .

tube p a ssi ng f rom e a ch org a n le a ds to the bl a dde r or to

, ,

the exte ri or a nd onthe othe r h a nd c ommu nic a tes with a

,

rel a tively sm a l l c a vity withi nthe kid n ey itself (Fig .

FI G 21
. .
l ongi tu dina l secti onthro ugh hum a nkidney sh owing
A, ,
ra di a ti ng tub es o p eni ng into th e m a i n ca vi ty communi ca ti ng wi th
th e bl a dd er B kid n
. ey t ub es en
, l a rged C termina l secti onof tube
.
,
to sh ow cu p l ik e d e p ressi onfill e d wi th a ca pill a ry l oo p whi ch e x ten
-
ds
as a n etw ork o v er th e a dj a cen t s ecti on .

I nt o this sp a ce th e w a stes a re p ou red frp m the m a i n

kid ney m a ss th rou gh sm a ll ope ni ngs e a ch O f which com
muni ca tes with a c a na l th a t a s it m a kes its wa y i nt o th e
, ,

i nteri or b re a ks up i nt o a l a rge numbe r O f ve ry sm a ll

,

subdivisi ons The kid ney is th us m a de up of m a ny t ree

.

like t h ough h oll ow b ra nches O pe ni ng on th e one h a nd

,

int o th e m a i nkid ney c a vity a nd onthe o the r te rmi na ti ng

1 06 R E S P I R ATI O N AN D DI S P OS AL OF WA S T E S
bli ndly ne a r th e o ute r su rf a ce O f th e org a n Th e te rm ina l.

se cti on o f e a ch tube thus d evel oped is th e a ctive a ge nt

i n rem o vi n g exc reti ons a nd ca lls for a b rief desc ripti on .

Ea ch O f the th ous a nds O f kid n ey tubes e nds i na n eu

l a rgement resembli ng th e rubbe r tube a nd te rmina l bulb
,

O f a c a me ra with on e side o f the bulb p ushed i nto f o rm

a cup like dep ressi on

-
. I n th e kid n ey this cup is filled
with a twisted c a pill a ry l oop a nd th e tubul a r secti onfor
, ,

a c on side ra ble dist a nce is c ove red with a netw ork of

c a pill a ries.

M ODE O F O P ERATI O N — N 0 t only d oes e a ch te rmi na l

.

cup with its i ncluded c a pill a ry l oop di ff e r widely i nst rue

tu re f rom th e a dj a ce nt tubul a r secti on bu t th e p rocesses
,

c a rried on i n e a ch a re equ a lly u nlike I n the cup a .

st re a m c onsisti ng esse nti a lly o f lymph is c onti nu a lly

, ,

filt e ri ng from th e c a pill a ry i nt o th e c a vity of th e tube

d ow n wh ich it fl ows to the exte ri or Th is is a p u rely
.

physic a l p rocess .

At th e s a m e time th e cells of th e tubul a r secti on a re

a bs orbi n g f rom th e bl ood of th e c a pill a ry netw ork cer
t a i n w a ste subst a nces ch iefly nit roge nous th a t a re u m
, ,

a ble to filte r th ro ugh th e te rmi n a l cup Th ese a re p as sed

.

th rough th e cells of th e tube to th e ce nt ra l c a vity whe re

they diss olve i nth e bef ore me nti oned lymph st re a m a nd
-

a re w a s hed a w a y i nit .
 S EC T I O N 4
A DJ U S T M EN T OF OR GA N I SM S
 C HAPTER X II

H OW PLAN T S AR EAD J U S T E D TO TH E I R
E N VI R O N M E NT

TH E E N V I R O N M E N T . pl a nt or a nim a l ca nlive its

NO
life e nti rely to itself Th e sum t o t a l O f a l l the exte rna l
.

influe nces a cti ng ona ny org a nism c onstitute its e nvi ron
ment N O org a nism ca nesc a pe the i nfl ue nce of tempe ra
.

tu re a nd m oistu re Th e du ra ti on a nd i ntensity of ligh t

.

is very imp orta nt to a ny g ree npl a nt Th e a m ou nt a nd

.

dist ributi on O f ra inf a ll a nd th e fe rtility of th e s oil a re

V it a l fa ct ors Th e c ompetiti on o f o th e r pl a nts or the
.

a bu n d a nce of pl a nt e a ti ng a nim a ls ma y dete rmi ne

-

wh ether a sp ecies ca no ccupy a p a rticul a r h a bit a t or not .

Anim a ls a re s omewh a t less di rectly depe nde nt on th e

p hysic a l f a ct ors O f thei r su rro u ndi ngs th a npl a nts but a re
neve rtheless subj ect to them i ndi rectly Tempe ra tu re .

usu a lly p roves a n imp ort a nt f a ct or l imi ti ng th e sp re a d

of a n im a ls Th e ki nd O f pl a nts occupyi ng a ny regi on
.

dete rmi nes th e food supply a nd th e na tu re O f shelte r

a v a il a ble to a n im a ls A ve ry I mp ort a nt p a rt of a n a ni
.

m a l s e nvi ronme nt is i ts rel a ti on to othe r a nim a ls It

’
.

must c ompete with s ome of them for f ood a nd a v oid

bec omi ng f ood for o the rs .

T H E N E CE S S I T Y FOR AD J U S T M E N T — With so m a ny
eleme nts of th e envi ronme nt pl a yi ng imp orta nt rOl es i n
the life of pl a nts a nd a nim a ls it is Obvi ous th a t a d j usti ng
mech a nisms a re requi red to e na ble th em to secu re or
m a i nt a in f a v ora ble rel a ti ons with thei r su rrou ndi ngs .

Pl a nts need to be a ble to a d j ust th e g rowth o f roo ts ,

stems a nd le a ves so a s to pl a ce these org a ns i na f a v ora ble

,

p ositi on to p e rf orm thei r seve ra l fu ncti ons f or if these

,

1 09
1 10 AD ! U S T M E NT TO E N VI R O N M E NT

a dj us tme nts a re no t p romp tly a nd p rope rly m a d e the

pl a nt pe rishes .

All a nim al s requi re food Ina few i nst a nces they ca n

.

rem a i n st a ti on a ry a n d h a ve a supply b rough t to them

by cu rre nts of w a te r, bu t i n fa r th e m a j ority o f c a ses
they must h a ve org a ns o f l oc om oti onto go inse a rch o f it
an d org a ns o f speci a l se nse to rec og nize it wh e nthey a re
i nits neighb orh ood . Th ey need these a ls o to a v oid bei ng
e a te nby o the r a nim a ls .

P LA N TS AN D A N I M A S CO N TR A S T
L ED —. I n c onformity
with th ese di ff e re nces betwee na nim a ls a nd pl a nts i nth ei r
rel a ti o nto th e e n vi ronme nt the re h a ve c ome i nth e Co u rse
o f ev oluti onto be g re a t a n d st riki ng st ructu ra l a nd fu nc
ti o n a l di ff e re nces betwee nthem Inge ne ra l a nim a ls a re
.

highly m o tile a nd pl a nts a re no t Pl a nts h a ve st rong

.

rigid b odies a n d a nim a ls do no t . Th e h ighe r a nim a ls

h a ve devel oped h ighly speci a lized se nse org a ns by me a ns
o f which they a re a cutely a w a re O f m a n y eleme nts o f th e
e nvi ronme nt wh i ch m a k e little or no imp ressi o n on the '

pl a nt M ore ove r th ey h a ve devel oped complic a ted a nd

.
,

efficie nt ne rv o us systems a nd b ra i ns th ro ugh wh ich th e

stimuli a ff ecti ng th e se nse org a ns b ri ng a b o u t immedi a te
an d e ff e ctive resp o nse by the m o t or org a ns .

T H E AD J U S T M E N T O F R OOT S TO T H E S O I L — A pl a nt .

wh o se roo ts f a iled to a ssume a suit a ble rel a ti o nto the s o il

w ould ve ry sh ortly pe rish Th is d a nge r is h oweve r eff ec
. .

, ,

ti v el y gu a rd ed a g a i n st i nseve ra l w a ys .

G EOTR O P I S M . Th e f orce o f gra vity is one O f th e chief

d i rective s timuli fo r roo ts This d oes no t me a nth a t roo ts
.

me rely h a ng d ow nona cc ou nt of th ei r W eigh t I t re a lly .

me a ns th a t s ome ro o ts t a ke a p ositi onp o i nti ng inth e li ne

O f gr a vi ty ,
wh ile o th e rs t a ke a p ositi on a t righ t a ngles
to i t . I n b o th c a ses g ra vity is th e e ff ective stimulus .

N O m a tte r i nwh a t p ositi o na ge rmi na ti ng seed is pl a ced ,

i ts roo t will tu rn d ow n T wo th i n
. gs a re to be noted i n
respe c t to th is tu rn i ng o f a roo t i nrefe re nce to g ra vity .

Fi rst it is th e d i rec ti on a nd no t th e f orce which is th e

,
 CH E M O T R OP I S M 1 11

e ff ective stimulus S ec ond th e tu rni ng is b ro ugh t a b out

.
,

by g row th a nd no t by c ont ra cti on a s i n th e muscles o f ,

an im a ls S uch tu rni ngs by g rowth i n resp onse to the

.

directi onof g ra vity a re tech nic a lly know na s geotrop ism

( Fig .

CH E M OTR OP I S M N ext to di recti ng roo ts i nt o the s oil

.

is the need to di rect them to th a t p a rt o f it whe re w a te r

'

an d th e necess a ry s oil nut rie nts a re to be f o u n d i n the

m ost suit a ble qu a ntity I t is a f a mili a r f a ct th a t roots
.

a re likely ( to fi n d sewe rs i n a dry s oil a nd cl og them

FI G 22 . . Geo tro pism

R o o ts Fi g La , S h ows a b ea n wi th a
Of . .

y o u ng roo t m a rk ed i nto l engths of 2 mm e a ch a nd th enpla ce d i na .

h oriz onta l p os iti o n Lb, sh ows th e sa m e roo t a da y l a ter A com

. .

p a ris onof th e two figur es sh ows th a t th e grea tes t a m o unt of e l onga

ti onh a s ta k enpl a ce i nth e 6 mm r e gi o nim m e di a tely b ehi nd th e roo t
.

ca p InFi g I I a , th e ti p wa s cu t Off a l o ng th e li ne a b a b o u t 3 0 mi nu te s
. . .

a ter th e ro o t wa s pl a ce d i np o si ti o n
f I I b sh ows t h a t th e ro o t n
. . . ev er

th el ess e l o nga te d a nd be nt d ow nto w a rd th e e a r th Fi g II I a , sh ows

. . .

a from whi ch th e ti p wa s rem o v e d b efore b ei ng pl a ced i np o si

r oo t

t i on I t e l o n
. ga te d Fi g III b b u t i t did not b end T h e three exp eri
,
. .
,
.

m ents togeth er sh ow th a t th e s timulus O f gra vi ty is recei ved a t the

ti p b u t th a t th e resp on se by gro w t h o ccu rs i nth e sligh tly older p a rt .

with a g re a t m a ss of rootlets Th a t th ey a re se nsi .

tive to w a te r a nd diss olved s a lts ca n a ls o be re a d ily

sh ow n by expe rime nt i n th e l a b ora t ory If seeds a re .

pl a nted i n a vessel so th a t th e y ou ng roots g row ou t

th rough the b ott om i nt o dry a i r they tu rnb a ck i nspite , ,
112 AD ! U S T ME N T TO E N VI R O N M E N T

of th e stim ulus of g ra vi ty , t ow a rd th e wet s o il i nth e ex

p osed b o tt om O f the vess el Th ough the expe rime nt is
less e a sy i t ca nneve rth eless be sh ow n th a t roo ts w ill
.

tu rn a w a y f rom a p oor s oil t ow a rd a fe rtile one Th i s .

s ort o f resp onse is k now na s Chemo trop is m .

T H ER M OTR OP I S M is th e resp onse to d i ff e re nces O f tem

tu e T O s o me s o rts o f pl a nts th is is imp ort a nt
p e r a r .
.

For ex a mple inth e c a c ti o f a rid regi ons i t keeps th e roots

,

ne a r th e su rf a ce of th e s oil whe re they ca nbest t a ke a d

v a nt a ge O f th e sc a nty ra i nf a ll wh e nit d o es c ome U nder .
l

expe rime nt a l c onditi ons th ei r roo ts ma y b e i nd uced to

tu rnd ow na nd pe net ra te deeply i nt o th e s o il if it is he a ted
from bel ow .

OT H ER T R O P I S M S O F R OOT S R o o ts a re a ls o se n
. sitive
to th e di recti on of ligh t This is u seful i n s ome c a ses
.
,

p a rticul a rly inroo ts which origi na te on p a rts of a pl a nt

exp osed to light Th us onth e English ivy roots origi
.
, ,

na te on th e d a rk side o f stems next th e w a ll (Fig 23 ) .

onwh ich i t is g rowi n g whe re th ey ca nsecu re a h old a nd

,

f a ste n th e sh oot secu rely i n pl a ce .

I t ca nbe sh ow nexpe rime nt a lly th a t roots a re se nsitive

to th e d irecti on O f a cu rre nt o f w a te r Th ey will a ls o.

resp on d to a n elect ric cu rre nt a nd to sti ll o the r s orts o f

stimuli I t seem s p rob a ble h oweve r th a t m ost o f th ese
.
, ,

a re o f little o r n o c onseque nce i nna tu re Th ey a re men .

t i on ed h e re me rely to ca ll a tte n ti onto the fa ct th a t p ro t o

pl a sm O fte nh a s u nutilized p o ssibilities bu t wh ich migh t
c onceiv a bly s ometime be o f u se to it u nde r new c o ndi
ti ons Th is f a ct sh o uld be b orne i nmi nd whe nre a d i ng
.

th e ch a pte r o n E v oluti on i n refe re nce to th e questi on

wh eth e r th e e nvi ro nme nt re a lly p roduces th e ch a ra cte r
i s ti cs O f o rg a nisms o r wh eth e r the ch a ra cte ristics th ey
a l re a dy h a ve dete rmi n e wh e th e r a nd wh e re th ey ca nlive .

T H E AD J U S T M E N T O F STE M S is p rim a rily con t rolled by '

resp on ses to ligh t a nd g ra vity (Figs 24 U nlike

.
,

ro o ts
,
they grow a w a y f ro m th e ea rth a n d t ow a rd th e
li gh t I n a ve ry y ou ng seedli ng eve n wh e re th e stem
.
,
 T E ND R IL S AND TWININ G S TEM S 1 13

a nd root a re sc a rcely a millimete r a p a rt they resp ond ,

opp ositely S i nce th e cells o f b oth h a ve bee n only re

.

cen tly de rived f rom the s a me si n gle cell the re must be

a ve ry e a rly di ffe re nti a ti on o f cells i n this respect .

Likewise a l a te ra l b ra nch origi na tes withi n a mi nu te

,

f ra cti on o f a n i nch f ro m a s tem tip a nd y et sh ows a

-
‘

resp on se to light a nd g ra vity which is di ffe re nt f rom th a t

o f th e m a i n stem si nce the f orme r grows h oriz ont a lly
,

an d th e l a tte r ve rtic a lly The resp onse to ligh t is c a lled

.

p ho to trop i sm It se r ves t.o b r i ng stems i nt o the m ost

FI G 23
. . nth e Stem of English Ivy whi ch origi na te d on
R oo ts o ,

th e d a rk side ne x t th e chim ney onwhi ch th e pl a n t grew N o te th a t

.

t h ese roo ts gro w a w a y fro m th e li gh t a nd to w a r d th e m o is t b ri ck

chimney .

f a v ora ble p ositi on for exp osmg th e le a ves to light

an d a ir .

T E N DRI LS AN D T W I N I N G STE M S a re guided i np a r t by

light a nd gra v i ty b u t their speci a l ch a ra cte ristics a re
, _

ch iefly due to thei r re a cti on to c ont a ct i e thi gmo trop ,

. .
,

i sm . A twi ni ng stem i n the c ou rse of its growth c a uses

th e tip p orti onto swi ng a ro u nd i na ci rcle Cont a ct with .

a suit a ble supp or t stimul a tes g row th i n su ch a m a nne r

th a t th e c oils of the stem a re w ou nd a b out i t .
114 AD ! U S TM E NT TO E N VI R O N M E N T

Te nd ril s beh a ve s omewh a t simi l a rly Whe na te nd ril .

c omes inc ont a c t with a supp ort it ma y re a ct a cc ord i ng ,

to its n a tu re by twi n ,
ing a b ou t it or p ressi ng its end i nt o
firm c ont a ct a nd f orming a sucke r like a tt a chme nt -
In .

s ome c a ses g ra pe for ex a mple a s s oona s a fi rm a tt a ch

, ,

ment h a s bee nm a de this itself a cts a s a stimulu s to c a use

a ce i l ing o f th e i nte rmedi a te p a rt of th e te nd ril so as to

FI G 24
. . Geo tro pism of Th ese b ea n pl a nts w ere grown
S tems .

i n a po t whi ch wa s k e p t righ t sid e up u ntil th e y w e re a b o u t 3 i n

- .

high a nd th enpl a ced oni ts si d e N o te th a t th e s tems h a ve p romp tly

.

be nt upwa rd to b ri ng th ems elves i n the li ne of gra vi ty .

d ra w th e m a i n stem i nt o cl ose r rel a ti o with th e n

supp ort .

A D J U S T M E N T OF L EA V ES is chiefly a cc omplish ed by ligh t

stimuli It is e a sy to dem onst ra te th is expe rime nt a lly
.
.

I f a p o tted pl a nt is pl a ced a few feet a w a y f ro m a ligh ted

wind ow a nd sh ielded f rom ligh t from o the r s o u rces a ll ,

its le a ves will p rese ntly be f ou nd f a cing th e wi nd ow

 AD J U S T M E N T O F R E PR O DU C TI O N 1 15

(Fig . If th e p ot is tu rned so a s to f a ce them a w a y

from th e ligh t th e le a ves will i n a sh ort t i me g row
,

a rou n d to f a ce it a g a i n E ve ry one is s omewh a t f a m ili a r

.

with this f a ct ou t of d oors The le a ves O f the ivy onthe

.

w a ll a re a ll f a ced ou t a nd a rra nged like a m os a ic so a s

to sh a de on e an othe r a s l i ttle a s p ossible Th e le a ves .

O f sm a ll pl a nt s like th e pl a nt a i nor d a ndeli onsp re a d ou t

i n a rosette f a ci ng upw a rd to the ligh t
,
It is less e a sy .

to obse rve th e le a v e s on t rees bec a use one is ra rely i na

FI G 25 . . Th ese pl a nts w ere growni na p ot pla ced nea r a wind ow

i n a n o t h erwis e d a rk en e d roo m I t will b e n
. o te d t h a t th ey h a v e

growni na di recti onwhi ch is s om ewh a t of a comp romise b etw eenth e

p osi ti on whi ch t h ey w ould h a v e ta k en i n resp onse to th e s timulus
o f gra vi ty a l o ne (i e if t h ey h a d b e en e qu a lly li gh te d o n a l l sid es )
. .

an d t h a t whi ch t h e y w o uld ta k e if i nflu ence d by t h e dir ect i o nof t h e

ligh t f rom one side only .

p ositi on to see them f rom th e di recti on of the i ncide nt

“

light but wh enh e is so situ a ted it will be see n th a t the

,

le a ves do fa ce the light .

AD J U S T M E N T O F R EP R OD U CT I O N This s ort of a dj ust .

ment is not so obvi ous to c a su a l i nspecti on bu t i t is ,

neve rth eless vit a lly necess a ry T rees which p roduce .

fl owe rs i n wi nte r a s th ey occ a si ona lly do h a ve sm a ll

,
'

,
1 1 6 AD ! U S T M E N T TO E N VI R O N M E N T

ch a nce o f m a tu ring seed F rom c onside ra ti ons o f th is .

s ort it is pl a in th a t the tree d oes re a ct to clim a tic c ondi

ti ons i n such a wa y th a t the rep rod uctive p rocesses a re
p rop e rly regul a ted Wh e npl a nts a re t a ke nf rom one re
.

gi on to a nothe r it no t i nf reque ntly h a ppe ns th a t they

g row vig orously but f a il to fl owe r or to set seed So me .

O f the n ecess a ry stim uli a re w a nti ng Th e c li m a te ma y .

FI G . 26 .
—A R ose tte of nged a s to expo se ea ch
Lea v es so a rra
lea f as fu lly a s p o ss ibl e to ligh t Al th o ugh n o t so o bvi o us the lea v es
.
,

o f m o s t pl a nts a re a ccu ra te ly pl a ce d to ta k e th e bes t a dv a nta ge o f

th e a v a il a ble ligh t .

be too wet or too dry too h o t or too c old th e ligh t too

, ,

inte nse or too we a k or s ome ch a ra cte ristic of the s oil

,

ma y be u nsuit a ble .

L a ck of w a te r stimul a tes rep roduct i onInm a ny ki nd s

of pl a nts Am ong th e Alg a e a re m a ny which h a ve two
.

meth ods o f rep roducti on a sexu a l a nd a nonsexu a l one ,

-

(Ch a p XX V I ) Whe nw a te r is f resh a nd a bu nd a nt they

. .
 IN T E R NAL C O R R E LATI O N S IN P LAN T S 1 17

g row ra pidly a nd set f ree multitudes of nonsexu a l si ngle -

m o tile rep roductive cells c a lled swimmi ng sp ores c a p a ble

O f immedi a te g ro wth in to new pl a nts On th e c ont ra ry .
,

whenw a te r bec omes sc a rce or u nsuit a ble for g rowth they ,

f orm by sexu a l p rocesses a ki nd O f resti ng sp ore which

is not m o tile a nd is su rro u nded by a h a rd thick w a ll th a t
p ro tects it wh e nth e p ond d ries up This sp ore p roduces .

a n ew pl a n t the next se a s on .

I N TER N A L C ORR EL ATI O N S I N P LA N T S a re a s necess a ry

a s thei r a dj ustme n t to the e nvi ronme nt A g re a t m a ny .

thi ngs which a re ordi na rily t a ke n for g ra nted a re i n

re a lity th e r esul t o f a ccu ra te c orrel a ti on T rees f orm .

myri a ds o f le a f buds eve ry ye a r bu t only a f ra cti on o f

,

them uri fol d i nth e sp ri ng If h oweve r th e fi rst c rop of

.
, ,

le a ves i s destroy ed th e n a nothe r l ot of buds u nfolds to

'

repl a ce them C le a rly eithe r th e u nf oldi ng of th e fi rst

.
,

buds must h a ve a cted a s a rest ra i ni ng i nflue nce on the

o the rs or thei r a bse n
,
ce a cts a s a stimulus for the cor ,

rel a ti onis a nevide n t f a ct .

I t is a c omm onpl a ce th a t roo ts devel op a t th e l ower end

o f cutti n gs a nd le a ves a t th e top N or d oes it a lte r th e
.

result if th e origi n a lly uppe r p a r t o f th e cutti ng be the

on e stuck i nth e s oil . This is no d oub t due i nl a rge p a rt
, ,

to th e f a ct th a t th e two e n ds a re subj ect to di ff e re nt

stimuli bu t not e nti rely so si nce roo ts do not devel op a t
, ,

a l l p a rts W h ich a re i nc ont a ct with th e s oil a nd so r eceive

th e sa me stimuli .

T H E M E C H A N I S M OF T R O P I S M S The recepti onof a n

.

exte rna l stimulus dep e nds on th e se nsitiveness or i rrit a

b i li ty o f the p ro t opl a sm Wh eth e r a resp onse will f oll ow
.

depe nds on th e c onducti ono f this stimulus to s ome p a rt

of the s a me cell or to o the r cells a t a dist a n ce wh e re s ome
o the r a ctivity o f p ro t opl as m ca nbe excited I nth e t rop .

ism of roots a nd stems th e stimulus is received by cells

ne a r th e tip a nd s ome s ort of ch a nge is p rop a g a ted
,

th rough seve ra l cells to a regi onfu rthe r b a ck wh e re th ere

a re i n duced g rowth ch a nges wh ich c a use the roo t to bend .
11 8 AD J U S T M E NT TO E N VI R O N M E N T

The be ndi ng is d ue to th e f a c t th a t th e cells o none Side

g ro w f a ste r th a n th o se o n the o th e r
.

The na t u re O f th e ch a nge is still s o mewh a t u nce rt a i n .

The re is s ome evid e nce to sh ow th a t it is a n imp ulse o f

s ome s ort c omp a ra ble to a nelect ric cu rre nt but ce rta i nly
not ide ntic a l with it C e rt a in oth e r f a c ts seem to p oi nt
.

to a w a ve o f ch emic a l ch a n .ge Instill o the r c a ses the re

is i ndic a ti on O f the a ctu a l m oveme nt O f a defi nite a nd
peculi a r subst a nce f ro m th e stimul a ted to the re a cti ng
regi on . Ina nim a ls subst a n
,
ces O f th is na tu e a e b eco
r r mi

in g f a i rly well k now n a nd a re c a lled h orm on es

.

T H E M E CH A N I S M O F C ORR E LAT I O N I N P LA N T S is eve n

less well u nde rst ood th a n th a t o f t ropism The re is
.

g ra du a lly a ccumul a ti ng a c onside ra ble b ody of f a cts

which seem to p oi nt to the m oveme nt O f h orm one sub
st a nces f ro m one p a rt of th e pl a nt to a nothe r wh ich a re
n
,

c once r ed i n c orrel a ti on (c omp a re C h a p X V I )

. .
 C HAPTER XIII

M O VE M E N T I N ANI M AL S

M O VE M E N T I N PR OTO Z OA Th e P rot oz oa a s we na me
.
,

th e one celled a nim a ls live i n w a te r whe re they eithe r

-
,

m ove a b o ut f reely or a re a tt a ch ed by s ome pa rt of thei r

p rot opl a sm to s ome s olid obj ect As we st a ted i n a n ‘

e a rlier ch a pte r (I I I ) th e re a re ve ry m a ny diffe re nt

,

ki nds O f P ro t oz oa which di ffe r gre a tly i n a ppe a ra nce a s

well a s h a bits We ca n disti nguish a m ong a l l of them
.

fo u r types of m oveme nt which se rve th e e nti re cell a s ,

disti nguished f rom m ovements of p rot opl a sm withi n

the cell .

A M (E B OI D M OTI O N Th e fi rst o f these is th e ch a ra c

.

teri sti c f orm o f m o ti onof th e ve ry simple org a nism know n

a s th e a m oeb a (Fig .and th is ki nd o f m o ti onis th e re ,

fore c a lled a m oeb oid m oveme nt Th e m o ti onc onsists o f

, .

a th rusti n g ou t of the p rot opl a sm i ns ome di rec ti onwith

a n equiv a le n t d ra wi ng i n of o th e r p a rts so th a t the ,

v olume o f th e cell is not ch a nged a lth o ugh th e o utli ne ,

of the cell is con ti nu a lly ch a nging Th e m oveme nt is not

.

ra pid . An a m oeb a usu a lly h a s to be w a tch ed c a refully

u nde r the mic ro sc ope to m a ke ou t wh ethe r it is m ovi ng
or n ot .These a m oeb oi d m ovements a pp e a r to be pu rely
ra n d om yet if a n a m oeb a is obse rv ed for a pe ri od O f
,

time it is usu a lly see nth a t it m a kes p ro gr ess i none di rec

ti on S O th a t ou t th ru sti n
,
‘
-
gs i n th a t di recti on must h a ve
bee nonth e wh ole m ore freque nt th a ni na ny oth e r We .

a pply th e exp ressi on a m oeb o id m o veme n t to a ny cell

m oti ons i n which th e p rot opl a sm is th rust ou t b odily
CI LI AR Y M OTI O N Th e sec ond ki nd of m oti on is
.
-

k now n a s cili a ry It receives th is na me f rom th e f a ct

.

1 19
1 20 M O V E M E N T I N ANI M AL S

th a t th e m oti onis a cc omplished by th e wh ipp ing of ve ry

ti ny th re a ds O f p rot opl a s m which p roj ec t from th e m a in
su rf a ce of th e cell Th ese p rot opl a smic th re a d s a re c a lled
.

cili a (s i ngul a r cil ium ) f rom their resembl a nce to t iny

,

h a irs (Fig .Th ey whi p ra pidly b a ck a nd f orth ,

FI G 27
. . Pa ra mmci u m or sli ppe r a nima l cu le sh ow i ng cili a
, .

eithe r i nu nis onor i nregul a r successi on th e m o ti onbei ng ,

a lw a ys m o re p o we r ful i no ne di recti onth a ni nth e o th e r .

Thus the org a nism if a free f orm is p ropell ed th rough

, ,

th e w a te r In P ro t oz oa th a t a re fixed to s ome O bj ect

.
,

th e cili a set up cu rre nts i n th e w a te r wh o se eff ec t is to

sweep th e food p a rticles i nt o c ont a ct wi th th e p ro t opl a s m
whe re th ey ca n be t a ke n i n .

F LA GELLATE M OT I O N — Th e th i rd typ e O f m oveme nt

is c a ll ed fla gell a te m o ti on In this f orm th e org a nism
.
 CO NT R A CTI O N 1 21

ha s at s ome p oi nt onits su rf a ce one or two p rot opl a smic

p roj ecti ons which a re l a rge r a nd st ronge r th a n cili a a nd
which a re c a lled fla gel l a si ng ul a r fla gell um
, (Fig .

Th ese fl a gell a s weep b a ck a nd f orth p ropelli ng the

org a n ism f orw a rd ve ry much a s a fish p ropels itself by
st rokes of i ts t a il .

CO N TR A CT I O N O F P R OT OP LA S MI C STR A N D S . The f ou rth

type of m oveme nt is f ou nd i nP ro t o z oa th a t a re a tt a ched ,

an d se rves to s a ve them
from th re a te ned h a rm In .

this type o f m oti on the re

ca n be see n i n th e p ro t o

pl a sm one or m ore thick

e ned s tra nds which re a ch
from the sp ot whe re the or
g a n i sm is a tt a ched o u t t o

w a rd th e f ree end (Fig .

I t is by the sh orte ni ng of
the p ro t opl a smic st ra nd or
st ra nds th a t the b ody is
pulled d ow n ou t O f h a rm s ’

wa y . This f orm o f m oti on .

is i nte resti ng bec a use th e

st ra nd o f p ro t opl a sm whi ch
FI G 28 C hl a myd o m o na s a
h a s the p owe r of sh orte ni ng one—
,
. .

cell e d pl a nt sh o wi ng fl a gell a.

r ep rese n ts the begi nni ngs

of th e me th od by which m o ti on is c a rried on i n a l l th e

highe r a nim a ls na mely th rough muscul a r c ont ra cti on

, ,
.

M OTI O N I N M A N Y CE LLED A N I M A L S — In the h ighe r

-
.

an im a ls b o th a m oeb o id a nd cili a ry m oti ons occu r to s ome

exte nt a nd i nm a ny i nst a nces a cc omplish ve ry imp orta nt
,

e ff ects ; For ex a mple i nou r ownb odies th e wi ndpipe a nd

,

b ronchi a l tubes a re li ned with cells which h a ve cili a on

th e su rfa ce f ronti ng onthe tubes These cili a whip con
.

sta n tly i n such a wa y a s to sweep up t ow a rd the th roa t

an y muc o us o r dust p a r ticles th a t m a y l o dge o n the m o ist
su rf a c e of the respi ra t ory p a ss a ges O e ca nre a dily a p
. n
1 22 M O V E M E N T I N ANI M AL S

p re ci a the imp ort a nce o f this c onti nu o us cle a ri ng ou t

te
o f th e ch a nnels by wh ich a i r e nte rs th e lu ngs Th e m ost .

imp ort a nt inst a nce o f a m oeb oid m oveme nt i nth e hum a n

b ody is i n c onnec ti o n wi th the w a nde ring O f th e white
bl ood c orpuscles wh ich h elp to p rotect th e b ody a g a i nst
,

i nfec ti onby seeki ng o u t a nd e ngulf

in g dise a se p ro d uci ng o rg a nisms -
.

The wa y i n which these c orpuscles

get f rom pl a ce to pl a ce a m ong th e
cells o f th e b ody 1 8 th ro ugh
a m oeb oid m oveme nt The chief .

o rg a n s of m oveme nt h oweve r , ,

a re

M U S CLES — All the l a rge m ove

.

me nt s o f the h ighe r a nim a ls a re

c a rried on by speci a lized mu scle
cells Th ese will be desc ribed b e
.

l ow He re we wish only to sa y th a t
.

t hey rep rese n t a m a rked ex a mple

o f the divisi on o f l a b o r wh ich wa s

FI G 29. .D i a gra m o f discussed i n Ch a pte r I I I M uscle .

t h e P o to z oa n Vo ti
a re c omp o sed o f livin
r r

ce ll a sh o w n i n t he
,
cells g p ro to

pl a sm which h a s the c ont ra ctile

,

no rm a l e x te nd ed p osi
ti o n a n d a ls o p a rti a lly
,
p rope rty highly d evel oped a n d th e
co n tra c te d T he p o r

s tra nd by oth e r p rope rties c orresp ondi ngly re

.

to p la s mi c
whi ch o nt a ti o nis p ro
c r c
du ced The fu nc ti ona l met a b olism
du ced is sh o w n e x te n d
.

in d ow n t h e s te m O f muscle cells c on t
sis s i n th e p er
g .

formi ng o f these c ont ra cti ons i n ,

o the r w o rds i n the c on

,
ve rsi on o f chemic a l e ne rgy
i nt o th e e ne rgy of m oti on .

T H E B I OLOGI CA L I M P ORTA N CE O F M OT I O N — Bef ore

-

g oi ng onto a fu rthe r desc ripti ono f muscul a r a cti on let ,

u s t a ke a m ome nt to i nqui re i nt o th e utility to a nim a ls

o f th e p o we r to m o ve .

(1 ) P rima ri ly M o ti oni nAni ma ls is a M ea ns of S ecu r

i ng Food — All a nim a ls requi re fo r thei r f oo d c o mplex
subst a nces which res ul t eithe r di rectly or i ndi rectly f rom
 B I OL O GI CAL I M P O R TAN CE OF M O TI O N 1 23

th e a ctivities o f the g ree ncells o f pl a nts A nim a ls eithe r .

m ove to pl a ces w he re these p rod ucts a re Or else c a use the

p roducts to c ome to them As a n ex a mple of the fi rst
.

type ma y be me n ti oned a ny g ra zi ng a nim a l which g oes

whe re g ra ss is a nd the n e a ts it As a n ex a mple O f th e
.

sec ond t a ke the oyste r wh ich is a nch ored fi rmly to th e

ro ck a n
,
d by m oveme nts o f cili a once rt a i nof its su rf a ces
withi n the shell c a uses a cu rrent o f w a te r to sweep
th rough its gill c a vities in such a wa y th a t minute f ood
p a rticles which the w a te r ma y c ont a i nca nbe sifted ou t
an d c onsumed .

( )
2 M o ti on i n t he S tru gg l e fo r E x i s te nc e — Wh e re
an im a ls a re a t a l l nume ro us c ompetiti on e nte rs i nt ro ,

du ci ng the eleme nt o f st ruggle eithe r of two a nim a ls,

for th e s a me pl a nt f oo d or o f s ome a nim a ls to use o th e r

an im a ls for f ood He re we fi nd a sec ond bi ol ogic a l neces
.

si ty f or m o ti on n ,
a mely the a v oid a n
,
ce O f i n j u ry M O .

ti ons for this pu rp ose ma y t a ke th e f orm eithe r of fligh t

or c o mb a t . Cl o sely rel a ted to these a re m o ti ons requi red
for m a ki ng a dj ustme nts to clim a tic c onditi ons A se a rch.

for sh elte r f ro m th e eleme nts or for w a rm qu a rte rs i n

wh ich to sleep a re ex a mples .

( )
3 M o ti on a n d P e rp e tu a ti o n o f t h e R a ce — N ext . i n
orde r cOmes a serI e S o f m o ti on s which a re rel a ted i mmedi ,

a tely or rem o tely to th e p e rpetu a ti on O f the ra ce

,
Th e .

pu rsuit O f a m a te st rife with c ompetitive suit ors the

, ,

ca re o f th e y o u n g a ll i nv olve m a ny m o ti ons a nd a ll h a ve
,

immedi a te be a ri ng onra ci a l pe rpetu a ti on Of m ore re .

m ote c onnecti on yet u nd oubtedly bel ongi ng he re a re the

, ,

m oti ons by which th e y ou ng o f b oth sexes st rive to m a ke

themselves a tt ra ctive Am ong civilized men the re a re
.

m a ny a ctivities wh ich h a ve to do with ra ci a l pe rpetu a ti on ,

a lth o ugh th e rel a ti onis O fte nn ot O bvi ous For ex a mple

.
,

the s a c rifi ces o f p a rents i norde r th a t th ei r child re nma y

be p rope rly educ a ted c ome u nde r the h e a d of c a re of the
y ou ng j ust a s t ruly a s d oes the t oil of a m othe r f or h er
i nf a nt s welf a re .
1 24 M O V E M E NT I N A NI M AL S

(4 ) T h e M o ti ons of P la y Fi na lly we must not

.
,

omit to m a ke me n ti onO f th e m o ti ons which a re m a de in

pl a y Th ese a re p a rticul a rly ch a ra cte ristic O f th e y o u ng
bu t a re by no me a ns m
.
,

issi ng i na d ults eithe r a m ong th e ,

highe r a nim a ls or ma n A f a c t th a t is w orthy O f con

.

side ra ti on is th a t wh il e the m o ti ons of th e fi rst th ree

g roups a re p ra ctic a lly u nive rs a l a m o ng a nim a ls i t is ex ,

tremely d ifficult to detec t with ce rt a i n ty a ny m oti ons of

pl a y except i n th e h ighe r a nd m ore intell ige nt f orms .

M o st ki nds of m a mm a ls pl a y bu t it w ould be h a rd to sa y
,

ce rt a i n ly wheth e r fish ever pl a y a nd one w o uld ce rt a i nly

,

be j ustified i n d oubti ng th e occu rre nce of pl a y a m ong

eithe r w orms or sta rfi sh es .

T H E R EGU LAT I O N O F M OT I O N (a ) R ecep o s

t r .
— It .

is quite evide nt th a t the me re p owe r to m a ke m oti ons h a s

ve ry little bi ol ogic a l imp ort a nce In orde r th a t m oti ons
.

sh a ll be effective they must be a d j usted to th e requi re

me nts of the situ a ti on Th is me a ns th a t the org a nism
.

must h a ve k nowledge of i ts e nvi ronme nt M o ti ons ca n .

not be di rected t ow a rd secu ri ng f ood u nless th e p osi

ti ono f th e f ood ca ni ns ome wa y imp ress itself a s a guide
fo r th e m o ti on I nge ne ra l te rms th e situ a ti onor en
. vi
,

ro n ment m ust h a ve a me a ns O f influe nci ng th e org a nism .

This c onsists i nxa l l a nim a ls except th e l owest of speci a l

, ,

org a n s O f one ki nd or a nothe r wh ich a re k now n a s th e

se nse org a ns AS a c onve nie nt si ngle w ord a pplied to
.
,

a l l th e v a ri o us ki n ds o f se nse org a ns th a t exist th e te rm ,

recep tor h a s bee n a d opted .

(b ) Adju s te rs Th e recepti ono f imp ressi ons f rom the

. .

su rro u nd i ngs will not by itsel f b ri ng a b o u t e ff ective mus

’

on l a r m oti on An a dd iti ona l step i s necess a ry na mely

.
, ,

th e est a blishme n t of c onnecti ons betwee n th e recept ors

an d th e o rg a ns th a t a re to c a rry o u t th e m oveme nts In .

o the r w ord s th e re must be a me a n s o f c ont rolli ng th e

m o ti o ns i nsuch a wa y a s to p roduce a suit a ble a nd effec
tive muscul a r resp onse a s a result O f th e imp ressi onm a d e
up o n th e recept o rs Th e a pp a ra tus for exe rcising this
.
 T YP E S O F E FF E CT O R S 1 25

c ont rol is known tech nic a lly a s th e a dju s te r mech a nism ,

which i n a ll th e highe r a nim a ls c onsists Of the ne rv ous

'

system .

(a ) Effectors
. Fina lly we h a ve the a pp a ra tus which
.

m a kes th e m oti ons to which i s given th e ge ne ra l

,

na me of efiectors Th e a dv a nt a ge of the w ord e ffect or “

”
.

ra the r th a n th e w ord muscles is th a t we h a ve eve n

“
,

i ns ome of the highe r a nim a ls m oveme nts c a rried onby ,

me a ns of cili a a nd a b roa d te rm like e ffect ors c ove rs such

,

m ovements while th e w ord muscle w ould f a il to do so .

TY P E S OF E FF E CTORS F rom th e p recedi ng p a ra g ra ph

.

i t is cle a r th a t cili a must be i ncluded i na compl ete listi ng

'

of th e ki n ds of e ffect ors f ou nd i n th e highe r a nim a ls .

For ou r p resent p u rp ose h oweve r i t is desi red to c onee n

, ,

t ra te th e a tte nti on on th e m ore highly Speci a lized a nd

m ore f a mili a r ki nds O f e ffect ors th e muscles ,
.

I nthe highe r a nim a ls th e re a re th ree distinct ki nds of

muscle tissue which will be menti oned b riefly he re a nd
desc ribed i nm ore det a il fu rthe r on
'

S K E ETA L
L M U S CLE S — U nde r this designa ti on a re in
.

cl uded mus cles wh i ch m ove th e b ones of th e skelet on ,

hence the na me Th ese muscles m a ke up th e g re a t bulk

.
'

O f ou r muscle subst a n ce They a re quick a cti ng a nd .

-

p owe rful a nd a re r esp onsible f or ou r m a in b odily

,

m oveme nts .

S O OT H
M M US C SLE —The sec ond type o f muscle
.
,

k now na s sm ooth muscle c a rries onm ost of th e i nte rna l ,

m oveme nts of th e b ody This kind O f muscle is much .

m ore sluggish th a nskelet a l muscle a nd is a ls o c omp a ra ,

ti v ely feeble In m o st o f the l owe r a nim a ls this is the

.

On ly typ e of muscle th a t is p rese nt Inthe h igh e r a ni .

m a ls it is f o u nd i n th e w a lls of the a lime nt a ry t ra ct i n ,

th e w a lls of th e bl ood vessels a nd i na few o the r pl a ces ,

wh ich need not be listed h e re .

H EAR T M U S CLE — In a ll of th e highe r a n im a ls th e

he a rt muscle di ffe rs from a ny othe r ki nd Of muscle a nd ,

must the ref ore be g i ve n a sep a ra te cl a ssific a ti on I t is .

1 26 M O V E M E N T I N A NI M AL S ‘

like th e skelet l muscle a inbei n

q uick a cti ng a nd p owe r
g -

ful ; it d iff e rs f rom a ll o the r ki nd s O f muscle i nits p rope rty

O f c on t ra c ti ng a nd rel a x ing a u t om a tic a lly I n a p a ra .

g ra ph a b ove wa s emph a sized the imp ort a nce of the con

t ro l of muscul a r m o veme nt This c ont rol requi res th a t
.

the muscles i nge ne ra l sh a ll be ina c tive excep t wh e n a o

ti v i ty is especi a lly c a lled fo r Th e h e a rt p rese nts qu ite
.

a di ff e re n t c a se si nce its p rope r fu ncti oni ng a s a norg a n

,

fo r pumpi ng bl oo d requi res th a t its m uscl e sh a ll c o nt ra ct

an d rel a x i n regul a r seque nce C o rresp ondi ng to this
.

diff e re nce i n req ui reme nt we fi nd th e d i ff e re nce i n b e

h a vi or a l re a dy me nti oned na mely th a t th e h e a rt muscle
, ,

w orks a ut om a ti ca lly wh ile th e o the r types o f muscle a re

,

u nd e r c ont rol a nd w ork only wh e n c a lled up onto do so .

M U S CLE F I B ERS All mu scles a re c omp osed of livi ng

.

cells the sh a pe of th e cells bei ng i n ge ne ra l cyli nd ric a l

, ,

th a t is th ey a re c omp a ra tively l ong a nd sle nde r For

,
.

thi s re a s o n they a re c omm on ly refe rred to a s muscle

fibe rs . The size v a ries Si ngle cells or fibe rs o f skelet a l
.
, ,

muscle ma y ra nge i nle ngth fro m 1 mm to 3 5 m m . .

i nch to 1 i nch ) ; a nd i n d i a mete r f rom 034 mm to ,

. .

.0 55 mm . i nch to i nch ) .

In th e high e r a n im a ls sm oo th mu scle fibe rs a re much

sm a lle r a ve ra gi ng no t m o re th a n 0 4 2 mm
, .i nch ) .

i n le n g th but i n th e l o we r a nim a ls i n which th ey con

, ,

sti t u te t h e chief type o f muscle subst a n ce they ma y ,

a tt a i n to l a rge r sizes .

M A K E U P O F S K ELETA L M U S CLE — Th e fibe rs a re

-
.

b ou nd t oge the r i n bu ndles or sh eets O f v a ri o us ki nds to

f o rm th e mu scle a s a wh ole A g o od id e a o f th e st ructu re
.

o f th e skelet a l muscles ca nbe O bt a i n ed f rom a nex a mi na

ti on O f ra w le a n m e a t wh e re it ca n be m a d e o u t th a t
,

i ndivi du a l fibe rs a re h eld t o geth e r i n bu ndles by me a ns

o f a bi n di ng tissue k now n a s c onnective tissue These .

sm a ll bu ndles a re b o u nd togeth e r i nt o s till l a rge r bu ndles ,

an d th ese a g a i ni nto still l a rg er th e c onne ctive tissue b e

,

c omi ng c o a rse r a s th e bu ndles th a t a re to be b ou nd to

 S M O O TH M U S CL E AN D H E A R T M U S CL E 1 27

ge the r a re l a rge r a nd l a rge r (Fig Th e a rra ngeme nt

.

o f the bu n dles with refe re nce to o ne a no the r a nd to the

muscle a s a wh ol e will be desc ribed i n m ore det a il i n a
~

subseque nt p a ra g ra ph He re we ma y sa y th a t the sh a pe
.

o f the muscle is dete rmi n ed by the a rra ngeme nt o f th e

bu ndles ; th a t is they ma y be pl a ced end to end m a ki ng
, ,

a l on g a nd sle nde r muscle ; or side by side givi ng a sh ort , ,

thick muscle Ex a mples O f th ese v a ri ou s sh a pes a re

.

nu me rou s a nd f a mili a r to m ost of us i n ou r ownb odies .

M A K E U P OF S M OOT H M U S CLE
-

AN D H EAR T M U S C LE Ins ome of .

th e l owe r a nim a ls the sm ooth

muscle fibe rs a re b ou nd t ogethe r i n
fa i rly defi nite muscles Inothe rs .
,

th ey f orm ra the r l a rge m a sses for ,

ex a mple the f oo t of a cl a m is a
,

m a ss c omp osed of sm oo th muscle Fm 30 A P i ece of . ,

fibe rs B ut i n th e highe r a nim a l s M us cl e S h owi ng Five

.

nd
the fibe rs of sm ooth muscle a re fofiffsedi f a figbg ffi
usu a lly b ou nd t ogethe r i nth e f orm fi ne bu ndl es T h e l a tter .
,

9 f sh ets a n d SO do not m a ke up
e ,

d V I du a l muscles wh ich ca n be
dissected ou t a s ca n p a rti cul a r skelet a l muscles Th us .

th e muscul a r p a rt O f the li n i ng of th e st om a ch c onsists

of two l a ye r s which ca n be sep a ra ted f rom e a ch o the r

but wh ich ca n not be disti nguish ed a s m a de up of

i ndividu a l muscles .

The fibe rs o f he a rt muscle a re b o u nd t ogethe r to m a ke

up a muscul a r b a g i nwhich it is difficult to disti nguish
i ndividu a l muscles .

T H E STR U CTU R E O F S K ELETA L M U S CLE F IBER —A .

si ngle fibe r of skelet a l muscle p rese nts u nde r the mic ro

sc ope a ve ry i nte resti ng a ppe a ra nce S i nce it sh ows a regu,

l a r seque n ce of c ross m a rki ngs or st ri a ti ons (Fig

-
.

But a pp a re ntly th ese c ross m a rki ngs do not h ave much

-

signific a nce Ca reful ex a mi na ti onof the fibe r S h ows th a t

.

i ts p rot opl a sm is of two ge ne ral s orts The re is a se ries .

1 28 M O VE M E N T I N ANI M AL S
Oi de nse p ro t opl as mic st ra nd s ru nni ng th e le ngth of the
fibe r bu t no t o ccupyi ng i ts e nti re bu l k ; th e sp a ces be
twee n a nd a mong these fibe rs a re filled with ve ry thin
w a te ry p rot opl a sm wh ich is p rob a bly no t much fi rme r
(i e visc ous ) th a nthe b ody fl uid or lymph which b a thes
. .

the fibe r on the outside Encl osing th e e nti re fibe r a nd

.

h oldi ng its p a rts in pl a ce is a delic a te memb ra ne or

sh e a th I t is p rob a ble th a t the i n
.

dividu a l c ont ra ctile st ra nds withi n

th e muscle fibe r a re c omp a ra ble
with th e p ro t opl a smic st ra nds
which we re desc ribed a t the begi n
ni ng O f the ch a pte r a s occu rring i n
m a ny of th e fixed p rot o z oa J ust .

a s th ese st ra n d s h a ve th e p owe r of
c ont ra cti ng forcibly so h a v e th ese
,

i nth e muscle fibe r a n d it is thei r

,

c ombi ned a c ti on th a t m a kes u p -

th e c ont ra cti on of the fibe r a s a

wh ole .

FI G 3 1 A S m a ll
.

P i e ce o f a S in
.

gl e M us cl e S M OO T H M U S C LE F I B ER — Th e
Fibe r At (a ) th e fibe r
.
fibe r O f smooth muscles is much
h a s be en crush ed a n d
simple r i n st ructu re th a n is th a t
twis te d to S h ow th e o u te r
sh ea th se p a ra te d f ro m th e O f th e skelet a l muscle Inf a ct on
.
,
p o to pl a sm be nea th At
a cc ou n o f its simplicity O f con
r
t
.

(b) th e d e nse a nd cl o se ly
pa ck e d l o ng i t u d i na l st ructi on it ma y be c omp a red di
s tra nds a re sugges te d rectl y with th e p ro t opl a smic con
,

to ge t h e r wi th th e ch a r
a cte ris ti c cr ss m a rki n
o - gs t ra cti ng st ra nd i n the P rot oz oa
.
,

a lth o ugh as a n a pp a ra tus for

d oi ng w ork it is p rob a bly a g re a t de a l m ore efli
'

cie nt th a n is th e c ont ra cti ng st ra nd i n the one celled -

o rg a n ism s

H EART M U S CLE F I BER is i nte rmedi a te i n st ructu re be

twee nsm ooth a nd skelet a l muscle Th e i ndivid u a l cells .

c o nt a i np ro t op l a smic st ra nd s simil a r to th ose i nskelet a l

muscle ; but the fibe rs i nste a d O f bei ng e ncl osed e a ch i n
.

i ts ownshe a th me rge t ogeth e r i nt o a c onti nu ous p ro to

,
 ATTA CH M E N T OF M U S CL E S

pl a smic netw ork so th a t it is difficult to s a y whe re one

,

fibe r le a ves Off a nd the nex t begins .

A TTA CH M E N T O F M U S CLES N e a rly a l l muscles th a t

.

b ring a b ou t b odily m o veme nts a s disti nguish ed from i n

te rna l m oveme nts a s of the ,

st oma ch or i ntesti nes m a ke ,

”
thei r c ont ra cti ons e ffective
th rough pulli ng up on h a rd
p a rt s (b ones i nhig he r a ni
m a ls sh ell st ructu res i n
,
-

l owe r ) Th e e ff ect is to
.

b ri ng a b out th e be ndi ng O f
j oi nts . M uscles a re a t
ta ch ed eith e r di rectly or
by me a ns o f te nd ons ; th ese
a re t o ugh a n d i nel a stic ex
te nsi ons o f th e c onnective
tissue which e ncl oses the
muscle fibe rs a nd bi nd s th e
bu ndles t ogethe r G ood ex .

a mples o f t en d ona cti ona re

f ou nd i n t h e h a nd (Fig .

Th e muscles wh ich
m ove the h a nd a re so bulky
th a t if th ey we re l oc a ted i n
um a nForea rm a n
the h a nd itself it w ould be FI G 32 H . . d
Ha nd ; to S h ow h ow mus cl es i n
too clumsy to be of se rvice t h e f orea rm a re a tta ch e d by l ong
.

d s o th e w ris t a n fi g rs
Th e a rra ngeme nt wh ich Is t e n on t d n e .

( ) group of mus cl es th a t b end th e

a ctu a lly p rese n
a
t is th a t the wris t a nd fingers .

muscles a re ou t of the
wa y i n th e f ore a rm whe nce th ey c onnect with thei r
,

p oi nt of a tt a ch me nt by l ong te nd ons which ca n be see n

an d felt re a dily i n the b a ck o f th e h a nd a nd a t th e
w rist .

JOI N T M OTI O N S S kelet a l muscles usu a lly O pe ra te a s

.
,

wa s s a id a b o ve by c a usi ng j o i nts to be n
,
d J oi nts a re of
.

m a ny kinds the m ost f a mili a r being the h i nge j oint such

,
1 30 M O V E M E N T I N ANI M AL S

as we h a ve a t elb ow a nd k nee (Fig . This is a

c omm on type O f j oi nt on ly i n th e h ighe r a nim a ls ,

but a ls o i n a l l o f the
g ro ups o f l o we r a nim a ls
th a t h a ve j oi nted limbs .

C ra yfish a nd l obste rs
(Fig . for i nst a nce
,

h a ve limbs with seve ra l

h i nge j oi nts i n them ,

th ese j oi nts bei ng set i n

’

di ff e re nt pl a nes so th a t
by c ont ra cti ng one or
th e o the r th e limb ca n
be be nt now one wa y
an d now a n o the r
. In
o u r o wnb odies f reed om

to m ove i n m ore th a n
o ne di recti on is ob
ta i n ed eithe r by a b a ll

an -
d s ocket j oi nt a s a t
th e sh o ulde r a nd hip ,

or by the ki n d O f j oi nt
we h a ve a t the w rist
whe re a numbe r o f
sm a ll b ones ca n slide
o ve r o n e an o the r i nv a

ri o u s di recti o n, s a nd so
pe rmit flexibility o f
m o ti o n M uscles a r e
.

a rra n ged a b o u t th e
j oi nts i nopp osi ng p a i rs .

Th e numbe r o f such
p a i rs d epe nd s on th e
v a riety o f m oti ons O f
FI G 3 3 Hum a nS k el e to n sh o wi ng
simpl e hi nge j oi nts a t elbow a nd k nee ; which th e j oi nt is ca
. .

ba ll a nd ock e t j i nts a t sh o uld er a nd p a ble Wh a t is me a nt

s o

hi p ; j o i nts m d e u p O f sm ll b o n es a t
.

a a

wris at n d ankle
by o pp o si ng p a i rs is
 M U S CL E FO R M AND DUT Y 1 31

th a t one muscle o f th e p a i r be nds the joi nt a nd th e O pp os ‘

in g muscle st ra igh t e ns it a g ai n Inthe c a se o f th e elb ow

.
,

th e muscle th a t be n ds th e j oi nt is the bi ceps a nd th e

o pp o si ng muscle th a t st ra igh te ns it is the t riceps The .

hip j oi nt which i s c a p a ble of m oveme nt i n seve ra l di rec

ti ons h a s m ore th a n one such p a i r of opp osi ng m uscles .

FI G . 34 — Lobs ter sh owi ng limbs h wi th s ev era l j oints

ea c .
, ,

M U S CLE F OR M I N R E LATI O N T O M U S CLE DUTY The .

f orce wh ich ca nbe exe rted by a ny muscle is simply th e

c ombi ned pull of its individu a l fibe rs S i nce a ny single .

fiber is o f mic ro sc op ic size a nd o f c onside ra ble fra gility ,

it is only by the re bei ng gre a t numbe rs of them t ogether

th a t muscles o f such p o we r a s ou r b odies p o ssess ca n be
obt a i ned Th e a rra ngeme nt of fibe rs i na muscle depe nds
.

on wheth e r th e muscle is on e i n which st re n gth is th e

chief n eed or one i n wh ich a mplitude of m o ti on is re
qui red S t rong muscles must ha ve m a ny fibe rs side by
.

S ide a n
,
d so a re th ick L ong muscles h a ve m a ny fibe rs
.

pl a ced end to end a nd so a re c a p a ble of g re a t dec re a se i n

le ngth thus p roduci ng l a rge m oti ons O f the p a rts to which
,

they a re a tt a ch ed .

T H E STR E N GT H O F M A N Y OF T H E M U S CLE S is rem a rk

a ble p a rticul a rly wh e n we c on
,
side r th e mech a nic a l dis
a dv a nt a ge u nde r which they w ork Th e st rongest .
1 32 M O V E M E NT I N ANI M AL S

muscle g roup i n the b ody is th a t by wh ich one rises on

-

t ip toe ; it is l o c a ted i n th e c a lf o f th e leg . D irect

tests S h ow th a t th e p ressu re th a t ca n be a pplied onthe
b a ll O f th e foo t by the c ont ra cti onof the c a l f muscles ma y
a m ou n t to a s much a s 4 00 p ou nds i na na dul t of a ve ra ge
st re ngth S o st rong a muscle a s
.

this must h a ve a gre a t m a ny fibe rs

j oi ned in the pull I f the numbe r
.

which a re necess a ry to exe rt this

f orce we re pl a ced side by side in
such fa sh i on th a t th ey al l p ulled
di rectly up on th e te nd on th e
c a lf w ould be a t le a st twice a s
bulky a s i t is Comp a ct ness to
.
,

geth e r with the necess a ry st re ngth ,

is secu red inthis muscl eby h a vi ng

the te nd on by which the mu scle
is a tt a ched to th e uppe r leg ex te nd
d ow n th rough th e muscle ne a rly
to its tip a n d h a vi ng th e fibe rs i n
serted on th e sl a n t (see Fig .

This m a kes p ossible th e p rese nce

FI G 35 D i a gra m
i n th e muscle o f a g re a t m a n y m ore
. .

S h o wing h ow a M us cl e
ma y be co mp os ed o f
m a ny Fibe rs wi th o u t
b ei ng u nduly bulky
fibe rs th a n c ould o the rwise be
a cc omm od a ted in the s a me bulk
.

L O N G M U S CLES — Th e re a r e
.

T h e upper te n d onex te nds .

w ell d o w n th rough th e s ome i nst a nces i n th e b ody whe re

mus cl e T he fibe rs ex te nd
sl a ntwise d ow nw a rd to m oveme nt must be e x e r t e d
.

th e su rf a ce o f th e mus cl e
th rough a c onside ra ble dist a nce
wh ere th ey a re a tta ch e d
t o a su rf a c te n
e d o nwhi ch but n eed no t be espe c i a lly p o we r

m erges be l ow i nto the ful A muscle wh ich m a kes this

.

A chill es te nd no f th e h e l
ki n d of m o veme nt must be l ong
o e .

bec a u se a si ngl e fibe r ca n sh orte n by only a ve ry

sm a ll f ra c ti ono f a ni nch he nce to get a t o t a l sh orte ni ng
,

o f seve ra l i n ches a g re a t m a n y fibe rs must be pl a c ed

en d to e nd .Th e best ex a mple O f this s o rt i s th e
so c a lled t a il o r s muscle which is th e muscle by wh ich
’
-
 CLA S S E S OF M O V E M E NT S 1 33

the leg is d ra w ni nt o the c ro ss legged p ositi on T o pl a ce -

.

the leg i n this p ositi o n r equi es

r a n n
exte sive m ti o o n ,

n
’

an d the ref o r e a l o g muscle The t a il.

o r s muscle is
in a f ct th e l on gest muscle i n the b o dy st a rti ,
n g a s it
d oes i n th e t ru nk a b ove the h ip j o int a nd exte nding
c
a ro ss the f ron O f
t th e thigh a n d o ve r th e k n ee t o a n
i nse rti on on the i nne r side O f the leg j ust bel ow the
knee .

C LA S S E S OF M OVE M E N T S — Alth ugh th e m veme ts

o .
o n
of a n y a n im a l a re b o th nume ro us a n d v a r ied th o se ,

th a t a re V isible from th e o utside f a ll i nto six cl a sses

‘

f
as o o ll ws : (a ) P os tu e r This is m o
. r e sig n ific a n t i n t w o

legged forms like ma na nd bi rds th a n i n th e f ou r legged

.

an m
i a ls which h a ve a ste a die r b a se A ma n .w h o is st a n d
n
i g e r ect eve
,
n th o ugh h e be a s o ldie r a t a tte n ti o n m a kes ,

ba l a nci ng mov ements a lm ost c onti nu ously Th ey

.
a re

usu a lly so slight a s to be obse rved only by c a reful w a tch

i ng .
(b ) Locomo ti on U nder th is h e a d w . e i n clude a l l
b odily m oti ons wh ose e ff ect is to t ra nsp ort th e b ody f rom
on e p o i t to a n
n o the r (c) Gra sp i ng wh ich i cludes
.
,
n a l l
m oveme nts by which m a te ri a ls a re seized or h a ndled
!

These m oveme nts a re of fi rst imp ort a nce i n th e secu ring

an d t a ki ng i no f f ood So o
. cl sely r el a ted t o th e g ra spi n g
m oveme nts th a t th ey c a nno t well be sep a ra ted a re th e ,

forcible outw a rd m oveme nts of th e a rm s wh ich t a ke pl a ce

i n st riki ng ( )
. d C h e w i ng a n d s w a l l o w i n g Th ese
.
a re

a g a in m o ti on s which a re p rim a rily c once rned with th e

t a king of food a lth ough i n m a ny of th e l ower anim a ls
,

the m oti ons O f biti ng e nte r a ls o a s a p a rt of c omb a t (e ) .

S en se p ercep ti on U nde r this h e a d a re i ncluded m ove

.

me nts of the eyes of th e exte rna l e a rs i n such a nim a ls

,

a s h a ve m ov a ble e a rs of th e n ,
o se ins n iffi ng a nd p e rh a ps,

a few o the rs (f) Voi ce p rodu cti on a nd brea thi ng All

.
.

the highe r a nim a l s m a ke m oveme nts O f b re a th i ng a nd

i nth ose a m ong th em th a t h a ve v oice th e m oveme nts O f
b re a th i ng a nd O f v oice p roducti on f a ll u nde r a single
he a d The re a re not m a ny m oti ons wh ich c a nnot be
.
1 34 M O V E M E N T I N AN I M AL S

a ssig ned to one o r a nothe r o f these cl a sses An i nte r

.

es t ing fa ct is th a t th e speci a l ized a ctivities O f civilized

'

men the a ctivi ties i n which they d iffe r m ost st rikingly

,

f rom thei r u ncivil ized fell ows fa ll m ostly eithe r i nt o cl a ss

,

( )
c ,
g ra spi n
,
g or cl a ss (f)
,
v o ice p roducti o.n We re a lize
the t ruth o f this wh en we c onside r th a t a ll use of t ools
an d in st rume nts i ncludi ng pe ns c omes u nde r th e
, ,

h e a d O f g ra spi ng a nd th a t it is by v oice p rod ucti on

,

th a t a ll ide a s which a re not c ommitted to w riting a re

c ommu nic a ted .
 CHAPTER X IV

ANI M AL SEN SE O R GAN S

AD J U S T M E N T S I N L OW ER A N I M A LS I nCh a pte r XI I .

th e ge ne ra l f a ct of th e se nsitive ness or i rrit a bility Of p ro

top l a sm wa s st a ted a n d the re a c ti onO f pl a nts to stimul a
,

ti ondiscussed Inth a t ch a p te r th e p oint wa s m a d e th a t

.

i nrespect to fu nd a me nt a ls pl a nt p ro t opl a sm a n d a nim a l

p rot opl a sm a ct simil a rly th e di ffe rences th a t exist de
,

pe ndi ng m a i nly on di ff e rences i n th e rel a ti on of the or

g a n i sm s to th ei r f oo d supply Thus it c o mes a b o u
. t th a t
i npl a n ts the resp onses to stimul a ti on t a ke th e f orm of
sl ow ch a nges du e u su a lly to growth wh ile i n a nim a l s
, ,

the resp onses a re ra p id a nd a re ca rri ed ou t by contra e

,

ti on s of p ro top la s m Th e very S imple a nim a ls th ose

.
,

th a t do not h a ve h ighly speci a lized ti ssiIes re a c t to such ,

fe a tu res i n th e e nvi ronment a s ligh t tempe ra tu re sub , ,

st a nces i n s oluti on g ra vity a nd one or two o th e rs i n

, , ,

defi nite fa shi ons wh ich dem ons t ra te cle a rly th eir se nsi
ti v en ess. For ex a mple th e p ro t oz oa n P a ra m oecium will
, , ,

t ra vel from h ighly illumi na ted regi ons to o the rs less

b rilli a ntly li gh ted I f a numbe r a re pl a ced i n a sm a ll
.

t rough of w a te r wh ich a t one end h a s a tempe ra tu re of

25 C (7 2 F ) a nd a t th e o th e r a tempe ra tu re eith er
°
.
°
.

highe r or l owe r by seve ra l degrees a ll th e P a ra m oeci a ,

will p rese ntly be f ou nd g a the red a t th e end whe re th e

tempe ra tu re is 25 C The a d j ustme nts thus m a de a re
°
.

on the wh ole f a v ora ble to th e org a n isms m a ki ng them ,

either th ro ugh b ri nging th em i nt o th e p rese nce of a b und

an t f ood or by re nde ri n g th em s a fe from h a rm Th e .

p rope rty of i rri t a bility is thu s see nto se rve a h ighly use
ful pu rp ose eve ni n th e l owliest types
,
.

1 35
1 36 ANI M AL S ENS E O R GAN S

R ECE PT OR S OR S EN S E ORGA N S m a de up i nth e highe r

a re

an im a ls o f g ro ups o f cells i n which speci a liz a ti on h a s

ta ke nthe f o rm o f i n te nsi fyi ng th e p rope rty O f i rrit a bility
a t the expe n se O f o the r fu nd a me nt a l p rope rties o f p ro t o
pl a sm The cells o f th e se nse org a ns c a rry on O f c o u rse
.
, ,

regul a r b a sic me ta b o lism but h a ve n,

o fu ncti ona l met a b
oli sm so fa r a s we k n ow excep t such a s is in v olved i nth a t
a lte ra ti on i n th e st a te o f th e p ro t o pl a sm whi ch results

f rom the stimulus a nd a fte r th e distu rb a nce h a s p a ssed

,
“

o n to o the r p a rts O f th e b ody the a ctivity o f th e se n se

org a n c omes to a n e n d u ntil i t is stimul a ted a g a i n .

C LA S S ES O F S E N S E OR GA N S Th e c omm o nmeth od o f
.

cl a ssifyin g the se nse org a ns is i n a cc ord a nce with th e

ki nds of imp ressi ons to wh ich they a re p a rticul a rly
susceptible In a cc ord a nce with th is s cheme we fi nd
.
,

th a t i na ll th e a nim a ls excep t th e ve ry simplest we h a ve

two m a i n cl a sses : (a ) i n te rna l ; (b ) exte rn al As th e .

names imply the i nte rna l se nse org a ns a re l oc a ted i nside

,

the b ody a nd a re a cted up on by stimuli a risi ng with i n

th e b ody ; the exte rn a l recept o rs o n th e o the r h a n
,
d lie ,

on o r n e a r th e su rfa ce a nd a re a c ted up on by stimuli

wh ich a re exte ri or to th e b ody These exte rna l stimuli
n
.

ma y be of v a ri ou s ki nds Inth e m a i n th ey c o sist eith e r

.
,

o f ch a n ges i n th e e nvi ronme nt immed i a tely i n c ont a c t

wi th the b ody or i nch a nges i nth e e nvi ronme nt a t a dis
t a nce which in s ome wa y a re a ble to imp ress the se nse
org a n s Th o se recep t ors which resp on
. d to immed i a tely
a dj a ce n t ch a n ges a re c a lled c ont a ct recep t ors For ex
m
.

am e the o rg a ns o f t ouch a re a ffected only by a ge nts

,

wh i ch c ome a g a i nst th e b ody Th ose recept ors which a re

.

a ff ected by distu rb a n ces a t a dist a nce a re c a lled dist a nce

se nse org a ns Th e org a ns O f sigh t a nd he a ri ng a re il lus
.

tra ti o n s o f th ese .

I N T ER N A L S EN S E ORGA N S We rec og nize inthe h ighe r

.

an im a ls a t le a s t five ki nds o f i nte rna l se nse org a ns S ome .

o f th ese a re k n ow nto be p rese n t a ls o i nm a ny O f th e l o we r

an im a l s ; with reg a rd to o the rs we a re no t q uite so su re .
 H UNG E R 1 37

The five will be listed a nd e a ch c onside red b riefly They

, .

a re the o rg a n s O f hu nge r thi rst p a i n muscle se nse a nd

, , , ,

equ ilib rium .

H U N GER . Th is is a se nse wh ose bi ol ogic a l imp ort a nce

is m a nifest s i nce with o ut it a nim a ls w o uld not be d rive n
,

to th e se a rch for a n d c onsump ti ono f f oo d a nd so w o uld ,

st a rve to de a th Whe n one re a lizes th a t o f a ll a nim a ls

.

ma nonly h a s le a rned th e c on necti onbetwee nthe t a ki ng

o f f oo d a n d th e a v oid a nce o f de a th by st a rv a ti on he ca n ,

a pp reci a te th e imp ort a n ce of the mech a nism which impels

an im a ls to ea t Th e org a nfor hu nge r i nma na nd i nthe
.

h ighe r a nim a ls is l o c a ted i nthe w a ll of th e st om a ch a nd ,

is stimul a ted by a p a rticul a r ki nd of muscul a r c ont ra cti on

i n th e st om a ch w a ll which gives r ise to wh a t is know n
a s a pa n g of h u nge r The se nse o f h u nge r is c omp l ic a ted
.

by the thi ng we c a ll appetite which is not a t rue se nse , ,

a s is h u n ge r bu t is ra th e r a mem ory o f the e nj oyme nt o f

,

f ood We do not know much a b ou t th e rel a ti onof a p p e

.

tite to hu nge r i n a ny O f th e l owe r a nim a ls but i n ou r ,

selves we rec ognize th a t a ppetite h a s a g oo d de a l to do

with a ro usi ng ge nui ne h u nge r In thi nki ng o f h u nge r .
,

an d O f a ppetite one h a s to remembe r th a t civilized ma n

,

h a s g one a l ong wa y i n his h a bits f rom h is s a v a ge p ro

ge nit ors a nd th e conditi ons for a rousi ng the se nse O f h u n
,

g er a r e c o rr esp on di ngly a lte r ed We k no w th a t.a ll ki nds

of a n im a ls wh ich h a ve o ro ve a b ou t i n se a rch o f f oo d
t

a re impelled u rge n tly to th a t se a rch by s ome me a ns At .

p resent i t is not k now nwhethe r th a t me a ns c onsists In ,

such l owe r f orms a s i nsects for ex a mple O f a n org a n O f

, ,

hu nge r i nthe st om a ch th ough such a norg a n is p re se nt

,

i n m a mm a ls a nd p rob a bly i nbi rds .

T H IRS T . This is the se nse wh ich impels l a nd a nim a ls

to t a ke w a te r from time to time f or th e p u rp ose O f re
pl a cing th a t which is l o st from th ei r b odies by ev a p ora
ti on a nd th rough th e ch a nnels of exc reti on In ma n .
,

an d p rob a bly a ls o i n th e highe r a nim a ls th e org a n o f ,

thi rst is l oc a ted i n th e muc ous li ning O f th e th roa t a nd ,

1 38 ANI M AL S ENSE O R GAN S

the se nse is ro used by insufficie nt m oistu re i nth a t reg i on .

This i nsufli ci ency o f m oistu re ma y be due to th e d rying

o u t o f the exp osed su rf a ce a s wh e n on ,
e sleeps with his

m outh o pe n; or ma y be due to a l owe ri ng o f the pe rce nt

a ge o f w a te r i nthe th re a t i nc on seq ue nce of l o ss o f w a te r
f rom th e wh ole b ody Th i rst due to th e fi rst c a use ca n
.

be a ll a yed by sw a ll owi ng s a liv a bu t th i rst c a used by i n

,

sufficie nt w a te r i nth e b ody ca nbe rel ieved only by t a ki ng

i n w a te r f ro m o utside I t is i nte resti ng to no te th a t
.

h um a n bei ngs ma y th ro ugh a ccide nt or dise a se l ose a ll

thei r o the r se nses bu t ve ry few c a ses h a ve eve r been
,

ob se rved i n which th e se n se of th i rst is l ost .

P AI N —The re a re org a ns for p a i n b oth with i n th e

.
,

b ody a nd onits su rf a ce so th a t p a inis i na m a nne r b oth

,

a ni n te rna l a nd a nexte rna l se nse I t is ordi na rily cl a s sed

.

with th e i nte rna l se nses bec a use i nth e expe rie nce o f h u
ma nbei ngs i t is rec og nized a s a ch a nge i nb odily st a te ,

an d no t a s a n e nvi ronme nt a l ch a nge For i ns ta nce if

.
,

the edge o f a k nife is p ressed a g a i nst th e ski n we thi nk ,

o f th e k n ife ; but a s s oona s it p e net ra tes to th e p oi nt of

stimul a ti ng th e p a i norg a ns it is no l onge r th e k nife th a t
we thi nk of bu t th e b ody th a t is hu rt P a i nis a w a rni ng
.

se nse i t is a roused only u nde r c onditi ons wh ich imply

,

h a rm Th e a dj ustme nts to p a i n a re Obvi ously a dj ust

.

me nts wh ose pu rp ose is th e a v oid a nce of inj u ry .

M U S C LE S E N S E. This is a se nse a b ou t which m ost of

u s k now ve ry little bu t which is o f th e ve ry g re a test i m
,

p o rt a nce i n th e a ctivities o f m a nki nd a nd a nim a ls I ts .

org a n s a re l oc a ted i n the muscles a nd a b out th e j oi nts

an d th ey fu rn ish i nform a ti ona s to th e deg ree o f c ont ra e
ti ono f th e muscles a nd th e p o siti onO f the j o i nts This
,
.

i nf o rm a ti onis imp ort a nt i nth a t it is th e b a sis o f success

ful l o c om o ti on Any a nim a l th a t m oves f rom pl a ce to
.

pl a ce by muscul a r e ffort must c ont ra c t its mu s cles i nco r

rec t seque n ce a n d with th e p ro p er deg ree o f f o rce with ,

refe re n ce to on e an o th e r O th e rwise th e a ctivities which

.

co nstitu te l oc om o ti o n will no t be p rope rly coOrdi na ted .

 E! UILI B R I U M 1 39

The muscl e se nse is th e me a ns by which th e m o ti ons a re

thu s a dj usted Ino u rselves little a tte nti onis p a id to it
.

bec a use th e se nse w orks so p e rfectly th a t we do not fi nd

it necess a ry to a tte nd to i t In c a se of i nj u ry to the .

sense a s for ex a mple i nth e dise a se k now na s l oc om ot or

, ,

a ta xi a (C h a p X V I ) the imp a i rme nt of l oc om oti on i s

.
,

v ery st riki ng .

EQ U I LI B RI U M Th is is a se nse f ou nd in a ll a nim a ls
th a t h a ve a speci a l p ositi on to which th ey will retu rn ,

if distu rbed For ex a mple so l owly a n a nim a l a s a

.
,

st a rfish if tu rned on its b a ck will with c onside ra ble

, ,

l a b or righ t itself I nsects c rust a ce a a nd a ll ve rt eb ra te

.
, ,

an im a ls h a ve this equ ilib ri um se nse a nd i n a ll h ighe r

, ,

an im a ls the re is f ou nd a speci a l eq uilib rium org a n In .

ma na nd i na ll ve rteb ra tes this org a nis l oc a ted i nth e ea r ,

the semi ci rcul a r c a na ls m a ki ng up th e chief p orti onof it

-
.

It is a norg a nwh ich is a ff ected by shifts i n th e p ositi on

o f the bo dy with rel a t i on to g ra vity a n d th e a nim a l i n ,

which the shifts o ccu r m a kes righ t ing m o ti ons u ntil it is

b a ck i nits a ccust omed p ositi on .

— The se a re th e se nses wh ich

!

T H E CO N TA CT S E N S Es .

a re a ro used by e n vi ronme nt a l c onditi ons a t the su rf a c e

o f the b ody Th ey i nclude t o uch tempe ra tu re a nd th e
.
, ,

ch emic a l sense so c a lled a s well a s p a i n The c ont a ct

,
-
,
.

senses h a ve a n inte resti ng rel a ti onsh ip to th e a ctivities

of a n im a ls i nth a t they m a ke up th e origi na l b a sis o f the
rel a ti on s O f a nim a ls to thei r e nvi ronment All a nim a ls .

a re subj ect to th o se in fl ue nces which c ome i nt o d irect

c ont a ct with them I t is only by sec ond a ry devel opme nt
.

th a t me a ns for c omi ng u nde r th e i nflue nce of c onditi ons

a t a dist a n ce c ome i nt o pl a y Fu rthe r refe rence to this .

i nte resti ng f a ct will be m a de l a te r .

T UC
O H — Th e se nse of t ouch h a s a v a st numbe r of
.

org a n s c onsisting of ti ny st ructu res embedded i nth e skin

an d dist ributed ove r ne a rly a ll p a rts o f it Th ese org a s . n
a re much m ore a bu n d a ntly p rese nt in s ome p a rts th a n
i n Othe rs bu t a b ou t the o ly su rf a ce O f a n
,
n
y size f rom
1 40 ANI M AL S E NS E OR GAN S

which th ey a re e nti rely a bse nt is th e f ront O f the eyeb a ll ,

which h a s th e se nse o f p a i nbut no se nse O f to uch Th e

n
.

to t a l umbe r O f t o uch o rg a ns i n ma n p rob a bly m ou nts

i nt o the milli ons T o uch is obvi ously a b a sis for i n
.

form a ti oni nc onnec ti on with ne a rly a ll ki nds o f a nim a l

a ctivity .Th e suit a bility of Obj ects for f ood or thei r ,

p o ssibility a s s ou rces of inj u ry a re dete rmi ned l a rgely

th ro ugh this se nse .

T H E T E M P ER ATU R E S E N S E is th e me a n s by which a n
an im a l pe rceives di ffe re nces o f tempe ra tu re betwee n its
ski na nd Obj ects wi th which it is in c ont a ct F rom the .

st a ndp oi nt o f th e org a ns i nv olved the re a re re a lly two ,

tempe ra tu re se nses si nce we h a ve a set o f org a ns for the

,

p e rcepti on of w a rmth a nd a no the r for the pe rcepti onO f

c old Th e tempe ra tu re se nse is u nd o ubtedly p rese nt i n
.

a ll w a rm bl oo ded a n
-
im a ls viz th e m a mm a ls a nd bi rds ;
,
.
,

wh e the r it exists i nth e c old bl ooded a nim a ls is u nce rt a i n

-
.

The sig nific a nce of th e tempe ra tu re se ns e is obvi ously

to s timul a te th e a n ima l to adj ust i tself to the m o st f a v or
a ble p o ssible tempe ra tu re Th e s timulus O f c old le ad s
.

to a c tivities which eithe r will m a ke th e a n im a l w a rm by

re a s o n o f i n c re a s i ng th e a m o u nt O f fu ncti ona l met a b
oli sm a n , d so the p roducti o n o f h e a t (see Ch a p V I I I ) .
,

o r will b ri n g the a nim a l i nt o a ne nvi ronme nt i nwhich it

is p ro tec ted f rom th e c old W a rm th onth e o th e r h a nd
.
, ,

i nflue nces th e a nim a l to rem a i nc omp a ra tively quiet a nd , ,

i f ex treme to seek a c ool sp o t

, .

T H E C H E M I CA L S E N S ES a re the me a n s wh e reby a nim a ls

a re a flected th ro ugh ch emic a l subst a n ces i n th e e nvi ron
me nt Only subs ta nces diss olved i n w a te r ca n a ct a s
.

s timuli to p ro t o pl a sm a nd so th e ch emic a l se nse re a cts

,

on ly to diss olved m a te ri a l Th e se nse is limited i nsc o p e

.

i nth a t n o t a l l d iss olved subst a n ces a ff ect it a lth ough th e ,

numbe r of subst a nces wh ich do not a ro use the se nse is

m
s a ll i ndeed in c omp a ris on with the e norm ous numbe r
by wh ich th e se nse is a roused .

In a qu a tic a n im a l s the o rg a n for th e chemic a l se nse

 TA S T E 1 41

c onsists of cells l oc a ted c onve nie ntly onthe su rf a ce O f the

b ody whe re the diss olved m a te ri a ls will c ome i n c ont a ct
wit h th em The cells which m a ke up the org a na re a l
.

w a ys exp osed di rectly tO th e o utside with no p rotective ,

ski nbetwee n (Fig T o lesse n th e likelih ood of i n

.

j u ry they a re O fte n l oc a ted i n little p ockets or h oll ows

o f s ome s ort In l a n d a nim a ls th e ch emic a l se nse ca n
.

be a roused b oth by subst a nces i n s oluti on i n w a te r a nd

FI G 36
. . D i a gra m of a n or g a n o f h emi ca l sense (a ) th e pl a ce
c .

wh ere th e sensi tiv e ce lls a re dir ectly exp ose d t o th e e xterior .

by m a te ri a ls which a re p rese nt i n the a i r i n the f orm of

ga s. It is quite t ru e th a t th ese l a tte r subst a nces dis
s olve i nth e m oistu re which ove rlies th e cells of the se nse
org a n bef ore a cti ng up on it so th a t st rictly spe a ki ng
, , ,

the ch emic a l sense i nl a nd a nim a ls ope ra tes a s i na qu a tic

an im a ls th ro ugh subst a nces in s oluti on .

T A S TE Inth e high e r a nim a ls th e org a ns for ch emi

.

ca l se n se a re of two ki nds to fit them for re a cti ng to

'

chemi ca l subst a nces which re a ch the b ody by th e two

diffe re nt ch a nnels suggested a b ove i e i n the f ood or ,
. .
,

d rink which is t a ke ni nt o th e m outh or inthe a ir which ,

1 42 ANI M AL SENSE O R GAN S

is b re a th ed i nth rough th e nose Inth e high e r a ni ma ls .

,

the n we find one o rg a n l oc a ted i n the m outh a nd the

,

o the r i nth e n o se ; th e f orme r I s c a lled the org a nO f t a ste ,

the l a tte r th e org a nO f smell Th e t a ste org a nis a ff ec ted .

on ly by m a te ri a ls which a re a ctu a lly p rese nt i n the

m outh Co nt ra ry to c omm onbeli ef i t h as a ve ry limi ted
.

ra n ge o f pe rcep ti on the re bei ng only f o u r t rue t a stes

, ,

na mely sweet s ou r s a lty a nd bitte r Fl a v or wh ich is

, , ,
.
,

usu a lly c onfused with t a ste is re a lly p a rt of the se nse O f

,

smell (see bel ow ) .

The sig nific a nce of t a ste is obvi ously to a i d i nf ormi ng

a fin a l j udgme n t a s to th e desi ra bility o f f ood th a t h a s
bee n c a p tu red or g ra sped bef ore sw a ll owi ng it I t is
, .

widely bu t by no me a ns i nv a ri a bly t rue th a t m a te ri a ls ,

which a re of a g ree a ble t a ste a re fi t for f ood wh ile ma ,

teri a l s th a t do n ot t a ste g oo d a re u n fi t f or f ood .

T H E D I S TA N CE R E CEPT OR S —U nde r this h e a d we i n .

elude th e th ree se nses o f smell h e a ri ng a nd sigh t Th e , , .

a dv a n t a ge which they c onfe r onth e org a nism p o ssess ing

them is obvi ous inc onnecti onwith a ll or ne a rly a ll O f its
a ctivities For ex a mple to le a rnof th e p rese nce o f f ood
.
,

a t a dist a n ce or O f th e th re a t O f h a rm a t a di st a nce
, _ ,

g re a tly f a cilit a tes th e resp onse o f th e o rg a nism .

S M ELL —.Th e org a n for smell a s st a ted a b ove is , ,

stimul a ted by m a te ri a ls th a t a re p rese nt i na i r i ng a se ous

form I t is l oc a ted i n th e muc ous l ini ng o f th e nose
.
,

whe re th e a i r th a t is b re a th ed in a nd ou t p a sses ove r it ,

an d th e g a se o us c o nsti tue nts o f the a i r a re thus a ble to

stimul a te i t One o r two p oi nts a b ou t th is se nse ma y be
.

me nti oned Th e fi rst is i ts ext ra ordi na ry delic a cy

. .

Wh e nwe thi nk o f th e a m a zi ngly m i nu te a m ou nts O f od or

o u s m a te ri a l whi ch suffice to a ro use th e se n se of smell i n
a g ood do g f o r ex a mple we re a lize the ex t ra ord i n
, ,
a ry sen

si ti v e n ess o f th e o rg a n A sec ond p oi nt is the e a se with

.

whi ch th e se nse is f a tigued Th e c onti nued p rese nce o f .

an y od o ro us subst a n ce i n th e a i r th a t is bei ng b re a thed

quickly f a ils to be pe rceived I t is this which e na bles p er .
 T HE O R GAN OF H E A R ING 1 43

s ons to rem a i ni npl a ces wh e re the a i r is se ri o usly viti a ted ,

with ou t knowing it I t a ls o expl a ins why f oul b re a th is

.

not smelled by the pe rs on h a vi ng i t .

Whe na i r th a t is b re a thed i nc ont a i ns Od orous m a te ri a l

it is rec ognized a s c omi ng f rom outside a nd the sens a
ti onis c a lled smell Whe nthe O d orous m a te ri a l is i nthe
.

m outh a s wh e n f ood is bei ng e a te n a nd a i r c ont a i ni ng

, ,

sOme O f it is b re a thed ou t ove r th e org a n of smell th e

sens a ti onis c a lled fl a v or It is e a sy to sh ow th a t fl a v or

.

is re a lly smell a nd not t a ste for wh e none h a s a c old i n ,

the he a d his f ood a s he s a ys l oses its t a ste wh e re a s wh a t

, , ,

i t re a lly l oses is its fl a v or Th e t rue t a stes sweet s ou r .

, , ,

s a lty a nd bit te r a re pe rceived re a dily no m a tte r h ow

, ,

seve re one s c old ma y be ’

.

FI G . 37 .
— D i a gr a m
hum a n ea r (a ) E xterna l ea r ; (b) ea r
of .

ca n a l ; (c) e a r dr um whi ch i s s e t i nvib rat i onby s o u nd w a ves ; (d) ch a in

of b on es by whi ch vib ra t i on s of ea rdr um a re tra nsmi tte d to sensi tiv e
p a rt of ea r ; (e) ca vi ty fille d wi th liqui d wi thi n whi ch a re p roto ,

pl a smi c recep tors whi ch a re a ff ecte d by vib ra ti ons ; (f ) one of th e

semi circul a r ca na ls p a rt of th e or ga n of e quilibri u m not directly
, ,

con cern e d i nh ea ri ng T h e entire stru cture excep t th e externa l ea r

.
, ,

is imb edded in sol id b one .

TH E ORGA N H EARI N G I s T H E EAR (Fig 3 7 — I t

OF .

c onsists of a n a pp a ra tus whe reby s ounds stimul a te p ro

tOp l a sm S ou n . ds a re du e to vib ra ti ons eith e r o f s olids ,

liquids or g a s es The m a j ority of the s ou nds wh ich

,
.

re a ch o u r ea r a re c a used by vib ra ti ons i nth e a i r Th ese .

ra on
V ib ti s st rike up on a d rum he a d — th e ea r d ru m - -

an d set it i nt o ex a ctly synch ronous V ib ra ti on Th e .

1 44 A NI M AL S EN S E OR GAN S

vib ra ti ons o f th e ea r d rum a re t ra nsmitted to a liquid

-

th a t is wi thi n th e se nsi tive p a rt o f th e ea r se tt ing th e ,

liquid in t u rn i nto vib ra ti o n a nd th e vib ra ti ons O f the ,

liquid a ff ect p ro topl a smic recept o rs which a re di rectly

exp osed to them .

S IGH T This se nse is a nexte nsi on a mplific a ti on a nd

. , ,

m odific a ti on O f th e resp onse O f p rot opl a sm to ligh t de ,

sc ribed i nCh a pte r X I I I nits simplest f orm ina nim a ls

.

it c onsists a s st a ted i n th e begi nni ng o f th e p rese nt

,

ch a pte r o f a ne ffect up o nth e p rot opl a sm of such s ort th a t

,

the a nim a l is impelled to m ove eith e r t ow a rd or a w a y

from th e ligh t Th is implies b o th re a cti onto th e p res
.

e nce of ligh t a nd s ome ki nd o f rec og niti ono f th e di rce

ti onf rom which i t c omes The re a re ve ry few ki nds of .

an im a ls th a t a re dev o id o f this type of se ns a ti on The .

se nse o f sigh t a s c omm only th ough t o f i ncludes a l ong

, , ,

wi th the p e rcepti on of ligh t rec og ni ti on O f th e fo rm o f ,

o bj ects . As s oon a s pe rcepti o n O f f orm is a ch ieved a

g re a t a dv a nce is m a de i nth e u tili ty o f th e se nse bec a use ,

th e re is now th e p o ssibili ty o f g a i ni ng a ccu ra te i nf orm a

ti ona s to th e n a tu re S ize di rec ti o n a n
,
d dist a nce O f th e
, ,

obj ec ts th a t m a ke up th e e n vi ronme nt This imp ort a nt .

k nowledge c a nno t be O b ta i ned wi th a ny th i ng like th e

sa me deg ree o f a ccu ra cy by a ll th e othe r se nses w orki ng
t oge th e r .

T H E P ER CEPT I O N O F F O R M requi res th a t th e O bj ect sh a ll

reflect ligh t up o n a se n si tive p ro to pl a smic su rf a ce a nd ,

th a t th e ligh t so refl ected sh a ll m a ke up on th is su rfa ce

a p a tte rn o r im a ge o f th e obj ect f ro m wh ich the ligh t

c omes Ina st ric t se nse a ny S peci a li zed recep to r wh ich

.
,

re a c ts to ligh t is a neye but i na l l th e h igh e r a n

,
im a l s a nd
i nm a n y o f th e l owe r a s well th e eye is a na pp a ra tus for
formi ng im a ges .

I M A GE F O R M AT I O N is a cc o mplish ed i nth e eyes O f a ni

m a ls ina t le a st two di ffe re nt w a ys ; th e comp o u nd eyes
of i n sec ts do i t i no ne wa y wh ile th e eyes o f ve rteb ra te,

an im ls
a — i ncludi ng ma n do i t i n a no th e r (Fig -“
.
 A C CE S S O R Y EYE S T R U C TU R E S 1 45

The l a tte r is the only wa y th a t will be c onside red he re .

The p rinciple is f a mili a r si nce it is m a de use o f i n or

,

di na ry ph ot og ra phic c a me ra s I t c onsists i n f ocu si ng

.

be a ms o f l ight by me a ns of a le ns In a ll the h ighe r

,
.

an im a ls th e front p a rt of the eye a cts a s a le ns th rough

which light reflected from th e obj ect is f ocused i n su ch
a wa y th a t the im a ge of the obj ect is f ormed up on the

FI G 38
. . D i a gra m v erteb ra te ey e (a ) cornea ; (b) crys ta lline
of .

l ens ; th ese to geth er m a k e up th e f ocussi ng a pp a ra tus ; th e l ens

s erves a ls o i na ccommoda ti on; (0 ) th e retina or sensi tive sur fa ce upon
whi ch im a ges a re f orm ed ; (d) th e b e gi nni ng of th e op ti c nerve ;
(e) the i ris .

se nsitive su rfa ce know n a s th e reti na wh ich is a t the ,

b a ck of th e eye i n li ne with the le ns .

A CCES S OR Y EY E STR U CTUR E S In th e eyes o f th e

.

highe r a nim a ls th e re a re one or two a dditi ona l p a rts which

ca ll f or b rief d es cripti on Fi rst is th e a rra ngeme nt for
.

ch a ngi ng th e f ocus or for a ccommoda ti ona s it is c a lled

,
.

This is necess a ry i n orde r th a t obj ects a t di ff e re nt dis

t a nces from the eye ma y be see n with equ a l cle a rness .

Eve ry one k nows th a t a n a dj ustme nt t a kes pl a ce i n h is

owneyes whenh e tu rn s from l ooki n g a t dist a nt obj ects
1 46 ANI M AL S EN S E OR GAN S

to l ook a t a b ook i n his h a nd The ch a nge O f f ocus is .

b ro ugh t a b o ut by a thicke ni ng of th e c ry st a l li ne le ns a nd ,

this intu rn results fro m th e c ont ra cti ono f ti ny m uscles

, ,

wi th i n th e ey e b a ll S ec ond is the a dj u stme nt o f the

.
,

di a ph ra gm k now na s the i ri s to regul a te th e a m ou nt of

ligh t e nte ri ng the eye Th a t th e p upil is s m
, ,

. a ll i nb righ t

ligh t a nd l a rge indim h a s bee nno ticed by ne a rly eve ry

on e . Th e ch a nge i n S ize o f th e pupil is th e result of
a d j ustme n ts i n th e i ris wh e reby ligh t i s eithe r l a rgely
,

excluded or a ll owed to e nte r f reely .

T H E P ER CE PT I O N O F C O LOR — A fe a tu re of sigh t n ot

yet t ouch ed up onis th e rec og niti ono f c ol or Th e a bility .

to pe rceive c ol or c ont ributes g re a tly to the utility o f sigh t

i nma n a s well a s to th e e nj oyme nt h e de rives f rom it
, .

Th e p rese nce o f b righ t c ol ors i n bi rds fish i nsec ts a nd , , ,

o the r f o rms which h a ve eyes h a s led to th e ge n e ra l beli ef

,

th a t th ey sh a re with ma n th e a bility to pe rceive c ol or .

Wh ethe r these l owe r f orms a ctu a lly do p ossess c ol or vi

si ono r no t is not e a sy to p ro ve a nd a t th e p rese nt time ,

we must a dmit th a t we k now ve ry little a s to h ow wide

sp re a d a m ong a nim a ls c ol or visi onis I no u rselves c ol or .
,

visi onis evide ntly due to speci a l p rope rties p ossessed by

th e se nsitive reti na l cells Th ese p rope rties a re no t p res
.

en t i nequ a l deg ree i nth e eyes O f a ll pe ople Col or bli nd

n
.

ness is a c onditi o i nwh ich s ome c ol ors usu a lly red a nd ,

g ree n a ppe a r a s neut ra l ti nts i nste a d of a s defi nite c ol ors

,
.

C ol or bli nd ness is f a i rly c omm on p a rticul a rly a m on g ,

m a les o f wh om f ou r ou t O f eve ry h u nd red o nth e a ve ra ge

, ,
.

a re c ol o r bli n d .

T H E P ER CE PT I O N O F D I S TA N CE is a ccu ra te on ly with i n
a c omp a ra tively limited ra n ge Th is is bec a u se i t de .

pe nd s up onth e c ombi ned use o f th e two eyes Wh e never .

on e l ooks a t a n e a r O bj ect h is eyes a re tu rned i nto s ome

exte nt i n orde r to b ri ng b o th o f th em to be a r di rec tly .

Th e tu rni ng i n is a cc omplished by th e muscles wh ich

m ove the eye b a lls a nd the j udgme nt o f h ow fa r th e eyes
-
,

h a ve bee n c onve rged is m a de th rough th e muscle se nse

 R E LATI O N OF S IGHT TO T O U CH 1 47

( p a ge 1 3 8) i nth e eye mu scles Whe n eve.r a nObj ect is so

f a r a w a y th a t th e eyes a re no t se nsibly c onve rged i nl ook

i ng st ra igh t a t i t this meth od o f j udgi ng dist a nce f a ils
, ,

an d such estim a ti ons a s a re m a de a re c onseque ntly less

a ccu ra te .Th ey dep e nd in f a ct up ona j udgme nt a s to
, ,

h ow fa r a w a y th e obj ect mu st be i n orde r to a ppe a r a s

sm a ll a s it d oes In o th e r w ords j udgme nt of dist a nce
.
,

is b a sed onk nowledge of size If th e Obj ect is of k now n

.

size a s a ma n th e dist a nce ca nbe estim a ted quite a ccu

n
, ,

ra tel y ; but if the size must be guessed a t th e dist a ce is ,

—
likely to be seri ously mis estim a ted .

T H E R ELATI O N OF S I GH T T o T O U CH A f a ct o f i n .
-

terest i n c on necti on wi th th e c onclusi ons h um a n beings

d ra w from th e sense imp ressi ons th ey receive is th a t sen
sa ti on s o f S igh t a re ge n e ra lly i nte rp reted i n te rms of
t ouch Th a t is to sa y th e ide a one re a lly f orms wh e nh e
.
,

l ooks a t a ny Obj ect is h ow i t w ould feel if h e c ould get his

h a nds on i t We a re t old th a t this is b ec a use t ouch is a
.

p rimitive se nse while sigh t is of much l a te r devel opme nt .

He nce the tende ncy to f orm j udgme nts i n te rms of the

,

sense which h a s fu ncti oned l onge r Wh a teve r the hist ory

.

of th e rel a ti on sh ip ma y be th e f a ct is u nde ni a ble th a t

,

no c omp reh e nsive ide a of a ny obj ect is to be h a d except

a s the mi n d de a ls with it from th e st a ndp oint of h ow it
w ould reve a l itself if it were h a ndl ed .
 C HAPTER XV

N E R VO U S CO R R E LATI O N

OR I GI N O F T H E N V US ER O SY S T E M — I nC h a pte r X I I I
.

the f a c t wa s p oi nted o u t th a t i norde r fo r the m o ti on s of

an im a ls to be c orrel a ted i na cc ord a nce with th e situ a ti ons
existi ng i n the e nvi ronme nt the st imul a ti on O f th e re
cep tors m ust be c on duc ted to th e eff ect ors a s we na me ,

th e muscul a r tissue by me a ns of a n a pp a ra tus wh ich is

,

c a lled th e a dju s tor mecha ni sm E a rly i n the b ook .

( C h a.p I I ) i t wa s st a ted th a t o ne o f th e fu nd a me nta l

p rope rties o f p rotopl a sm is sensi ti veness or i rri ta bi li ty ,

a p rope rty by vi r tue o f which the o rg a n ism resp onds to

stimul a ti on A fe a tu re of this p rope rty is th a t the dis
.

tu rb a n ce set up by th e stimulus sp re a ds th ro ugh th e p ro

top l a sm so th a t p a rts a t a dist a n

,
ce sh a re i nth e resp onse .

Th e ad stor mech a nism evide ntly depe nds on th is ten

de ney O f distu rb a nces to sp re a d th rough p ro t opl a sm to
p oi nts m ore or less dist a nt f rom th e stimul a ted p o i nt .

In i nt roduci ng th e ge ne ra l t opic o f ne rv o us c orrel a ti on ,

whi ch c once rns itself with the a cti on of th e a dj ust or

mech a nism th e v a ri ous st a ges betwee nth e simplest p os
,

sible a rra ngeme nt a nd the highly c omplex st ru ctu re o f

th e highe r a nim a ls will be o u tli ned a s b riefly a s p o ssible .

COR R ELAT I O N I N P R OTO Z OA —Th a t eve n so simple a n

.

o rg a n ism a s a p ro t o z oa nsh ows defi nite a d j ustme nt to its

e nvi ronme nt is sh ow n by th e p a ss a ge o f distu rb a nces
f ro m th e su rf a ce o f th e p ro t o pl a sm th rough out th e m a ss
an d so to th ose speci a l p a rts O f i t which a re c a p a ble O f
m o ti on Thus i nP a ra m oecium wh i ch swims by me a ns
.
, ,

o f cili a ,the vig or O f st roke or O fte nits di recti on ca nbe

, ,

see nto v a ry a cc o rdi ng a s th e org a nism c omes i nt o c ont a ct

wi th one th ing o r a nothe r a s it swims a l ong Th e cili a .

1 48
 P R I M ITIV E M E TA Z O A 1 49

Wh ose m oti ons need not a ctu a lly c ome i nt o

a re a ffected
c ont a ct with th e Obj ect th a t c a u ses the ch a nge in thei r
st roke Usu a lly it is th e ve ry tip of the P a ra m oecium s
.
’

b ody which m a kes the c ont a ct a nd the distu rb a nce is

t ra nsmitted from this p oi nt th rough th e b ody to th e c i li a .

He re we h a ve i n simplest f orm a mech a nism for

a dj ustme n t .

FI G 3 9
. .
— D i a gra m
illus tra te h ow a sensi tive cell
to , R, on th e
sur f a ce ma y lie i n di r ect conta ct wi th a contra ctil e cell
, ,
E, u nder
nea th a dis t urb a nce of R b eing tra nsmi tte d directly to
,
E .

CORR ELAT I ON P R I M I TI VE M ETA Z OA

IN M a ny
Celled Ani ma ls ) — Am ong th e ve ry simples t Of th e
.

m a ny celled a nim a ls a re to be f ou nd f orms i nwhich th e

-

speci a liz a ti on O f fu ncti onh a s p roceeded to such a p o int

th a t the cells wh ich a re se nsitive to exte rna l i nflue nces
a re n ot themselves c a p a ble o f m o ti onbu t lie i ni mmedi

a te c on t a ct with oth e r cells which h a ve th e p rope rty O f

m oti onh ighly devel oped a nd which rep rese nt th e p rimi
tive f orm of s mooth muscle cells (see C h a p X I I I ) . .

The rel a ti on o f the se nsitive cells to th o se wh ich ca n

a chieve moti on is ill ust ra ted i n th e di a g ra m (Fig .
1 50 NE R VOU S CO R R E LATI O N

I t will be see nth a t th e c ont a c t O f one with th e o the r i s

so cl o se th a t a distu rb a nce o f the se nsitive cell to wh ich ,

we ma y now g ive the na me of recep tor si nce it is speci a l

i z ed fo r receivi ng stim ul i ca nbe t ra nsmit ted di rectly to
,

the effec tor (muscle ) cell so a s to excite i t to a ctivity .

The next step i n th e devel opme nt is see n i n Fig 4 0 .

,

whe re the recep t or a t th e su rf a ce is sep a ra ted by a sh ort

dist a nce f rom the muscle cell ; th e ga p is b ridged by

FI G 4 0
. . D i a gra mill us tra ting h ow a p roj ecti o nof s ensi tive cell ,

R, ma y b ridge a sm a ll ga p to contra ctile cell E ena bling a

, ,
dis
t urb a nce o f R to be tra n smi tte d a cross the ga p to E .

a n outg rowth O f th e recept or cel l which e na bles i t to

est a blish c ont a c t wi th th e e ffec tor Th e nex t step which
.
,

is illust ra ted i nFig 4 1 is one i nwh ich th e dist a nce b e

.
,

twee n the recep t or a nd th e e ff ect or is too g re a t to be

b ridged by a simple exte nsi ono f th e recept or a nd so we ,

fin d an o th e r cell m a ki ng th e c onnecti on This cell wh ich

.

c omes betwee n th e recept or a nd the eff ect or cells a nd ,

se rves to c a rry mess a ges a c ross f rom the recep tfii to th e ‘

 CO R R E LATI O N I N HIGH E R FO R M S 1 51

effect or c onstitutes th e begi nning of a defi nite a dj ust or

,

mech a nism .

C ORRELAT I O N I N H I GH ER F OR M S As one g oes up th e

.

an im a l sc a le bey ond th e m ost p rimitive multicellul a r

types i nc re a sed c omplexity of st ructu re is sh own i n v a
ri ou s di recti on s Only two a re to be st ressed he re Th ey
. .

a re (1 ) i n c re a sed numbe r of recept ors a nd e ffect ors a nd ,

( 2 ) a device f or pe rmitti ng flexibili ty o f c onnecti on be

FI G 4 1
. . D i a gr a m t o S h ow h ow wh enth e dis ta nce fro m R to E
is to o grea t t o b e b ridge d by a simpl e o u t gr ow th of R a na ddi ti ona l
cell , A , e sta blish es th e conn ecti on
. S u ch a n i nterm e di a ry cel l ,
spe ci a liz e d f or con du cti on, represents th e b egi nni ng of a d efini te
a dj ustor m ech a nism .

tweenthe recept ors a nd e ff ect ors Inth e l a st p a ra gra ph

.

the org a nism wa s pi ctu red a s th ough i t h a d only one re

cept or a nd one e ff e ct or S O th a t th e only ne rv ous system
n
,

requi red wa s a si gle cell c on necti ng th e recept or to th e

e ff ect or A s a m a tte r o f f a ct one d oes not h a ve to go
.
,

very fa r up the a nim a l sc a le bef ore e nc ou nte ri ng forms

which h a ve m a ny recep t ors a nd likewise a g oodly numbe r
1 52 N E R VO U S CO R R E LATI O N

o f efl ecto rs On e wh o studies c a refully th e a ctivities o f

.

a no rg a nism o f this kind will pe rceive th a t the a d j ust or

mech a nism must be m ore c omplic a ted th a na nything th a t
h a s bee npictu red th us fa r For it ca nbe sh ow nth a t a
.

si ngle eflector ma y be pl a yed up on now by one recept or

an d now by a no th e r Als o th a t a ny give nrecept or ma y
.
,

i nflue nce s ometim es one a nd s ometimes a no the r of th e

e ffect ors E vide ntly th e a dj usto r system i nsuch a na ni
.

ma l c onsists of s ometh i ng m ore th a nne rve cells ex te ndi ng

di rectly f rom individu a l recept ors to i ndividu a l e ffect ors ,

si nce th a t a rra ngeme nt O ff e rs no me a ns by wh ich a ny re

cept or c o uld est a blish c onnecti onwith a ny eff ect or oth e r
th a nthe p a rticul a r one with which its a d j ustor ne rve cell -

c onnects .

FI G 42
. D i a gra m to s u gges t th e orga n
. i z a ti ono f a simpl e n erv o u s

sys te m From the rece p to rs , R a nd R , sens o ry nerv e ce lls , S , l ea d

.
’

to th e ga ngli o n,
G Th e n ce m o to r n
. erv e ceUs , M , l e a d t o t h e e ff e c to rs ,

E an d E I t is se e n th a t wi thi n t h e ga ngli o ne a ch o f the se n sory

’
.

nerv e cells h a s conta ct wi th b o th m o to r nerv e cells .

TH E S TR U CTUR E N ERV OU S S Y S TE M :
OF TH E Th e
meth od by which flexibility o f c onnecti on betwee n re
cep to rs a nd eflectors is est a blish ed i na nim als is by me a ns
O f a n a rra ngeme nt c omp a ra ble to a teleph one i nst a ll a
ti on Eve ry recept or h a s ne rve cells le a di ng f rom it to
.
-

a regi on i n th e b ody which we ma y c a ll a n e rve ce nte r ,

or to use th e t ech n
,
ic a l na me a ga ngli on (see Fig ,
.

The d istu rb a nces a roused i n th e recept ors will be con

d u cted f rom th em to this ce nte r i nste a d o f d i rectly to
a neff e ct or E a ch e ff ect or intu rnh a s ne rve cells le a di n
. g -
 S IMPLE R E FL E X ACTI ON 1 53

to it which c ome di rectly f rom the g a ngli on i nste a d o f

f rom a recep t or Th us it is p ossible fo r ne rv o us distu rb
.

a uces to p a ss f ro m r ecep t or to effect or but o n ly by wa y

,

O f the ce n te r or g a ngli on a nd only by p a ssi ng th rough

,

two n e rve cells I t is cust om a ry to c a ll th e ne rve cell

-
.
-

which extends betwee n the recept or a nd th e g a ngli on a

sen sory n e rve cell a nd th e one which exte nds f rom the
-
,

g a ngli onto th e e ff ect or a mo tor ne rve cell Th e a rra nge

-
.

me nt which is desc ribed is e a sily u nde rst ood bec a use a s ,

st a ted a b ove it is c omp a ra ble to a teleph one i nst a ll a ti on

,
.

J ust a s i n th e l a tte r mess a ges c oming i n ove r one li ne

ca nbe di rected ou t ove r a n o the r a nd th ei r c o u rse ca nbe
direc ted from the ce nte r so th a t th e o utg o ing li ne is now
,

on e an d now a no the r the ne rve i mpulses c omi ng i nt o th e

,

g a ngli on from th e recep t ors a re p a ssed th rough it a nd

ou t i n t o p a thw a ys le a di ng to one or a no the r e ff ect or .

S I M P LE R E F LE X A CT I ON — A f a c t o f th e beh a vi or o f
.

an im a ls which ca n be cle a rly dem onst ra ted by c a reful

study is th a t th e stimul a ti onO f a give n co ta ct recept or n
ne a rly a lw a ys resu l ts i nth e a cti onO f a p a rticul a r eff ect or
'

th e resp on se O f the e ffect or c omi ng with no m ore del a y

a fte r the stimul a ti on o f th e recep t or th a n is n ecess a ry
for the ne rv ous imp ulse to p a ss f rom th e recep t or up to
the ce nte r a nd f rom th e ce nte r ou t to th e e ffect or S o .

fa r a s the m a j ority O f a cti ons of th e simple a nim a l go ,

on e migh t dispe n se with th e g a ngli a a nd supp o se th e re

cept or a nd e ffect or to be di rectly c onnected a s th ey a re ,

i n the ve ry p rimitive f orms desc ribed i n a f orme r p a ra

g ra ph E vide ntly the re a re withi nth e ce nte r or g a ngli on
.

p refe rred or u su a l c onnecti ons betwee nsens ory a nd m ot or

nerve cells so th a t u nde r ordi na ry ci rcumst a nces th e ner
-

v ous distu rb a nce a rrivi ng from a ny p a rticul a r se nsory

cell is p a ssed on a t once to a p a rticul a r m ot or cell a nd
a rou ses a p a rticul a r e ffec tor to a ctivity
,
A c a se of th is
.

s ort i n wh ich a given a cti on f oll ows th e stimulus Of a

,

give nrecept or with regul a rity i s ca lled reflex a cti on a nd

, ,

the p a rticul a r seque nce of ne rve cells betwee n the re

-
1 54 N E R VO U S CO R R E LATI O N

cept o r which is stimul a ted a nd th e e ffect or which ordi

ma rily c omes i nt o a cti on a s a result o f its stimul a ti on
is k now n a s a reflex p a th We ca n im a gi ne a n a nim a l
.

wh ich h a s a nywhe re f rom twe nty to one h u nd red recep

t ors a nd from h a lf a d oze nto f orty e ff ecto rs exhibiti ng
,
- -
,

a g re a t v a riety O f simple reflex a c ti on s a s one recep tor or ,

an o the r is stimul a ted a n d so on e or a n o the r e ff ect or i s

th row n i nt o ope ra ti on .

N ERV O U S OR GA N I Z AT I O N I N J OI N T ED A N I M A LS Inth e .

p a ra g ra ph a b ove a situ a ti on is pictu red as existi ng i n a

f a i rly simple met a z oa nwh ose ne rv ous system c onsists o f
a si n gle g a ngli on with o utlyi ng ne rve cells c onnecti ng it -

with th e recep tors onthe one h a nd a nd with the eff ect ors ,

o n th e o th e r .C omplexity o f b od ily st ructu re c omes

a b o u t i n th e a n im a l ki ngd om i n v a ri o u s w a ys but th e ,

on e which is ch a ra cte ristic O f m o st O f th e h igh e r a n i m a ls

is th e device O f j oi nti ng th a t is O f h a vi ng th e b ody m a de
‘

, ,

up o f a seque n ce O f j o i nts or segme nts I n its simplest .

f orm such a j o i nted b ody h a s e a ch divisi on ex a ctly like

eve ry o th e r a nd such a b ody ca n be rep rese nted a s i n
,

Fig 4 3 wh ich is simply Fig 4 2 repe a ted ove r a nd ove r

.
,
. .

In a n a n im a l so c onst ructed e a ch j o i nt h a s a l l th e
o rg a n s necess a ry for the m a i nte na nce O f life a nd such a ,

mech a nism c ould be th ough t O f a s a c ol ony of rel a tively

i ndepe nde nt o rg a nisms N eedless to sa y p ra ctic a lly a ll
.
,

O f th e j oi n ted a nim a ls go bey ond this simplicity a nd th e re ,

is speci a liz a ti on a m ong th e j oi nts (i n respect to m ost

o rg a n s ) so th a t s ome j oi nts c a rry ono ne b odily fu ncti on
an d s o me a no th e r S O fa r a s th e ne rv ou s org a n
. iz a ti on
is c once rned h oweve r th e speci a liz a ti onis i nm a ny c a ses
,

not ve ry m a rked As Fig 4 3 sh ows th e st ructu re i n

. .
,

such org a nisms c onsist s esse nti a lly of a reduplic a ti on

o f th e a rra n geme nts a l re a dy de sc ribed for a simple
o rg a n ism th a t i s to sa y e a ch segme nt h a s its
, ,

re cep to rs ,
it s eff ect ors its ga ngli on with se ns ory
, ,

ne rves p a ssi ng f ro m th e recept ors to th e g a ngli o n a nd

m ot or ne rves f rom th e g a ngli o nto the eff ect ors Inorde r .
 NE R VOU S O R GANI Z ATI O N
’

1 55

for aj o inted a nim a l to fu ncti onwith a ny degree of suc

cess a l l of th e segments must w ork i nh a rm ony He nce .

the m a chi ne ry O f ne rv ous c orrel a ti oni nth ese must be so

org a nized a s to extend from j oi nt to j oi nt As Fig 4 3 . .

sh ows the a rra ngeme nt is to c onnect e a ch g a ngli onwith

,

its neighb ors on eith e r side by ne rve cells which exte nd -

di rectly f rom g a ngli on to g a n gli on S i nce these a re

.

neithe r se ns ory cell s nor m ot or cells bu t c onnecti ons we ,

give them the na me O f connecti ng cells Th e p rese nce .

FI G . 43 . il lus tra ti ng th e ner v o us sys tem i na j o inted

D i a gra m -

an im a l In a ddi ti on to th e s tru ctu res sh ow n i n figure 4 2 th e con

.

necti ng nerve cells (0 ) a nd (c ) a re sh own by whi ch nerv o us dis

'
,

t ur b a nces ca n p a ss f ro m j oi n t to j o int
. For t h e s a k e of S impli ci ty
p a rt O f th e po ss ibl e nerv e connecti ons i n th e s econd ga ngli on a re
o mi tte d .

of these c onnecti ng cells Ope ns up a dditi ona l p o ssibilities

o f c omplexity i n th e m o veme n ts O f ne rve imp ulses for ,

now a distu rb a nce origi na ti ng i na ny recep t or a nd p a ssi ng

n
f rom it to the g a ngli onis ot c onfi ned i nits ou tw a rd p a th
to th e m o t or ne rves le a di n g di rectly f rom th a t g a ngli on
to th e e ff ec tor bu t ma y p a ss o ve r the c on
,
necti ng ne rve
cells to a na dj oi ni ng g a ngli ona nd so to a ne ffect or i nthe
1 56 N E R VO U S C O R R E LATI O N

a d j oini ng j oi nt or ma y p as s f rom th e g a ngli ona l ong th e

,

wh ole le ngth o f the b ody a ro using th e m ost dist a nt ef

,

fectors to a cti on .

CO M P LE X R E F LE X A CT I O N I t will be cle a r th a t i na
.

j ointed a nim a l with its multiplicity o f recept ors a nd e f

fectors a nd its nume ro us g a n gli a the re is much m ore sc ope
for c omplexity o f n e rv ous a cti onth a n i nth e s imple un
’

j oi nted forms An a dditi ona l fe a tu re o f the situ a ti on

.

a ls o c on t ributes to the c omplexity na mely th a t the re is

, ,

no th ing to p reve nt m ore th a n one recept or f rom bei ng

stimul a ted simult a ne ously In fa c t th e u su a l th ing is
.
,

for a numbe r of recept ors to be stimul a ted a t once This .

me a ns th a t seve ra l st re a ms o f ne rve impulses a re p ou ri ng

i nt o the s a me g a ngli on if th e a nim a l be of th e simple
type with on ly one j oi nt or i nt o v a ri ous g a ngli a i f the
,

an i m a l is c omp osed o f seve ra l j oi nts In eithe r c a se.

p rope r a dj ustme nt requi res th a t th e st re a ms o f i nc omi ng

impulses be so c orrel a ted th a t the o u tg oi ng impulses
to which they give rise sh a ll th row th e eff ect ors i nt o
a cti on i n a wa y th a t i s orde rly a n d a d j usted to th e
requ i rem e nts Of th e situ a ti on P a rt o f th e fu ncti on o f
.

the g a ngli a is th us to c orrel a te the st re a ms o f i nc omi ng

ne rve impulses i nt o p rope rly org a nized ou tg oi ng st re a m s
to th e e ff ect ors . Betwee n the simple a nd d i rect p a ss a ge
of a n e rv ous impulse f rom a give nrecep t or to a p a rticul a r
e ff ect or a nd this org a niz a ti ono f st re a ms o f impulses to
,

b ri ng a b ou t th e c omplic a ted yet orde rly a ctivities o f

m a ny e ff ect ors is a l ong step a nd we must c onfess a t once
,

th a t we do no t k now h ow th e p rocess o f c orrel a ti on is

a ctu a lly c a rried o u t I t is no t a t a l l difficul t to p rove
.

th a t th e re is such a p ro cess a nd th a t i t fu ncti ons ve ry

efficie ntly bu t much bey ond th a t th e i nvestig a ti ons o f
,

bi ol ogists h a ve not yet e na bled them to go .

I M PORTA N CE O F T H E F RO N T E N D O F T H E B ODY I f the .

a ctivities o f a j o i n ted a nim a l be c onside red a s it m oves

f ro m pl a ce to pl a ce it will be re a lized th a t its f ront end
,

is ordi na rily th e p a rt th a t comes fi rst i nt o a new e nvi ron

 ADVANTAG E OF DI S TAN CE R E CE P T O R S 1 57

me nt a nd is the ref ore th e fi rst to be a ff ected by wh a tever

is new i nth a t e nvi ronme nt Obvi ously the recept ors a t
.
,

the front end will be m ost f reque ntly stimul a ted a nd thei r
stimul a ti on will be of g re a test sig nific a nce to th e or
n i sm I t is i n a cc ord a nce with th is p ri nciple th a t we
g a .

fin d th e m ost imp ort a nt recep t ors a t the front en d ; a nd

p a rticul a rly is it t rue of ne a rly a ll a nim a ls th a t such
di s ta n ce recep tors a s they p ossess a re i nth a t p a rt of the

b ody Animp or t a nt c onseque nce of this g roupi ng of th e

.

dist a nce recep t ors i n one regi on is the c orresp onding i n

c re a se i n size a nd c omplexity of th e front g a ngli a Th e .

dist a nce recept ors a re c omplic a ted org a ns c omp osed o f

nume ro us p a rts a nd r equi ri ng m a ny se ns ory ne rve cells -

to c on nect them with th e ce nt ra l g a ngli a ; the result is

th a t the g a ngli a a t th e front end of th e b ody a re usu a lly
the l a rgest of a ll th a t the a nim a l p o ssesses a nd ona cc ou nt ,

of thei r c on necti on with the dist a nce recep t ors th ey a re

a ls o th e m ost imp ort a n t He re we h a ve th e begi nni ng
.

o f th e b ra i n n a mely a g ro up O f g a n
, ,
gli a with a nespeci a lly
l a rge numbe r of se ns ory ne rves le a di ng i n a nd o f c ou rse ,

a n equiv a le n t supply of o utg oi ng ne rves eith e r m ot or , ,

le a d ing di rectly to effect ors or c onnecti ng le a di ng to

, ,

o the r g a n gli a a nd so i ndi rectly to m ore dist a nt e ffect ors

, , ,
.

M ore ove r it is of p red omi na ti ng imp ort a nce bec a use it

,

is the g a ngli ong roup with wh ich a ll th e di s ta nce recep

-

tors c ommu n ic a te a nd i nt o which th e ref ore a re p ou red

, ,

the ne rv ous impulses b ri ngi ng i nform a ti onO f f a ct s i nth e

e nvi ronment a t a dist a nce f rom th e b ody su rf a ce These .

f a ct ors of numbe rs O f ne rve c onne cti ons a nd O f r el a ti on

with the dist a nce recept ors i n c ombi na ti on m a ke the
, ,

b ra ina t once th e d omi na nt p a rt o f th e ne rv ous system .

T H E A DV A N TA GE OF T H E D I S TA N CE R E CE PTOR S Wh en .

the c ont a ct recept ors a re stimul a ted the resp onse is a s ,

p rompt a s th e w orki ng O f th e mech a nism will p e rmit .

Obvi ously this p romp t ness is a nesse nti a l p a rt O f a dj ust

,

ment to c ont a ct stimuli si nce if th e s ou rce O f th e s ti mu

,

l a ti onis i nimmedi a te c ont a ct with th e b ody immedi a te

1 58 N E R VOU S CO R R E LATI O N

a ctivity is likely to be th e o nly ki nd th a t wil l se rve the

an im a l Wh e n th e dist a nce recep t ors a re stimul a ted
.
,

onthe o the r h a n d th e a dj ustme nt need no t be immedi a te

, .

I f th e stimulus is i ndic a tive o f d a nge r th e re a cti onma y

be del a yed s omewh a t a nd yet th e org a nism ma y su rvive .

This int roducti onO f a p ossibility o f del a y i nresp onse to

'

st imul a ti on pe rmits a g re a t exte nsi on of ne rv ous a cti on

inth a t it ope ns th e wa y for th e m a ki n g o f a ch oice a s to
wh a t th e resp onse is to be In simple reflex a cti ons .
,

a s h a s bee nrep ea tedl y sa i d n o o ch oice e n

,
te rs th e resp onse ,

foll ows immedi a tely a nd a u t om a tic a lly up onthe stimul a

ti on Th e p ossibili ty of ch oice implies onth e o the r h a nd
.
, ,

th a t the resp onse need no t be im medi a te a nd a ut om a tic

bu t ma y be del a yed s omewh a t a nd ma y be subj ect to
regul a ti o n . We ca nth i nk O f only one b a sis for th is regu
l a ti on which w o uld be o f a ny v a lue wh a tev er to the or
g a n i s m n
,
a mely wh a t h
,
a s h a ppe n ed to it bef ore ; so th a t
ch oice to be w orth a nyth i ng must be f ou nded on p a st
, ,

expe rie nce A ve ry little reflecti onwill sh ow cle a rly h ow

.

v a st is th e exte nsi onO f a na nim a l s ra nge o f a dj ustme nt

’

i f i t ca nc a ll p revi ous expe rie nce int o pl a y a s a p a rt O f the

mech a nism for gu idi ng i ts p rese nt beh a vi or .

M A CH I N ERY FOR U S I N G P A S T EX P ER I EN CE In a n .

e a rlie r p a ra g ra ph th e f a ct wa s b ro ugh t ou t th a t dist a nce

recept ors c ommu n ic a te d i rectly with th e f ront g a ngli on
o f th e n e rv ous system th e b ra in S ome O f the ne rve
,
.

cells O f th is front g a ngli onh a ve a peculi a r p rope rty no t

f ou nd i nth e o the r ne rve cells i n the b ody na mely th a t
-
, ,

wh e nne rve impulses c ome i nt o th em th ey a re no t p a ssed

ou t immed i a tely a s is th e c a se with th e o the r n e rve cells
'
-
,

bu t a re st opped a nd h eld Th e result o f th is p rope rty is

.

to i n te rrupt reflex a c ti on a nd p reve nt its c ompleti on .

These ne rve cells h a ve th e a dditi ona l peculi a rity th a t the

-

ne rv o us d istu rb a nces th a t a re thus held with i n them

m a ke s ome ki nd O f pe rm a ne nt imp ressi onup onthem so ,

th a t u nde r suit a ble c o nditi ons impulses p revi ou sly re

cei ved ca nbe p a ssed ona g a i n Th us a reflex a cti onth a t
.
 CO N TA C T R E CE P T O R S I N ME M OR Y 1 59

is st a rted a t one time ma y be c ompleted l a te r Th is .

a bility o f b ra i n cells to st ore n e rve impu lses is c a lled

memory a nd is the b a sis f or th e use o f p a st expe rie nce
,
.

I t S h ould be remembe red th a t a ll expe rie nce is b a sed

up onth e stimul a ti onof recept ors a nd the m a chi ne ry o f,

mem ory is a m a ch i ne ry for st ori ng th e imp ressi ons which

these stimuli imp a rt for use a t s ome futu re time An
,
.

imp ressi ononce st ored i nth e f orm O f mem ory ma y se rve

a s the st a rti n g p oi nt for a ctivity no t once but repe a tedly ,

thus m a ki ng a S i ngle mem ory a ble to b ri ng a b out m a ny

a cti on s a nd so c ont ribu ti ng g re a tly to th e u tility O f
mem ory .

CON TA CT R E CE PT ORS P ART I CI PATE I N M E M OR Y — In

p ra ctic a lly a ll a nim a ls wh ich h a ve a b ra i n th e c ont a ct
recept or s a n d p rob a bly th e int e rna l recept ors a s well h a ve
i ndi rect c ommu nic a ti on by me a ns of c onnecti ng ne rve
cells with th e b ra i n (Fig . so th a t wh e n a n y o f the
se nse org a ns a re stimul a ted it is p rob a ble th a t ne rv ous
impulses will p a ss i nt o th e b ra i n In th e c a se o f m ost
.

c ont a ct recept ors this connecti onwith th e b ra i nw ould be

c a lled a sec ond a ry c onnecti on th e p rim a ry c onnecti on
,

bei ng with the m ot or ne rve cells Wh e nrecept ors h a vi ng

-
.

this ki nd of c onnect iona re stimul a ted the ordi na ry reflex

resp on se f oll ows a t once a ut om a tic a lly a nd a t th e s a me
, ,

time the re is a flow O f ne rv o us impulses i nt o th e b ra in .

Th e se which p a ss to th e b ra i nregiste r the re a s mem ories ,

j ust a s do the impulses c omi ng m ore di rectly f rom the

dist a nce recept ors Th us a l l th e ch a nnels o f expe rie nce
.

bec ome a v a il a ble for th e fu tu re guid a nce of th e a nim a l .

A S S O CI AT.I O N — M em ories a re m a de se rvice a ble by a

fu rthe r p rope rty O f th e b ra i ncells to whi ch we give the
-

na me of a ss oci a ti on This is th e g ro upi ng t o gethe r o f

.

mem ories th a t a re rel a ted so a s to form o th e r a nd m ore

m
c omplex m e ories Th e best wa y to S h ow the si gni fi
.

c a nce of a ss oci a ti on is by illust ra ti on It is cust om a ry .

to use for th e p u rp o se o f illust ra ti on th e c a se of a b a by

th a t st a rts life with ou t a ny b a ckg rou nd O f expe rie nce
1 60 N E R VO U S CO R R E LATI O N

a nd with o ut mem ories S upp ose th a t before

th e ref ore .

th e eyes o f the b a by a n ordi na ry bell is held Th e .

FI G 44
. T h e H uma n N erv o u s S ys tem , sh o wi ng the m a i nn
. e rv e

tru nks commu ni ca ti ng wi t h the sp ina l co rd , whi ch e x te nds d o wn

wa rd from the b ra i n i nto the b o dy Wi thi n the spi na l co rd th e
.

se ns ory nerv e s fro m co nta ct o r i nte rna l re ce p to rs ma y m a k e di re c t

co nnec ti o ns wi th m o to r ne rve s to e ff e c to rs a nd a ls o by m ea ns o f
, ,

co nnect i ng nerv es ma y m a k e i ndi rec t co nnec ti o nwi th th e b ra i n

,
.

Ma rtin Hu ma nBody
. .
 A C TIVIT Y I N H IGH E R A NI M AL S 1 61

b a by sees th e bell or i no the r w ords th e im a ge of th e bell

, ,

is formed up on the se nsitive su rf a ce o f the b a by s eyes ’

,

This im a ge stimul a tes se ns ory ne rves by which i n tu rn

ne rv ous impulses a re c onveyed to the b ra i n wh e re the
se ns ory imp ress i oni s registe red a nd st ored a s a mem ory
This is pu re mem ory of th e a pp e a ra nce o f a ce rt a i nOb
j c et a nd ca n h a ve no me a ni ng i n th e ch ild s mi nd At
,
’
.

an o the r time th e bell is ru n g th e child h e a rs i t a nd a n

,

o the r recept or is stimul a ted a n o th e r set o f n

, e rv ous i m
pulses is a ro used a nd i na n e nti rely diff e re nt p a rt of the
,

b ra i n a mem ory i s registe red this bei ng a mem ory of a

,

ce rt a i n s o u nd S till l a te r th e child gets its h a nds up on

.

th e bell a nd receives a se ries of se ns a ti ons of t ouch which

a re registe red a s mem ori es I na still di ffe ren t p a rt of the
b ra i n Thus th e ch ild h a s th ree sep a ra te a nd disti nct
.

mem ories a ll rel a ti ng to th e s a me obj ect By s ome p ro

n
.
,

cess withi nth e b ra i nwhich we do no t a t a ll u de rst a nd ,

these th ree rel a ted mem ories a re a ss oci a ted In o the r .

w ords th e child pe rceives th a t th ey a l l bel ong t ogethe r

,
.

Thus it f orms wh a t we c a ll a nide a which is a mem ory ,

o f th e bell a s a wh ole We must remembe r th a t th e b e

.

gi nni ng O f this p roces s i na b a by is ona ve ry sm a ll sc a le

an d is p rob a bly no t c a rried onve ry efli ci entl y so th a t it ,

ma y t a ke th e child s ome time to f orm s a tisf a ct ory a ss oci a

ti ons We a ll k now f rom ou r ownobse rv a ti on h oweve r
.
, ,

th a t a t a ve ry e a rly a ge th e norm a l inf a nt is f o u nd to be

equipped with a c onside ra ble ra nge of ve ry useful
a ss oci a ted mem ories .

TH E CO N TR OL OF A CT I V I TY I N T H E H I GH ER A N I M A LS .

(a ) R eflex F rom wh a t h a s bee ns a id i nth e p a ra g ra ph s

.

a b ove it sh o uld be cle a r th a t th e a ctivities O f a n y a nim a l

wh ose b ra i nh a s th e p rope rties o f mem ory a nd a ss oci a ti on
ma y be g ove rned i n a t le a st two disti nct w a ys In the .

fi rst pl a ce much of th e a ctivity ma y be pu rely reflex

,
.

By this is me a nt th a t se ns ory stimul a ti on ma y le a d a t

on ce a n d a ut om a tic a lly to ce rt a i n defi nite m oveme nts .

More Of ou r own m oti ons th a n we re a lize a re o f this

1 62 N E R VOU S CO R R E LATI O N

ch a ra c te r For ex a mple th e a ct o f w a lking a lth ough

.
, ,

ve ry c omplex is i na tech nic a l se nse a p u re reflex Th is

,
.

me a ns th a t the muscul a r c ontra cti ons which m a ke up th e

a ct a re i n iti a ted a nd regul a ted by immedi a te se ns ory
s timul a ti on Th e se ns ory stimul a ti onwh ich ope ra tes i n
.

this p a r ticul a r reflex h a s a numbe r of s ou rces th e p res ,

su re o f th e s oles O f th e feet up onthe g rou nd the muscle ,

se nse a roused f ro m th e c ont ra cti ons O f the muscles a nd

th e be nd i ng of the j oints th e se nse of equilib rium b rough t
,

int o pl a y a s th e h e a d swi ngs with th e m oti ons O f the

b ody the se nse of sigh t if one h a ppe ns to be w a lki ng
,

with th e eyes op e n a ll c ont ribute stim uli which a re cor

,

r el a ted with in a p a r t O f the ce n t ra l ne rv ous system set

a side for th a t p u rp os e a n d c a used to d isch a rge a na cc u
ra tely timed a n d p rope rly g ra ded se ries o f stimu l i to
c a use the muscles to pe rform i nseque nce a nd to the de
g ree th a t is necess a ry for successfu l w a lki ng .

( )
b V o li ti o na
. l C o nt ra sted with a n a ct o f this s ort ,

wh ich is pu rely reflex a re m a ny a ctivities which a re

,

sp oke nOf a s vo lu nta ry na mely a s c a rried ou t by th e will

, , .

Wh e na v oliti ona l a ct is a na lyzed i t is f ou nd to be guided

by a c omb i na ti on o f th e immedi ate e nvi ronme nt with
such a ss oci a ted mem ories a s a pply to th e c a se i nh a nd .

I t i s c onceiv a ble th a t a v oliti ona l a ct migh t be th e resul t

wh olly o f mem ory no immedi a te stimulu s h a vi ng a ny
,

thi ng to do with st a rting it Off Y et study of the a ctivi

.
,

ties O f a nima l s a s well a s ou r owna ctivities sh ows th a t

i na v a st m a j ority O f c a ses th e a cts th a t we c a ll v oliti onal
depe nd up ona c ombi na ti onof immedi a te se ns ory sti mu
l a ti onwi th a ss oci a ted mem ory S i nce a ll a ss oci a ti ons a re
.

buil t origina lly up on se nse imp ressi ons i t is p ossible to

t ra ce th e origi nof a l l a c tivities b a ck to recept ors Thus .
,

s tric tly spe a ki ng th e b ra in d oes no t origi na te a ctivity

,

b u t me rely c a uses to go onto c ompleti onch a i ns o f eve n ts

wh ich h a ve th ei r o rigi ni nse nse o rg a ns .

I N S T I N CT I nm a n y o f th e l o we r a nim a ls but m o st
st riki ngly exemplified i ni n
.
,

se cts a re to be see nnume rous

,
 B E HAVI OR 1 63

c omplic a ted a ctivities which a re desc ribed a s i nsti ncts to

distinguish them f rom a cts of v oliti on The c omplexity .
,

orde rlin ess a nd utility o f these i nsti nctive a cti ons a re

,

ofte n a st o u n di ng a nd to one Wh o h a s no t l ooked i nt o

,

them it migh t seem th a t they c a nno t i ndic a te a nythi ng

less th a nthe exe rc i se of a ve ry high deg ree o f i ntelligence .

As a m a tte r Of f a c t c a reful study dem onst ra tes c onclu

,

si vely th a t th ese so c a lled i nsti nctive a cti ons no m a tte r

-
,

h ow c omplic a ted a n d orde rly a re p u rely reflex i n origi n

,

an d c ont ro l The a nim a ls which c a rry them ou t do so

.

onthe b a sis o f immedi a te se n s ory stimul a ti on th e a cti on ,

f oll ows immedi a tely up ona nd is dete rmi ned by the sum
t ot a l of the se ns ory stimuli which the a nim a l expe rie nces .

It must be a dmitted th a t m a ny f a cts a b ou t i nsti nct a re

not a t a ll u nde rst ood a t p rese nt For ex a mple we ma y.
,

t a ke th e i nsti nct Of mig ra ti on i nbi rds I t is cle a r th a t.

the stimulus f or mi g ra ti on is a n immedi a te s timulus ,

b a sed i n p a rt p rob a bly on ch emic a l ch a nges th a t go on

i nthe b ody of th e bi rd itself a nd i np a r t on fe a tu res O f
the e nvi ronment B u t th e a ctu a l executi onof th e migra
.

ti on p a rticul a rly such fe a ts a s the fligh t of bi rds f rom

,

Al a sk a to a ti ny o utlyi ng isl a nd o f th e H a w a ii a n g roup ,

an d the re h itt ing up onthe isl a nd with u ne rri ng a ccu ra cy

a t the en d O f hu nd red s o f miles O f fligh t with n o si n
gle
l a ndm a rk i nV iew is e ntirely u nexpl a ina ble a t th e p rese nt
,

time .

B EH A VI OR D ETER M I N ED BY N ERV OU S ORGA N I Z ATI ON .

The questi on of whethe r i n a ny give n species o f a nim a l

a ss oci a ted mem ory (v oliti on ) S h a ll d o mi na te o ve r pu r e
reflexes (i n stinct ) or whethe r th e r everse sh a ll be the
,

c a se is dete rmi ned by th e m a ke u p O f th e ne rv ous system

,
-

from bi rth C e rt a i nki nds of a nim a ls i nhe rit from thei r

.

an cest ors a ne rv ous org a niz a ti on i n wh ich a ss o ci a ted

mem ory pl a ys a nimp ort a nt p a rt o the rs i nhe ri t a ne rv ous
system i nwh ich c omplic a ted reflexes dete rmine th ei r b e
h a vi or Hum a nbei ngs bel ong i n th e fi rst O f these c a te
.

gori es i n ,sects i n th e sec ond M a ny f orms p a rticul a rly

.
,
1 64 N ER V O U S CO R R E LATI O N

th e l owe r ve rteb ra tes rep rese nt combi na ti ons o f th e two

,

ki nds O f o rg a niz a ti o n It is w o rth no ti ng th a t wh e nwe

.

a sce n d to the ve ry top o f the sc a le to ma n himself we , ,

‘

fin d instincts i n the se nse i nwhich we u su a l ly spe a k o f

,

them a l m ost wh olly w a nti ng v oliti ona l a c ts t a ki ng

, ,

their pl a ce .

H A B IT FOR M ATI O N I t h a s bee nsh ow nh ow ina ni

.
,

m a ls th a t p ossess the p rope rty Of a ss oci a tive mem ory a ny ,

give na ct ma y be b a s ed up onth e c ombi ned effect of im

’

medi a te se ns ory stimul a ti on a nd such a ss oci a ted mem

ori es a s a re relev a n t Assumi ng th a t th e a ctivity thus
.

set i nt o m oti onp roves useful th e recu rre nce o f th e s a me ,

c ombi na ti on of stimuli w ill na tu ra lly b ri ng a b ou t the

s a me resp onse I f th e situ a ti onis one which recu rs fre
.
.

que ntly the re will be su ccessive repetiti ons of the s a me

,

ki nd of a ctivity The resul t is to est a blish wi th i n the

.

ne rv ous system excepti ona lly re a dy c onnecti ons betwee n

th e se ns ory a ss oci a ti on a nd m ot or regi ons i nv olved
, , ,

so th a t with th e recu rre n ce o f th e se ns ory stimul a ti onthe

pe rform a nce of th e p a rticul a r a ct f oll ows m ore a nd m ore
ce rt a i nly Thi s is wh a t we c a ll ha bi t forma ti on I t will
. .

be see nth a t it a m o u nts to a supp ressi onof th e v oliti ona l

p a rt of th e c ont rol thu s m a k ing th e p rocess a pp roa ch
,

th e st a tu s of a p u re reflex N O bette r illust ra ti on o f.

h a bi t form a ti on c ould be give n th a n th e d rivi ng of a n

a ut om obile Th e begi nne r receives fi rst a se ries O f sen
.

s ory imp ressi ons i nth e f orm O f i nst ru cti ons a nd obse rv a
ti ons o f o the r d rive rs ; from th ese h e f orms a se ries o f
a ss o ci a ted mem ories
; a nd wh e n h e begi ns to d rive his ,

equipme nt for pe rformi ng th e a ct successfully c onsists O f

these mem ories t ogethe r with th e se ns ory stimuli which
,

c ome to him a s th e ca r m oves a l ong Th e c orrel a ti on .

o f these i n t o c orrect muscul a r a c ts is a t fi rst ex t remely

d iffi cult ; a ll th e mem ories th a t h a ve bee na cqui red must
be c a lled i nt o the mi nd eve ry time a new m oveme nt on
the p a rt o f th e d rive r bec omes n e cess a ry With th e p a s .

s a ge of time this necessi ty fo r rec a lli ng d efi nitely o ne s ’

 CO N S C I O U S N E S S OR AW A R E N E S S 1 65

i nst ructi ons diminish es a nd ultim a tely th e p rop er d ri ving

,

m oti ons f oll ow the receip t of the se ns ory stimuli a s

g

p romptly a nd a s a ut om a tic a lly a s i n a ny pu re reflex .

Once th is situ a ti on is re a lized we sa y the h a bit is fully

-

f ormed The re a re of c ou rse a ll p ossible st a ges i nh a bi t

.
, ,

f orm a ti on depe ndi ng onthe c omplexity o f th e mem ories

,

inv olved a nd on th e f requency of th e repetiti on of the

stimuli .

U
C O N S C I O S N E S S OR A WA R N
E E S S — I t sh ould be cle a r
th a t th e p rocess of a ss oci a ti ng mem ories m a kes up wh a t
we c a ll th ough t or i ntellectu a l a ctivity D oes it f oll ow
,
.

th a t eve ry a n im a l which is a ble to f orm a ss oci a ti ons h a s

the p owe r O f th ough t ? In a nswe ri ng this questi on a
!

fe a tu re O f i ntellectu a l a ctivity or th ough t must be t a ke n

i nt o a cc o u nt which h um a n bei ngs a nd p rob a bly m ost of
the high e r a nim a ls p ossess bu t c once rning which we a re
ex tremely d oubtful whe n it c omes to the l owe r a nim a ls
this is the p rope rty f a mili a rly k now na s consci ou sness ,

an d to which psych ol ogists give th e na me a wa reness .

The me nt a l p rocesses of h um a n bei ngs a re a cc omp a nied

ve ry l a rgely by a k nowledge of Wh a t i s b ei ng th ough t
'

Thus we a re i nthe h a bi t of a ssumi ng th a t the p rocess of

a ss oci a ti on a lw a ys i n v olves c onsci ous ness or a w a re ness .

This h oweve r is by no me a ns th e c a se ; th e re is g ood

, ,

p roof th a t th e p rocess O f a ss oci a ti ng mem ories ca n a nd

d oes go on with o u t c onsc i ousness bei ng a t a l l i nv olved .

Thus th e f a ct th a t l owe r a nim a ls ca na ss oci a te mem ories

d oes not p rove th a t they a re a w a re of th ei r a ss oci a tive
p ro cesses .

T H E FEE LI N GS OR E M OT I O N S The re is a typ e O f ner

.

v ous a ctivity to which we give th e na me of feeli ng or ,

em o ti on wh ich is not the s a me a s th e ne rv ous a ctivity

,

of the i n tellectu a l p rocesses a lth ough th e two ki nds a re

cl osely inte rw ove n Th e em oti ons a re of th e utm ost i m
.

p ort a nce to h um a nbei ngs bec a use b o th th e j oys a nd th e

,

s orrows of life a re l a rgely w ra pped up i nthem We k now .

by Obse rva ti onth a t m a ny of th e a nim a ls l owe r th a nma n

1 66 N ER V O U S CO R R E L ATI O N

expe rie nce em oti ons o f one ki nd or a no the r One na tu r .

a lly in qui res th e refo re wh a t is the bi ol ogic al s ignific a nce

, ,

o f em o ti on Ina ttemp ti ng to a nswe r this ques ti o nit is

.

necess a ry to no te th a t th e em o ti ons f a l l na tu ra lly i nt o

two opp osin g g roups (a ) the a g ree a ble (b ) the dis a g ree
, ,

a ble .I f these two cl a sses a re a na lyzed i t is f o u n d th a t

th ose of the fi rst cl a ss a re a ss oci a ted eithe r with c ond i
ti ons f a v o ra ble to b odily well be ing or with ra ci a l p er -
,

p e tu a ti on; th e sec on d cl a ss of feeli ngs a re on the othe r ,

h a nd see n to be a ss oci a ted with c onditi ons wh i ch a re

'

inimi c a l to b od ily well bei ng This is cle a rly see ni f we

-
.

list th e m ost imp ort a nt o f th e em oti ons o f th is cl a ss .

They a re : f righ t a nge r em o ti ons c onnected with p a in

, , ,

em o ti ons c onnected with h u nge r em o ti ons o f w orry or ,

an xiety This d oes no t exh a ust the list of th e dis a gree

.

a ble em o ti on s but c ove rs th ose which a re of p rim a ry

bi ol ogic a l imp ort a nce In th e nex t p a ra gra ph will be
.

sh ow nh ow th ese em oti ons a ctu a lly rel a te themselves to

sel f p rese rv a ti on
-
.

B ODI LY C H A N GES I N E M OTI O N — For a l ong time i t .

h a s bee n k n ow n th a t eve ry ge n ui ne em o ti on is a ec om
p a n i ed by s ome s o rt o f b odily ch a nge F a mili a r illu s .

tra ti on s a re th e sh edd i n g o f te a rs i ntime o f s orrow th e ,

blush o f emb a rra s sme nt th e smile of ple a s u re I f th e ,

.

em oti ons h a ve a ny bi ol ogic a l sig nific a nce it w o uld be ex

p e ct e d to be i n c o n necti o n W i th these b o dily ch a nges
wh ich a cc omp a ny them T o get a t th e me a ni ng o f th e.

em o ti ons th e ch ief b odily ch a nges a cc omp a nyi ng the five

,

d is a g ree a ble em oti ons listed a b ove a re set d ow na nd th ei r

sig nific a nce b riefly c onside red T a ki ng them i n orde r .

th ey a re a s foll ows

“ The h a i r st a nd s onend .

N The pupils a re d il a ted .

N Th e f a ce is p a l e .

A The m outh is dry o n a cc ou nt of suspe nsi onof th e

a cti ono f the s a li v a ry gl a nds .
 B ODIL Y C HANG E S IN E M O TI O N 1 67

5 . Th e he a rt be a t is g re a tly quicke ned .

6 . Th e digestive functi onis c ompletely suspe nded b oth ,

the sec reti onof the digestive j uices a nd the muscul a r

m oveme nts O f the a lime nt a ry t ra ct bei ng supp ressed .

All th ese b odily ch a nges a re rec og nized a t once a s ch a r

’

a cteri sti c o f such em o ti ons a s f righ t or a nge r Th e .

an a lysis o f them reve a ls th a t with o u t excep ti onth ey a re

a ss oci a ted with the p rese rv a ti on of th e b ody i n time of

immedi a te eme rge ncy .

Th e e recti on of th e h a i r a lth ough of no imp ort a nce

,

i nma nwith h is sc a nty equipme nt o f h a i r will be e a sily ,

rec og n ized a s a defi nite a i d to self p rese rv a ti on i n such

-

an im a ls a s d ogs or c a ts i n which thei r u su a l a ppe a ra nce

,

of mild f rie n dli ness is t ra nsformed i nt o m a ligna nt feroc

ity by th e S imple p rocess of c a usi ng th e h a i r to st a nd
onen d .

The dil a ti onof th e pupils p rob a bly c ont ributes to the

a cute n ess of visi onou t o f the c orne rs of the eyes a nd th us ,

is of a ssist a nce to figh ting a nim a ls Th e f o u r o the r .

b odily ch a nges i n th e list a ll c ont ribu t e to one end ,

na mely th e f a v ori ng O f met a b olism i n th e b ra i n a nd i n

,

th e skelet a l muscles a t th e expe nse of othe r p a rts of the

b ody Whenth e f a ce tu rns p a le i t me a ns th a t th e bl ood
.

to which i t owes its c ol or h a s bee ndive rted from th e ski n

to the b ra ina n d muscles The suspe nsi onof th e a cti on
.

of the s a liv a ry gl a n ds a nd of the o th e r digestive org a ns

e na bles the bl ood wh ich w o uld ordi na rily be requi red f or
thei r nou rishme nt to be sh ifted to th e b ra i na nd muscles .

The quicke ni ng O f the he a rt be a t me a ns g re a tly imp roved

-

ci rcul a ti on of th e bl oo d wh ich a g a i n be nefits th e b ra i n

,

an d muscles Whe n one rec a lls th a t i n time O f eme r

.

ge noy a n a le rt b ra in a nd a ctive muscles a re vit a lly i m

p ort a nt to self p rese rv a ti on whe re a s digestive a ctivity or
-
,

the ruddi ness o f th e ski nma y be dispe nsed with for th e

m ome nt with o u t h a rm it is cle a r h ow th ese b od ily
,

ch a nges a id i n self p rese rv a ti on

-
.
1 68 N E R VOUS C O R R E LATI O N

T H E R EP R OD U CT I V E I N S T I N CT
. I nne a rly a ll ki nds of
an ima ls rep roducti on a nd the refore ra ci a l pe rpetu a ti on
, ,

depe nds onthe fusi ono f two speci a l sex cells a negg p ro ,

du ced i nth e o v a ry o f the fem a le a n d a spe rm p rod uced

,

i nthe testis o f th e m a le (see Ch a p XX V I ) . I norde r to .

i nsu re ra ci a l pe rpetu a ti onth e re must be a d efi nite gu a r

an ty th a t this fusi on will occu r Th e mech a nism for
.

a cc omplishi n g i t i nv olves c orrel a ted muscul a r m oveme nts ,

which a re set i nm oti ona nd gu i ded j ust a s a re a l l such

‘

m oveme nts th rough th e ope ra ti ono f th e ne rv ous sys tem

,
.

Th e c orrel a ted m oveme nts m a ke u p a c omplex reflex a o

ti on which resembles othe r reflex a cti ons i nbei ng b a s ed
,

on se n s ory stimul a ti on This stimul a ti on is i n p a rt ex

.

te rna l a nd i n p a r t i nte rna l th e l a tte r p a rt risi ng f rom

,

the org a ns of rep roducti on th emselves Ch emic a l regu .

l a ti on (see nex t ch a pte r ) is p rob a bly a f a ct or a ls o The .

c ombi ned e ffec t of these i nflue nces is to a ro use a nexceed

in gly p owe rful em oti onfully a s st rong a s a ny c onnected
eithe r wi th secu ri ng f ood or wi th esc a p i ng h a rm By th is .

em o ti on th e stimuli a re vig orou sly rei nforced a nd th e

org a n isms a re impelled to rep roduce Whe n one rec a lls .

t ha t it is i nc onceiv a ble th a t a ny of th e l owe r a nim a ls ca n

or d oes a pp reci a te the c on necti onbetwee nm a ting a nd the
subsequ e nt a ppe a ra nce o f y ou ng th e V it a l bi ol o gic a l i m
,

p o rta nce o f th e rep roductive i nsti nct i s cle a rly see n .

M ore ove r sh ould th e e ff ect of civiliz a ti onbe to we a ke n

,

th e effects O f th is i nsti nct se ri ously i n th e h um a n ra ce ,

the re w o uld be no e ffec tive m otiv a ti on to p reve nt th e

dis a ppe a ra nce o f m a nkind f rom th e e a rth .
 C HAPT ER X VI

C H E M I CAL CO R R E LATI O N

T H E N E CE S S I T Y FOR GRAD UA L AD J U S T M E N T S . Inth e

ch a pte r j ust finished the me a ns wh e reby a nim a ls a d a p t
themselves p romp tly a nd efficie ntly to th e v a ried requi re
me nts of a n eve r ch a nging e nvi ronme nt h a ve bee n ou t
-

li ned The ge ne ra l t opic of th e a dj ustme nt o f org a nisms

.

h a s h oweve r no t bee n c ompletely c onside red u ntil a c

, ,

c ou nt h a s bee n ta keno f the m a ny gra du a l a dj ustme nts

which org a nisms h a ve to m a ke N ume ro us ex a mples of .

these gra du a l a dj ustments wil l be c onside red inth e p a ra

gra phs bel ow He re by wa y O f illust ra ti on need be
.
, ,

me nti oned only two : i npl a nts th e ri pe ni ng of th e f ruit

,

is a na dj ustme nt which se rves th e bi ol ogic a l requi reme nts

o f th e p l a nt an
,
d it is a na dj ustme nt wh ic h g o es ongra du
»

a lly .I t is not only gra du a l bu t it is a ge nui ne a dj ust

me nt th a t is to sa y the ch e mic a l ch a nges wh ich c onsti
, ,

tute ripe ni ng occu r a t a defi nite time i n rel a ti on to th e

devel opment o f th e seed a s well a s to th e ge ne ra l g rowth
,

a ctivities o f th e pl a nt A gr a du a l a dj ustme nt i na nim a ls

.

is the a d j ustme nt o f symmet ri ca l growth I f one st ops .

to thi n k a b o ut it h e ma y well i nqui re h ow i t h a ppens

th a t the two e a rs wh ich a re sep a ra ted fro m e a ch o th er
,

by the width o f the h e a d m a na ge to grow a t a u nif orm

,

ra te a n
'

d to st op th ei r gr owth a t th e s a me time so th a t ,

i nthe m a j ority o f pe ople th e two e a rs a re o f th e s a me size .

He re is j ust one of t h e nume rous a dj ustments th a t occu r

i na nim a l g rowth a nd with o u t which missh a e na nd dis
p
torted b odies w ould i nevit a bly result I t is cle a r th a t a.

me ch a nism like th e ne rv o us system is not well a d a p ted

for m a ki ng a dj ustme n ts of this ch a ra cte r To m a i nt a i n .

1 69
1 70 C H E M I C AL CO R R E LATI O N

a nunce a sing flow o f ne rv o us impulses to eve ry g rowi ng

p a rt th ro ugh o u t th e pe ri od o f grow th a nd for no o th e r
,

pu rp ose th a nthe regul a ti onof th a t g rowth w o uld seem ,

to in v olve a nexcessive a m ou nt of ne rv ous a ctivity if a ny

simple r me a ns of a cc omplish i ng the s a me result c ould
b e fo u nd .

C H E M I CA L ADJ U S TOR S A s a m a tte r o f f a ct exp eri

.
-
,

me nt h a s sh ow n th a t g ra du a l a d j ustme nts i n org a nisms

a re m a d e th rough th e a ge n cy o f ch emic a l subst a nces .

The g re a t a dv a nt a ge o f this meth od is th a t a chemic a l

regul a t o r ca n be c on st a ntly p rese nt a s a c onstitue nt of
th e p rot opl a sm a nd so ca nexe rcise its i nflue nce c onti nu
,

ou sl y a n d u nif ormly Th a t p rot opl a sm is susceptible o f

.

chemic a l st im ul a ti on h a s bee n p oi nted o u t repe a tedly .

B A S I C AN D GR OW T H M ETA B O LI S M C H I E F LY A FF ECTED .

I t is pe rh a ps w orth while to no te th a t wh ile chemic a l

regul a ti onma y a ff ect a l l th e ki n ds o f met a b olism of wh ich
o rg a n isms a re c a p a ble b a sic m et a b olism a nd th e met a b
,

oli sm o f g ro wth a ppe a r to be m o st defi n itely subj ect to

this ki nd of i nflue nce Fu ncti ona l met a b olism i n con
.
,

t ra st i s typ ic a lly u n
,
de r th e c ont rol o f th e ne rv ous system .

Whe no ne c onside rs th a t the u tility of fu ncti ona l met a b

ol i sm is the immedi a te a d a pt a ti o no f th e org a n ism to its
e nvi ronme nt i t bec omes cle a r th a t ne rv ous regul a ti on
, ,

which b ri ngs a b o u t p romp t re a cti ons to e nvi ronme nt a l

ch a nge s is o n th e wh ole bette r suited to th e c ont rol o f
,

a ctive functi on th a n is th e sl owe r a cting ch emic a l

regul a t or .

C H E M I CA L CORRELAT I O N I N P LA N T S — R efe re n . ce was

m a d e to this ph e nome non a t the cl ose o f C h a pte r X I I .

He re only a few f a mili a r illust ra ti ons will be ci ted A .

ch rys a n themum pl a nt if left to i tsel f will p rod uce m a ny

buds e a ch o f which will d evel op i nt o a m ode ra te sized
,
-

ch rys a nthemum bl oss om By d est royi ng a l l th ese buds

.

bu t o ne a si ngle l a rge bl oss o m ca n be obt a i ned Ev i .

dentl y the re i s he re i nth e pl a nt s ome type o f c o rrel a ti on

which so fa r a s we know must be chemic a l I f the g row
, ,
.
 C H E M I C AL C O R R E LATI O N 1 71

ing tip o f a b ra nch 1 8 i nj u red s ome bud a l ong the stem

, ,

wh ich ma y h a ve bee n l ong d orm a nt a nd which w o uld

ordi n a rily c on ti nue d orm a nt will st a rt i nt o a ctivity a nd
,

c a rry onth e g rowth of the stem a s a wh ole I f a cu tti ng .

of ge ra n ium is th rust i nt o suit a ble s oil roots will devel op ,

oni t a t sp o ts which ordi n a rily w o uld neve r p roduce such

st ructu res All these a re ex a mples of ch emic a l regul a
.

ti on N othi ng is k now na s to the na tu re or s ou rce o f th e

.

chemic a l subst a nces wh ich a ctu a lly se rve a s stimuli for

b ringi ng a b ou t th e ch a nges .

CH E M I CA L CORR EL AT I O N I N A N I M A L S In a nim a ls .

a l l the m a chi n e ry of a dj ustme nt is m ore h ighly org a niz ed

th ani npl a nts a nd SO we fi nd i nthem a defi nite mech a n
,

ism for chemic a l c orrel a ti on This mech a nism c onsists

.

o f the p ro ducti on of sp eci a l ch emic a l subst a n ces which

a re p ou red ou t i n t o the tissu e fluids a nd wh ich exe rt
defi nite e ff ects on met a b olism Th ese subst a nces h a ve
“

m
.

c ome i nt o p ro i ne nce i nrecent ye a rs a nd bid f a i r to a s

sume eve n g re a te r imp ort a nce i n th e p ublic eye a s the
ye a rs go on I t is w or th while th e ref ore to bec ome fa m
.
, ,

ili a r with th e tech nic a l na me wh ich h a s bee n a pplied to

them Th ey a re k now na s h ormones f rom a G reek w ord
.
,

me a ning lite ra lly to excite a nd refe rri ng to thei r p rop

erty of fu n cti oni ng a s ch emic a l stimuli .

H OR O N E M A N UFA CTU R I N G C LAN D s

M - — All of the higher .

an im a ls h a ve i nthei r b odies cell m a sses wh ose fu ncti ona l

met a b olism c onsists i nth e m a nu fa ctu re of p a rticul a r h or
m ones Th ese cell m a sses a re S imil a r to the gl a nds a l
.

re a dy desc ribed by which digestive j uices a re ma n

, u fa c

tu red ; the on ly imp ort a nt di ffere nce betwee n them a nd

the m ore fa mili a r gl a nds is th a t th e m a te ri a ls whic h they
p roduce a re p o u red ou t di rectly i nt o the bl ood st re a m to
be ci rcul a ted a b ou t th e b ody wh e re a s the sec reti ons o f
‘

th e gl a n d s o f th e o the r type such a s th e s a liv a ry gl a nds

, ,

a re p a ssed by me a n s o f S peci a l tubes or ducts i nt o th e di

gesti v e t ra ct or i nthe c a se o f th e swe a t gl a n
,
ds onto th e ,

su rf a ce of the S ki n Bec a use th e gl a nds th a t a re now b e

.
1 72 C H E M I CAL CO R R E L ATI O N

ing desc ribed h a ve no ducts they a re o fte nrefe rred to a s

the du c tless gla n ds .

H OR M O N ES W H I CH A FFE CT B A S I C M ETAB O LI S M A .

f a ct o f g re a t imp ort a nce is th a t innorm a l a nim a ls so l ong

a s th e tempe ra tu re c on ti nues c ons ta nt the b a sic meta b o
l i sm g oes ona t a ve ry u n iform ra te Th e tempe ra tu re he re .

refe rred to is the cell tempe ra tu re a n d n o t th e ex te rnal

tempe ra tu re S o in w a rm bl ooded a nim a ls wh o se cells

.
-
,

live i na u niform tempe ra tu re b a sic met a b olism p roceeds

,

wi th o u t much a lte ra ti oni nra te da y i na nd da y o u t Ev i,

.

d ence h a s bee nobt a i ned of rece nt ye a rs to p rove th a t th e

a ctu a l ra te of b a sic met a b olism is dete rmi n ed by th e i nflu
e nce o f a g roup o f h orm ones a nd th e re a s onth a t i t h old s
,

a t a f a i rly u n iform level is bec a use these h orm ones a re

m a nu f a ctu red a nd p ou red ou t by thei r gl a nd s a t a u ni
f orm ra te S h ould th e gl a nds bec ome dis orde red so th a t
‘

thei r o utpu t is eithe r i nc re a sed or d imi nished the ra te ,

o f the b a sic met a b olism is a lte red to c orresp on d .

FI G 4 5
. . T he Thy roid Gl a nd T sh ow ni nrel a ti o nto
, ,
th e l a ry nx ,

L, a nd t he tra c h ea (w i ndpipe ) W .

T H E T H Y RO ID H OR M O N E T o illust ra te the eff ec t o f

.
-

h o rm o nes o nb a s ic met a b olism a si ngle ex a mple th e best ,

k now no f a ll will be ci ted Th e thy ro id gl a nd lies a t th e

, .

f ront a nd sides o f th e wi ndpipe (Fig 4 5 ) whe re it ordi .

na ril y fits i nt o th e c ont ou r of the neck a nd so is not no

 H OR M O N E S 1 73

ticed ; but s ometimes it bec omes e nl a rged a c onditi on

k now na s goi tre i nwhich c a se the ch a nge i nsize o f the
neck is re a dily pe rceived Th e thy roid gl a nd ma nu fa c
.

tu res a h orm one which h a s bee n ide ntified chemic a lly

an d h a s bee n given the na me o f th yroxi n The ra te o f,

the b a sic met a b olism o f th e b ody cells is dete rmi ned by

the a m ou nt of this subst a nce i nth e bl ood st re a m Whe n .

th e thyroid dep a rts from its norm a l p roducti on of

thy roxi n so th a t the bl ood c ont a i ns too little or too much
,

of the h orm on e the met a b olism is a ffected one wa y or

,

the oth e r a nd the i ndividu a l i nwh om this h a ppe ns is no

l onge r norm a l A desc rip ti onof th e b odily ch a nges thus
.

c a used is give ni n th e next ch a pte r whe re the effects of

,

distu rb a nces i n c orrel a ti on a re t a k en up .

H OR M O N E S A FFE CTI N G GR O WT H M ETAB OLI S M —R e .

fe re nce wa s m a de a t th e begi nni ng of th e ch a pte r to the

fa ct th a t g rowth is a ki nd o f p rocess which le nds itself
p a rticul a rly to ch emic a l regul a ti on O ur k nowledge o f
.

the v a ri ous h orm ones wh ich c ont rol g rowth is ve ry i n

c omplete but two illust ra ti ons will be cited to sh ow
,

s omethi ng of th e meth od of a cti on of the g rowth

regul a ti n g a ge ncies .

T H E H OR M O N E S O F T H E P I TU I TARY B ODY I n th e
.

sp a ce betweenth e roo f o f th e m o uth a nd the b a se of the

b ra i na nd beh i nd th e na s a l p a ss a ges is a h orm one m a nu -

fa ctu ri ng gl a nd k now na s th e p i tu i ta ry b ody This is a

.

c omplex st ructu re a nd p rob a bly sec retes seve ra l h or

.

m ones We a re inte rested he re only i nthe one which is

.

k now n to i nflue nce g rowth The m a nne r i n which this

.

p a rticul a r i nflue nce is exe rted is th rough a n e ff ect up on

the size of th e skelet on I t is a f a ct th a t on the wh ole
.

th e size of a n a nim a l is dete rmi ned by th e S ize of his

skelet on th e s oft p a rts a d a pt i ng themselves of necessity
,

to the g rowth of th e b on es G i a nts a nd dw a rfs h a ve

.

a lwa ys bee nm ore or less c omm on but the c a u se o f o ve r

,

or u n de r growth h a s only rece ntly c ome to light I t is

-
.

now believed th a t a c onside ra ble pe rce nt a ge o f the gi a nts

1 7 4: C H E M I CAL CO R R E LATI O N

th a t a re exh ibited i nsh ows o we thei r excepti o a l size to n

o ve r a c tivity o f t his gl a n
-
d d u ri ng th e no rm a l g ro wth
pe ri od o f ch i ldh o od The co ntra ry c ond i ti on o f dw a rf
.

i sh ness is th o ught to be th e resul t o f u n de r a ctivi ty o f -

th e gl a n d .

T H E I N T ER S T I T I A L H OR M O N E I nte rmi ngled with the

.

st rictly rep rod uctive p a rts o f the o rg a ns o f rep ro ducti on

a re cells which a re gl a n dul a r i n fu ncti on a nd to wh ich
h a ve bee ngive nthe na me of i nte rsti ti a l gl a nd cells Th e .

h o rm one which they sec rete is o f p a rticul a r imp ort a nce

i n gr owth p a rticul a rly i n th a t st a ge which i s c once rned
,

wi th th e ch a nge f rom y o uth ful ness to m a tu rity The re .

a re ce rt a i n b odily ch a n ges ch a ra cte ristic o f this pe ri od

con sisting i nth e devel opme nt o f the so c a lled sec ond a ry -

sex ua l ch a ra cte rs ” Am ong them a re i ncluded such

.

ch a nges a s th e a lte ra ti ono f th e v o c a l c ords wh ich c a uses

th e v o ice to b re a k a n d the d evel opme nt of h a i r on th e
,

f a ce The h orm one o f th e i nte rstiti a l gl a nd is of speci a l

.

i nte rest bec a use obse rv a ti on of i ts e ffects fi rst suggested

th e ide a o f ch emic a l c orrel a ti on Alth ough c a st ra ti o no f
.

d om es tic a nim a ls a nd c a p oniz a t i on of f owls h a d bee n

p ra cticed for ce ntu ries a nd th ei r e ff ect s up o nth e devel o p
me nt o f th e sec ond a ry sexu a l ch a ra ct e rs h a d bec ome a
m a tte r of eve ryd a y k nowledge it rem a i ned for the b i ol o
,

gist a b o ut th e middle o f th e l a st ce ntu ry to expl a i nth e

, ,

c o nnecti on F rom th e studies thu s i niti a ted c a me othe rs

.

whi ch h a ve give n us a ll th a t is now k now n a b out th e

h o rm one mech a nism .

H OR M O N ES A FF ECT I N G F U N CT IO N A L M ETA B O LI S M .

Al th o ugh a s a l re a dy p o i nted o u t fu ncti o na l met a b olism

, ,

i na n im a ls is fo r th e m ost pa rt u nde r ne rv o us ra the r th a n

.

ch emic a l c o n t rol the re a re s o me i nst a nces i n which the

,

fu ncti ona l m et a b olism is a ffe cted by h o rm o nes so this ,

type o f regul a ti o nc a n no t be a l to ge th e r neglected .

T H E P A N CR EATI C H OR M O N E The p a ncre a s is p rim a

.

rily a no rg a nfo r th e m a n u f a ctu re o f a dige s tive se creti on

wh ich is p o u red ou t i nt o the intesti ne th ro ugh a d uc t ;
 H O R MO N E S 1 75

but he re a nd the re a m ong th e cells of the p a nc re a s p rope r

a re cells o f a di ffe re n t type th a t a re now k now nto m a nu
f a ctu re a h orm one which they p ou r o u t int o the bl ood .

This h orm one h a s to do with th e utiliz a ti ono f fuel by th e

muscles a nd thus pl a ys a n imp ort a nt rOl e i n the fu nc
,

ti on a l met a b olism by which m o ti on is p roduced I t is .

fo u nd th a t wh e nthis h orm one is deficie nt the a bility o f

the muscles to m a ke use of sug a r is se ri ously imp a i red .

U nde r ordi na ry ci rcumst a nces p rob a bly from eigh ty to

,

on e hu n d red p er ce nt of the fuel wh ich th e muscles bu rn

i nm a ki ng thei r m o veme nts is sug a r I t is evide nt the n
.
, ,

th a t a n imp a i rme nt of th ei r a bility to utilize sug a r is a

se ri ou s m a tte r In th e nex t ch a p te r a na c co u nt is give n
.

o f the dise a se k n o w na s di a be tes wh ich result s f ro m defi

,

ci en cy i nthe p a n c re a tic h orm one .

H OR M O N ES A FFE CTI N G B OT H B A S I C AN D F U N CTI ON A L

M ETAB O LI S M —Alth o ugh for p u rp oses of desc rip ti on a
.

fa i rly defi nite disti ncti onh a s bee nd ra w nbetweenthe dif

ferent ki nds of met a b o lism it is cle a r th a t they a re m ore
,

or less cl osely i n te r rel a ted so th a t wh a teve r a ffects one

-
,

ca n sc a rcely a v o id a ffecti n g th e o the r i n s ome m a nne r

It w
.

o uld be expected th a t h orm ones which i nfluence a ny

type of met a b olism w o uld b ri ng a b o u t m odific a ti ons i n
o the r types a s well The re is h oweve r a t le a st one speci
.
, ,

fi c h orm one which is k now nto exe r t a p a rticul a r ki nd of

influence on b a sic met a b olism a nd a no the r i nfluence on
defi nite ki nd s o f fu ncti ona l met a b olism For this re a s on .

i t dese rves speci a l me nti on .

T H E H OR M O N E O F T H E A DR E N A L S In cl o se c ont a ct
.

with e a ch kid ney is a sm a ll gl a ndul a r st ructu re k nown

a s the a dren a l body This b ody p ro duces a nd p o u rs ou t
.

i nt o the bl ood a h orm one to which v a ri ou s na mes h a ve

bee ngive n th e m ost f a mili a r of wh ich is a drena li n This
,
.

subst a nce fi rst c a me i nt o p romi ne nce whe nit wa s di scov

e red th a t a little of it d ropped on a w o u nd w ould d ra w
!

t ogethe r th e edges o f th e i nj u red bl ood vessels a nd so -

st op th e bleedi ng I f a sm a ll qu a ntity o f the subst a nce is

.
1 76 C H E M I C AL CO R R E LATI O N

i nj ected i nt o a veina nd dist ributed th ro ugh o u t th e b ody ,

it d ra ws t o geth e r th e w a lls o f the ti ny bl ood vessels a nd -

so c a uses a m a rked rise i n bl oo d p ressu re -

A t fi rst this
.

a cti o no nth e bl ood vessels wa s th ough t to be th e speci a l

-

fu nc ti on of a d re na li n bu t i t wa s ve ry s oon sh own th a t
,

wh a t a d re na linre al ly d o es is to stimul a te th e ti ny ne rves

which te rm ina te i nth e sm ooth muscles i nth e w a l l s of th e
bl ood vessels a nd so c a use these sm oo th muscles to con
-
,

t ra c t Th e a cti on of a d re na li n i n cl osi ng sm a ll w ou nds

.

an d inra isi ng bl ood p ressu re i s th us i ncide nt a l to i ts re a l

-

fu ncti ono f ro usi ng sm oo th muscles to a ctivi ty An i m .

p orta nt f a c t a b o u t th is a c ti on is th a t the a m ou nt o f
a d re n a li nn o rm a ll y p rese nt i n th e bl ood is no t e nough to
b ri ng this a b out Th e re must be a ni nc re a se a b o ve th a t
.

which is ord ina rily p rese nt i f the bl ood p ressu re is to be

-

r a ised .

T H E R EGU LAT I O N OF A DR E N A LI N S E CR ET I O N — Th e
fa c t j ust me nti oned h a s c a used the questi onto be a sked ,

wh e th e r the re is a ny c onditi oninwh ich the a d re na l gl a nds

bec ome m ore a c tive th a n usu a l a nd p ou r o u t i nt o th e
bl o od l a rge e no ugh a m o u nts of ad re na lin to c a use the
sm oo th muscles o f the bl ood vessels to be exci ted S uch
-
.

a situ a ti on h a s bee n sh o wn to a rise i n c o n nec ti on with

th e u nfa v ora b l e em o ti on s desc ribed i n th e l a st ch a p te r
'

whe re it wa s see nth a t i n time of st ress the b ody ra llies

its p owe rs to meet a neme rge ncy Th e p ossible c onnec
.

ti on o f a d re na li n with th is eme rge ncy re a c ti on c a me to

ligh t whe n it wa s f ou nd th a t no t only a re th e sm oo th
muscles o f th e bl ood vessels stim ul a ted bu t a l l the o the r
-
,

e ff ects listed onp a ge 1 66 a re b ro ugh t to p a ss th ro ugh th e

a cti o n o f a d re n a lin i n the b o dy Th us supp osi ng th a t
.
,

the a d re na l gl a nd s p o u r o u t i nc re a sed qu a ntities o f

a d re n a li ni ntimes o f eme rge n cy th e re is a Chemic al rei n
,

fo rceme nt o f th e e ff e cts p roduced by th e ne rv ous system ,

with th e p resum a ble a dv a nt a ge of i ncre a si ng one s a bility ’

to co pe wi th th e si tu a ti o n Th e re is a g re a t de a l of evi
.

de nce to S h ow th a t th e a d re na ls a re u nde r the c ont rol of

 T HE E FFE C T OF AD R E NALI N 1 77

the ne rv ous system a nd a re a ctu a lly excited to i nc re a sed

,

a ctivity in times o f em o ti on a l st ress .

TH E E FF E CT O F A DR E N A LI N O N M U S CU L AR P O W ER .

I t h a s l ong bee n k now n th a t a pe rs on i n time of eme r

e nc displ a ys u n suspected st re n gth Th e f a mili a r ph ra se
g y .

th e st re ngth of despe ra ti on refe rs to this f a ct Th e re .

is s ome re as on to believe th a t this inc re a sed muscul a r

p owe r ma y be due to the i nflue nce of a d re na li ni n a u g
menti ng the met a b olism of th e muscles S i nce a pe rs on .

i ntime o f st ress a ctu a lly h a s i nhis bl ood m ore a d re n a li n

th a na t o the r times the st re ngth of desp e ra ti on ca nbe

,

thus expl a in ed .

TH E E FF E CT OF A DRE N A LI N O N B A S I C M ETA B O LI S M .

! uite r ece n tly it h a s bee n S h o w n th a t a d r e na li n bel on gs

i nthe g ro up of h orm ones th a t regul a te b a sic met a b olism .

The meth od of dem onst ra ti ng th is f a c t is to i nj ect i nt o

the veins o f a pe rs ona sm a ll a m o u nt of a d re na li na nd the n
to me a su re his resti n g met a b olism Th e e ffect of the i n
.

j ec ti on is to i nc r e a se th e met a b olism c on side ra bly a b ove

m
the nor a l level S i nce no exte rna l w ork is d one i nb a sic
.

met a b olism , th e e ffect o f i nc re a si ng it by i nj ecti ons of

a d re n a li nis to c a use the b ody to p ro duce m ore h e a t th a n

it othe rwise w o uld (Ch a p V I I I ) . Th e re is a ls o evi

.

de nce th a t wh e nth e a m o u nt of a d re na li ni nth e bl ood is

less th a nnorm a l a s for ex a mple f oll owi ng i nj u ry to the
, , ,

a d re n a l b odies ,
th e b a sic met a b olism is a ls o less th a n
norm a l .

TH E R E LATI O N O F H OR M O N E S T o R A CI A L C H AR A CTER
I S TI CS.
—V e ry rece ntly th e suggesti on h a s bee n m a de
th a t the t ra its wh ich disti nguish one ra ce o f h um a nbei ngs
from a nothe r a nd wh ich we c a ll ra ci a l ch a ra cte ristics
, ,

ma y be due to a di ffe re nce i n th e h orm ones wh ich th e

i ndividu a ls of th e di ffe re nt ra ces p roduce The imp ort .

an ce of the suggesti on lies i n th e fa ct th a t si nce ra ci a l

ch a ra cteristics a re h e redit a ry bei ng t ra nsmitted f rom
,

p a rents to child re nge ne ra ti ona fte r ge ne ra ti on the me a ns ,

of i n he rit a nce SO fa r a s ra ci a l ch a rac te ristics a re con

1 78 C H E M I C AL C O R R E LATI O N

ce rned g oes b a ck o f th e ch a ra c te rs themselves to th e

,

gl a nds i n which th e h orm ones a re m a nu f a ctu red T o .

i llust ra te th e st ra igh t bl a ck h a i r a nd reddish ski n o f

,

the Am e ric a n I ndi a n a s c ont ra sted with th e yell ow h a i r

,

an d f a i r ski n o f th e N orthe rn E u rope a n ra ces a re d ue, ,

a cc ordi n g to this view to d i ffe re nces i n h orm ones but

, ,

the di ffe re nces i n h orm one p rod ucti on a re themselves

i nhe rited .

S U M M AR Y . Th rough th e c ombi ned a cti ono f ne rv ous

an d chemic a l c orrel a ti on a ni m a ls devel op symmet ric a lly
,

an d thei r b odily p rocesses p e rf orm thei r fu ncti ons h a r

moni ou sly ; they m a ke th e necess a ry a d j ustme nts to th e
e nvi ronme nt b oth i ntime o f quiet a nd i ntime of st ress
, ,

an d c a rry onth e a ctivities esse nti a l to ra ci a l rep roducti on .

A f a ilu re i na dj ustme nt me a ns to the i ndivid u a l imp a i red

, ,

efficie ncy b odily i nj u ry dise a se or d e a th ; to th e ra ce

, , , ,

f a ilu re i npe rpetu a ti on wh ich to s ome exte nt no m a tte r

, ,

h ow slight diminish es th e b re a d th o f th e f ront which th e

,

nex t or succeedi ng ge ne ra ti ons must p rese nt to thei r eh

vi ron ment if the ra ce a s a wh ole is to su rvive .
 C HA P TER X VII

DI S E A S E S DUE TO M ALAD ! U S T M E NT

D I S EA S E M AY B E D E F I N ED i n ge ne ra l te rms nor as ab

ma li ty i n th e fu ncti oni ng o f livi ng p rot opl a sm ; i n o th e r

w ords a s a c onditi onof th ings i nwhich met a b olism is not
,

p roceedi ng a s it S h ould M et a b olism (see Ch a p V I I I )

. .

is a n a ff a i r of th e individu a l cells Th e distu rb a nce of

.

met a b olism wh ich c onstitutes dise a se is the refore i nthe

'

l a st a na lysis a ls o a n a ff a i r of i ndividu a l cells B ut a s

, .
,

the p revi ous ch a pte rs i n th is secti onh a ve t ried to m a ke

.

cle a r th e cells wh ich m a ke up th e b ody do not fu ncti on

,

i ndepe nde ntly ; wh a t one cell d oes ma y i nfluence the a o

ti vi ty o f cells eve n a t g re a t dist a nces withi n th e b ody .

It is evide nt the refore th a t m a l a dj ustme nt or th e f a il

, , ,

u re of the v a ri o us cells to i n te ra ct p rope rly is a fruitful ,

Ca use of dise a se For ex a mple if th e cells th a t m a ke up

.
,

the he a r t muscle do not fu nc ti on p rope rly the ci rcul a

ti onof bl ood is imp a i red a nd th e cells th ro ugh ou t th e b ody
will su ffe r th e ex a ct degree depend ing on h ow fa r the
,

t ra nsp ort a ti on System f a lls S h or t of norm a l efficiency .

TH E M A LAD J U S T M E N T S f rom wh ich cells su ff e r a re ofte n

the immedi a te resul t o f imp rope r fu ncti oni ng of the
mech a nisms for c orrel a ti on Thus we h a ve a g roup of
.

dise a ses due to imp a i red h orm one c ont rol a nd a nothe r ,

g roup due to imp a i red ne rv o us c ont rol Th e m a l a dj ust .

ment ma y be nut riti ona l th a t is to sa y th e m a te ri a ls

, ,

necess a ry for met a b olism ma y f a il to be fu rnish ed

o f p rope r s orts or i n c orrect a m o u n ts A thi rd g roup .

o f m a l a dj ustme n ts ma y result from mech a nic a l i n

j u ri es o f on e s or t or a no the r a lth o ugh with reg a rd to
,

these it sh ould be no ted th a t the immedi a te results of me

ch a n i ca l i nj u ry a re no t ordi na rily cl a ssed a s dise a s es .

1 79
1 80 DI S E A S E S DUE TO M ALAD J U S T M E N T

The re a re ce rt a i nrem o te e ff ects o f inj u ry h oweve r wh ich , ,

a re p rope rly c a lled d ise a se bec a use th ey i n v olve fa ilu re i n

th e inte rn a l a dj ustme n ts of the cells Th ese will be con .

s idered b riefly i n a subseque n t p a ra g ra ph .

Fi na lly the re a re th e nume rous a nd se ri o us m a l a d

,

j ustme nts which a rise f rom the st ruggle for exis te nce
a m on g c ompeti ng org a nisms a nd wh ich a re f a mili a r i nthe
so c a lled i n
-
fecti o u s dise a ses For c onve nie nce i n p re
.

sen ta ti on th e dise a ses which f a ll u n

,
de r this l a tte r h e a d ,

i nste a d o f bei ng included h e re will be c onside red sep a

ra tel y i n the secti on on
,

A ss oci a ti ons of Org a nisms ”

(Ch a p XX )
. .

N ATU R E OF D I S EA S E M et a b olism is m a de up of th ree

.

diffe re nt st a ges (1 ) the t a ki ng up o f m a te ri a ls by the

,

cells (2) th e u tiliz a ti on of th ese m a te ri a ls i nth e chem

,

ic a l p ro cesses wh ich go on withi n th e cells a nd (3 ) th e ,

disch a rge of th e w a ste p roducts wh ich r esult f ro m th e

chemic a l p ro cesses D ise a se is a distu rb a nce eith e r i nth e
.
,

a mou n t o f met a b olism i n wh ich c a se the qu a n

,
tities of
m a te ri a l c onsumed th e v olume o f th e ch emic a l a ctivity
, ,

an d th e a m ou nt o f w a ste p roducts p o u red o u t ma y be

i nc re a sed o r di mi nish ed ; or i t ma y be a ch a nge i n th e
cha ra c ter o f th e met a b o lism i nwh ich c a se th e subst a n
,
ces
u tilized by th e cells will be d i ff e re nt th e ch a ra cte r o f the ,

chemic a l p rocesses will be ch a nged a nd th e w a ste p rod

’

nots p o u red o u t will be of d i ff e re nt s orts I t is t rue o f .

,

c ou rse th a t i nm a ny dise a ses — a nd pe rh a ps i n m o st

,

b o th th e a m ou nt a nd ch a ra cte r o f th e met a b olism a re

a lte red .

S Y M PTO M S O F D I S EA S E By defi niti on a sympt om o f

.
,

dise a se is a b odily m a nifest a ti on resulti ng f rom th e dis

tu rb ed met a b olism wh ich c o n stitutes th e dise a se itself .

A h e a l thy b ody fu ncti oni ng norm a lly h a s ce rt a i nrep re

, ,

sen ta ti v e a ppe a ra n ces wh ich a re rec og nized a s a t tributes

o f h e a lth I n ma n fo r ex a mple th e re is a ce rt a i nra n
.
, ,
ge
o f ra pidity of h e a rt be a t wh ich is ch a ra cte ristic o f g o o d
-

he a l th Whe nthe h e a rt ra te is outside this ra nge eithe r

.
,
 S YM P T O M S DUE TO C HANG E S 1 81

sl owe r or m ore ra pid it usu a lly c onstitutes a symp tom

,

o f dise a se . In so c om p lic a ted a b ody a s th a t o f ma n ,

sympt oms a re of v a ri ous s orts bec a use distu rb a nces of

met a b olism ca np roduce so m a ny di ff e re nt e ff ects I t is .

ou t o f th e questi onina b ook o f this s ort to m a ke a com

l e te l ist o f the sympt o ms o f th e di ff e re nt ki nds o f dis

p
e a se yet s ome of th e m ore imp ort a nt ma y be me nti oned
,

i norde r to sh ow th eir rela ti onto th e u nde rlyi ng de ra nge

me nts of met a b olism .

S Y M PTO M S D U E T o C H A N GES I N R ATE OF M ETA B OLI S M .

—P rob a bly the c omm onest distu rb a nce of met a b olism

is th a t i nwh ich the re is a whippi ng u p of th e chemic a l -

a ctivity c a usi n
,
g m a te ri a ls to be used up a t m ore th a nth e
usu a l ra te ; th e chemic a l p rocesses withi nth e cells go on
m ore ra pidly th a nnorm a l a nd a n i nc re a sed v olume of
,

w a ste m a te ri a l is p ou red ou t The c omm onest sympt om

.

o f this p a rticul a r c on diti onis th a t rise i nb ody tempe ra

tu re which c onstitutes fe ver I t is t rue th a t th e tempe ra
.

tu re of th e b ody ca nrise to th e feve r p oi nt with o u t met a b

ol i sm a ctu a lly bei n g inc re a sed th ro ugh i nte rfe rence wi th
,

the l oss from th e b ody of the h e a t th a t is p roduced i nnor

ma l met a b olism bu t i nth e v a st m a j ority o f feve rs the re
,

is a n unde rlyi ng i nc re a se i n th e b a sic met a b olism An .

oth e r symp t om r el a ted to feve r is th e ra pid w a sti n g a wa y -

of the b o dy .Obvi o usly a ny one c onsumi ng m a te ri a l ra p

,

idly must su ffe r b odily w a st a ge u nless h e repl a ces the ma

teri a l with equ a l ra p idity ; bu t i n m ost feve rs f oo d con
sumpti on is sligh t or a bsent Acc omp a ny i ng feve r a re .

such symp t oms a s th e ra p id he a r t ra te menti oned e a rlie r ,

a h ot dry S ki n h e a d a che a n
, ,
d th e ge ne ra l feeli ng o f ill
,

being know na s ma la i se Inc a s es whe re th e dise a se t a kes

.

th e form o f dec re a sed met a b olism the re is usu a lly a l owe r

i ng o f b o dy tempe ra tu re a cc omp a nied by feeli ngs o f
,

l a ssitude a nd sl owed he a rt a cti on .

SY M PT OM S D U E T o A LTER ATI O N I N T H E C H AR A CTER

O F M ETA B O L I S M a re o f so m a n y ki nd s th a t only a few
ca n be me n ti oned Th e sympt oms o f m a ny ne rv o u s
. .
1 82 DI S E A S E S DUE TO M AL AD J U S TM E N T

dise a ses co me u nde r th is he a d a s do sympt oms o f such

,

dise as es a s di a betes to be sp oke no f sh ortly I t is p rob

, .

a ble th a t th e v a ri o us a ches a n d p a i ns which used to be

c a lled rh eum a tism a re sympt oms bel ongi ng to th is c a te
g ory a nd i ndic a ti ng th a t th e ch a ra cte r of the met a b olism
i ns ome o r a ll o f the cells o f th e b ody is d ep a rti n g f rom
norm a lity N ume rous symp t om s t ra ce a ble to deficie ncy
.

i ndi et i nrespect to vit a mi n s (Ch a p X ) a ls o c ome unde r .

this h e a d .

I PA
M I R ED H OR OM N E C ON T OR L — I n Ch a pte r X V I i n
.
,

wh ich chemic a l c orrel a ti on wa s c onside red the norm a l ,

a cti ono f s ome of th e h orm on es wa s desc ribed a nd i ncon ,

necti onwith th e desc rip ti onit wa s p oi nted ou t th a t dis

turb a n ces i nh orm on e p roducti onc ould no t f a il to b ri ng
a b o u t a lte ra ti ons i n th e met a b olic a ctivities of s ome or
a ll o f th e cells o f th e b ody D ise a ses of this ch a ra cte r
.

a re u n fortu na tely f a i rly p rev a le nt Th ey a re d i fficul t to

.

de a l with bec a use up to th e p rese nt time a lm ost nothi ng

i s know n a s to th e u nde rlyi ng c a use o f imp a i rme nt of
h orm one p roducti on I t is k now n th a t h orm one m a nu
.

fa ctu ri n g gl a n ds bec ome eith e r u nde r a ctive or ove r -

a ctive bu t why th e gl a n
,
ds S h o uld thus dep a rt f rom thei r
usu a l deg ree o f a ctivity or wh a t regul a tes th a t a ctivity
,

i n th e fi rs t pl a ce is for th e m ost p a rt u n
,
k now n Evi .

dently th e re is he re a g re a t field for fu rthe r bi ol ogic a l a n d

medic a l i nvestig a ti on T 0 illu st ra te h ow dise a se is c a used
.

by imp a i red h orm one c ont rol s ome ex a mples a re ci ted i n

p a ra g ra ph s bel ow .

U N DER A CT I V I T Y OF T H E T H Y R O
-
I D — S ometimes it .

h a ppe ns i n i ndividu a ls th a t th e thy roid gl a nd bec omes

imp a i red a nd p roduces less th a n the norm a l a m ou nt o f
thyroxi n; th e resul t is to dep ress th e b a sic met a b olism
an d for th e i ndividu a l to S h ow a se ries o f sympt o ms which
a re th e di rec t c on seque nce o f this dep ressi on; a m ong
these sympt oms ma y be me nti oned a dimi nish ed u tiliz a
ti ono f f ood Obvi ously if th e re is less met a b oli sm less
.
,

f ood is requi red S ince the i ndividu a l is quite likely to

.
 UND E R —A C TIVIT Y OF TH Y R O ID 1 83

ea t a t b out the usu a l ra te he is a lm ost su re to dep osit

a ,

the su rplus inhis b ody i nth e f orm o f fa t I t is pe rh a ps .

not g oi ng too fa r to sa y th a t i n ge ne ra l pe rs ons wi th a

te nde ncy to obesity a re pe rs ons i n wh om the thy roid
gl a nd te nd s to be S lightly less a ctive th a nnorm a l .

Anoth e r sympt om which displ a ys itself p a rticul a rly ,

whe re the deficie ncy of thyroxi nis m a rked is a dec re a se ,

inth e efficie n cy of th e nerv o us sys tem N e rv ous a nd i h .

tell ectu a l sluggish ness a re evide nt i n seve re c a ses bei n

,
g
so p ron ou n ced a s to a pp roa ch imbecility Th e re a re sev .

e ra l hum a n st ra i ns know ni nwh ich th e thyroid gl a nd is

FI G 4 6 . D ra wing fro m a ph oto gra ph t o S h o w th e a p pea ra n

. ce o f

child , 23 m on ths ol d , wi t h d efi ci ent t hyro id , a nd th e s a m e chil d a f ter

a dmi n is tra ti on o f S h eep s t hyroid for 1 1 m onths
’
.

he redit a rily eithe r se ri ously defect ive or wh olly i na ctive .

Child re ni nh e riti ng th is deficie ncy a re d oomed from bi rth

to p a rti a l or c omplete idi ocy if na tu re is a ll owed to t a ke
i ts c o u rse One of th e st riki ng medic a l t riumphs of th e
.

l a tter p a rt o f th e l a st ce ntu ry wa s th e disc o ve ry th a t ex

t ra cts of the thy ro id gl a nd of me a t a nim a ls c o uld be a d
ministe red a s a p a rt o f th e diet to pe rs ons with deficient
thy roid a nd th a t th e b ody w ould thus obt a i n e nough of
,

the h orm one for its requi reme nts Complete cu res h a ve
.

bee n a nd a re bei ng w rough t thus i n c a ses i n which de

fici ent thy roid is the only se ri ou s a b norm a lity (Fig .
1 84 DI S E A S E S DUE TO M ALAD ! U S T M E N T

OVER A CT I V ITY O F T H E T H Y RO ID
-
The reve rs e ch a nge
.

i nth e thyroid ma y a ls o ta ke pl a ce a n d excessive a m o unts

o f sec reti onbe p roduced Whe nth is h a ppe ns the ge ne ra l
.

level of b a sic met a b olism is ra ised , the symp t oms ex

h i b i ted bei ng th ose resul ting f rom i nc re a s ed met a b olism .

I nc re a sed f ood supplies a re na tu ra ll y requi red to c a rry

o nth e h eighte n ed met a b olism a nd su ffe re rs f rom this con
d iti on a re i n v a ri a bly excepti ona ll y h e a rty e a te rs not ,

withst a ndi ng wh ich they a re u su a lly em a ci a ted Th e .

i nc re a sed met a b olism o f th e ne rve cells h a s as w o uld be

-
,

expected th e c ont ra ry e ff ect to th e l owe ri ng of me ta b

,

oli sm th e n
,
e rv ous p rocesses a re speeded up U nfortu .

na tely th is d oes not b ri ng a b ou t a s one migh t exp ect

di ng imp roveme n
, , ,

a c orresp o n t in i ntellectu a l p owe r On .

the c ont ra ry t h e resul t is a ra the r m a rked i rrit a bil ity

, ,

t ogethe r with a te nde ncy to inst a bility of disp ositi on

which i n sev ere c a ses a m ou nts to me nta l de ra ngeme nt
.
.

I t is evide nt th a t efficie nt ne rv ous fu ncti oni ng requi res

th a t th e b a sic met a b olis mo f th e ne rve Cells be pitched on -

neithe r too l ow nor too high a sc a le Th e qu a lity o f opera

.

ti ono f th e ne rv ous mech a nism is not dete rmi ned by the

ra te a t which i t w orks bu t by the u n de rlyi ng deg ree of
excelle nce o f th e a pp a ra tus itsel f .

P ITU ITA RY G LA N D I M PA I R M E N T — The re is a dise as e

f rom wh ich a dul t h um a n bei ngs s ometimes suff e r i n ,

wh ich v a ri o us b ones resu me g row th a fte r h a vi ng a tt a i ned

th e norm a l a d ult c o nditi on One b one th a t is c om monly
.

a ff ected is th e l owe r ja w Th e p a tie nt fi rst bec omes

.

a w a re th a t s ometh i n g is w rong by o bse rvi ng th at his

tee th n o l on ge r meet th a t h is l owe r ja w h a s g row nto a n
,

exte nt suffi cie nt to th row th e l o we r t eeth o u t o f a lig n

.

me nt with th o se o f th e uppe r ja w Th e b ones o f th e ex

.

tremi ti es ma y a ls o resume g ro wth e n l a rged k nuckle

,

j o i nts a nd e nl a rged fi nger a nd toe j oi nts bei ng a result

-
.

S tudy o f this d ise a se sh o ws th a t it is due to a n i nc re a se

i nth e a ctivity of th e p i tui t a ry b ody .

D I AB ET ES This dise a se which resul ts f rom deficie ncy

.
,
 I M P AI R E D N E R V O U S CO NT R O L 1 85

of the p a nc re a tic h orm one is u nf ortu na tely c omm ona nd

,

a ppe a rs to bec ome m ore so . A v a st a m o u nt o f study h a s

beendev o ted to it o f rece nt ye a rs a nd while no imp ort a nt
,

p rog ress h a s bee nm a de i np reve nti ng th e h orm one ma n -

u fa ctu ri n g cells f ro m g o ing w rong o r i nr epl a ci ng th e defi

cient h orm one a s is d one whe nt h e thy ro id gl a nds f a il
,

to fu n cti on by a dmi niste ri ng m a te ri a l i nth e food it h a s

, ,

been f o u nd p o ssible i n m a ny c a ses to de a l successfully

with th e dise a se th rough a st ric t regul a ti on o f the diet .

Si nce the e ffect of p a nc re a tic h orm one deficiency is to i m

p a ir th e a b i li ty of the tissues to use sug a r th e meth od of
,

t re a tme nt is obvi ously th rough regul a ti ng th e diet so th a t

the p a tie nt ca nget a l ong with out sug a r; Up to the p res
en t this is the di recti oni nwh ich p rog ress h a s bee nm a de
, .

I M PA I R ED N ER V O U S CO N TR O L — Wh e n one c onside rs
.

the c omplexity of th e a dj ust or mech a nism i n the h ighe r

an im a ls t ogeth e r with th e depe ndence of fu ncti ona l me
,

ta b ol i sm on th e ope ra ti ono f this mech a n ism th e p ossi

,

b ili ti es of distu rb a nces i n met a b olism a risi ng th rough

ne rv o us imp a i rme nts a re see n to be m a nif old ; Th e
imp a i rme nt of ne rv o us c ont rol ca n be a tt ributed i n ,

gene ra l to one of two th i ngs a ctu a l dest ructi onof ne rv

, ,

o u s tissue whe r eby n erve p a thw a ys a re b roke n or a lte r ,

a ti on s i n th e met a b olism withi n the n erve cells wh e reby

they a re c a used to fu ncti oni ns ome a b norm a l m a nne r .

I M PA I R M E N T S D U E T o T I S S U E D E S TR U CTI O N The re
a re v a ri o us dise a ses wh ose symp t o ms a re th e di re ct result
o f the dest ructi ono f n e rv ous tissues Am ong th ese is a
.

dise a se p revi ously me nti oned l o c om ot or a t a xi a m ore cor

, ,

rectl y c a lled ta bes i nwh ich th e re is dest r u cti on o f n

,
e rve
tissue i nth e l owe r end of th e S pi na l c ord with c onseque nt
i nte rrupti onof imp ort a nt se ns ory p a thw a ys f rom th e legs .

Inthe dise a se i nf a ntile p a ra lysis th e re is a ls o a destru c

,

ti on of ne rve tissue withi n the sp i na l c ord i n th is c a se

,

a ff ecti n g m ot or ne rve cells a nd thus c a usi ng muscul a r

-

p a ra lysis Ina p oplexy the re is i nj u ry to th a t p a rt of th e

.

b ra in whe nce p roceed th e ne rv o us impulses by wh ich

1 86 DI S E A S E S DUE TO M ALAD J U S T M E N T

v oli ti ona l muscul a r a cts a re c a r ried on a n i nj u ry which ,

a g a inresults i np a ra lysis .

I M PA I R M EN T D U E T O A LTERATI ON O F N OR M A L M ETAB O
LI S M .
— Th e h igh e r b ra in fu nc ti ons na mely a ss oci a ti on
, , ,

v oliti on a pp reci a ti on(a s of a rt or li te ra tu re ) the f a culty

, ,

o f re a s on wh ich is th e high est devel opme n

,
t of the p rop
erty o f a ss oci a ti o n th e feeli n
,
gs or em o ti ons the c ondi ,

ti ono f c on sci ous ness or a w a re ness a ll a cc omp a ny a nd in

,

a l l p rob a bility depe n d on th e fu ncti ona l met a b olism i n

the highe r ne rve cells I t foll ows th a t a lte ra ti ons i nth is
-
.

met a b olism a re likely to result i n pe rve rsi ons of these

high me nt a l fu ncti ons On a cc ou nt o f th e delic a cy a nd
.

c omplexity of th e ne rv ous i nte ra cti ons wh ich c onstitute

me nt a l fu ncti oni ng a sligh t distu rb a nce o f met a b olism
,

ma y a nd freque ntly d oes le a d to m a rked d ep a rtu re f rom

norm a l me nt a l a nd em oti ona l a ctivity a nd such distu rb ,

an ce is likely to m a nifest itself i n tu rn in a lte ra ti ons i n

b odily fu ncti oni ng Conditi ons i n wh ich this situ a ti on
.

p rev a ils to a m a rked d egree a re f a mili a r to medic a l p ra e

ti ti o ners a n d a re g ro uped u n de r th e ge ne ra l na me o f
h ys teri a . Th e chief ch a ra cte ristics o f th is c o nditi on
a re : fi rst a ne n h a nceme nt o f the b odily e ff ects o f em oti on ,

so th a t vi ole n t outbu rsts o f joy s orrow or a nge r result

, ,

f rom t rivi a l c a uses ; a nd sec ond a l oss o f th e se nse of p ro

p orti oni nm a king a ss oci a ti ons so th a t f a ct ors which to a
,

norm a l i ndividu a l a re eithe r o f mi nor imp ort a nce or a re

deemed u nw orthy of i nclusi oni n th e a ss oci a tive p rocess
a re ex a gge ra ted by th e hyste ric to bec ome o f sup reme i m

p ort a nce Wh e n one rec a lls th a t th e u se o f a ss oci a ted

.

mem ory a s a guide to c onduct will evide ntly c a use th o se

a ss o ci a ti ons whi ch a ppe a r m o st imp ort a nt to t a ke fi rst
pl a ce i n th e regul a ti on o f c ond uct it is cle a r th a t this
,

ex a gge ra ti ono f mi nor or u nw orthy a ss oci a ti ons with th e ,

a cc omp a n yi ng mi nimizi ng o f re a lly imp ort a nt mem ories ,

is b ou nd to le a d to dep a rtu res f rom norm a l beh a vi or .

N EU RA S T H E N I A is th e tech nic a l na me o f a c o nditi on

c omm only c a lled ne rv ou s p ros tra ti on I t is a no the r con .
 T R E AT M E NT OF N E R V O U S DI S O R D E R S 1 87

diti on due to imp a i rme nt of met a b olism i nthe ne rv ous

system usu a lly di rectly t ra ce a ble to ove r f a tigue p ro
,
-
,

t ra cted ove r Work or c onti nu ous w orry I t is ch a ra cte r

-
, .

i z ed by b odily restless ness by i na bility to obt a i n ,

ref reshme n t in Sleep a nd by u ndue a t te nti on to se lf

, .

N eu ra sth eni cs usu a lly th i nk a b o ut themselves a nd thei r

b odily c onditi onto the exclusi onof m ore i nte resti ng a nd
m ore imp ort a nt t opics
,
.

D I RE CT T R EATM E N T O F N ERV OU S D I S ORDER S —I na s .

much a s the physi ol ogic a l b a sis of ne rv o us dis orde rs lies

i ndistu rbed met a b olism of ne rve cells the l ogic a l meth od -
,

of t re a tme n t is obvi ously to ta ke such me a su res a s will

rem o ve the c a use o f distu rb a n ce Fro m wh a t h a s bee n .

s a id i np revi ous ch a p te rs i t will be cle a r to th e stude nt

th a t this is a ve ry difficult m a tte r owing to ou r impe rfect ,

knowledge of th e norm a l fu ncti oni ng of the cells of the

c ent ra l ne rv ous system B u t wh erev er th e dist u rbi ng
. .

c a use ca n be f ou nd a nd rem oved rec ove ry o ccu rs We , .

ma y the ref ore c onfide ntly expect th a t with a dv a nci ng

, ,

k nowledge of the ne rv ous system will c ome th e disc ove ry

’

o f p ro cedu res a pp rop ri a te to the t re a tmen t of dise a ses of

this typ e .

TREAT M E N T OF N ERV O U S D I S ORDER S BY P S Y CH OT H ER

AFY . In hyste ri a a n d to a slightly less exte nt i nneu
,

ra sth en i a b o th th e i ntellectu a l a nd em oti ona l st a tes of th e

,

p a tient a re a b norm a l chiefly i nthe di recti onof th e exa g

,

gera ti on of ide a s or em o ti ona l m a n ifest a ti ons which i n ,

thei r p rope r pe rspec tive a re h a rmless e nough bu t wh en ,

d omi na nt a re b o th u ndesi ra ble a nd u nwh ol esome Ass o .

ci a ted mem ories are est a blished a s h a s bee n emph a s ized

p revi ously i nthe cent ra l ne rv ous systems a s a resul t of

,

the i np ou ri ng of se ns ory stimuli th rough th e org a ns of

sense I t f oll ows the re fore th a t they ma y be i n p a rt
.
, ,

c ont rolled by c ont rolling the se ns ory stimuli to which the

p a tie nt is subj ected .

Th e t ra ined physici a nu nde rst a nds th is fully a nd wh ile ,

seeki ng the re a l u nde rlying c a use of distu rbed ne rve cell -

1 88 DI S E A S ES DUE TO M ALAD J U S T M E NT

met a b olism i norde r to c o rrect it a ls o t a kes p a i ns to p re ,

sc ribe e ffectively to th e p a tie nt such a g ree a ble a nd i nte r

esting se ns ory s timul a ti ons a s will d ive rt his mi nd f rom
th e u nwh oles ome a ss o ci a ti ons a nd repl a ce them with new
an d f a v ora ble ones .

M a ny well i nte nti oned bu t ignora nt cults h a ve bee n

-

f ou nded ona nimpe rfec t u nde rst a ndi ng of thi s me th od o f

tre a tme nt . Th ey of c o u rse a chieve a ce rt a inme a su re of
, ,

success i nde a li ng with ne rv ous dis orde rs by pu rely p sy

ch ic me a ns bu t equ a lly o f c o u rse do no t remedy the
, , ,

a ctu a l c a use Th ey c onstitute m ore ove r a public

.
, ,

me na ce i n m a ny i nst a nces bec a use th ey c a rry thei r ,

me th od o f t re a tme nt o ve r to dise as es o f a l l s orts wh e re

th e psych op a thi c eleme n t is eithe r e nti rely l a cki ng or
of n egligible imp ort a nce a s for ex a mple sm a llp ox or
, , ,

diphth e ri a .

M A L NUTR I TI O N By m a l nu t riti on is me a nt a ny f a il
.
;

u re o f p ro t o pl a sm to obt a i n th e m a te ri a ls wh ich a re

esse nti a l to its successful met a b olism Th e m ost obvi ous .

c onditi o n of this s ort is simple st a rv a ti on which is no t ,

o rdi n a rily i n cluded a m ong dise a ses S t rictly spe a ki ng .

,

dise a ses o f m a lnut riti ona re d ue to l a ck of one or seve ra l

o f th e p a rticul a r items wh ich th e cells req ui re For .

ex a mple g rowi ng child re n a re s ometimes i nsufficie ntly

,

supplied with lime s a lts or i t ma y be th a t for s ome oh

,

scu re re a s on thei r p ro t opl a sm is u na ble to u tilize lime

s a lts eve n wh e n supplied i n a bu nd a nce I n eith e r c a se .
,

norm a l b one g row th is imp ossible giving rise to d ise a ses ,

o f which ri cke ts i s a type with excessively b owed legs

,
s

an d o th e r i ndic a ti ons th a t th e b ones h a ve no t obt a i ned

sufficie nt lime I n Ch a pte r X i nte resti ng f ood a ccess o
.
,

ries k n ow na s vit a mi n s a re me nti oned Th e re a re a nu m .

b er o f dise a ses k n ow nwhich a re due to sc a rcity o f these

i nthe diet I t ma y h a ppe nth a t th e mech a n

. ism for t ra ns
fe rri ng o xyge n(see Ch a p X I ) bec omes i nefficie nt th ro ugh
.

th e f a ilu re o f th e b ody to m a n uf a ctu re a dequ a te a m o u nts

o f h ema gl o b i n to p rovide for the requi red t ra n sp o rt o f
 P O I S O NI NG 1 89

o xyge n Th e dise a se a nemi a is a nexp ressi onof this con

.

diti on .

P OI S O N I N G . By fa r the g re a te r numbe r of ills to

which flesh is h eir a re the result of di rect inte rfe re nce with
the met a b olism of p rot opl a sm th rough the a ge ncy of p a r
ti cu l a r chemic a l subst a nces which bec a use they a ff ect
,

p rot opl a sm i n this m a nne r a re know n a s p ois ons

,
.

P ois onous subst a nces a re ve ry c omm oni nna tu re it bei ng ,

a cu ri ous fe a tu re o f pl a n t met a b olism th a t a g re a t m a ny

ki nds of pl a nts p roduce a nd st ore with i n s ome p a rt of
thei r st ructu re chemic a l subst a nces wh ich a re h a rmful to
an im a l p ro t opl a sm The re a re a ls o a numb er of i norg a nic
.

or mi n e ra l c omp ou nds known th a t a re p ois onous to p ro

top l a sm . Whe re th e b odily distu rb a nce is due to the
t a ki ng i nt o th e st om a ch o f a k no w np o is onous subst a nce ,

th e c onditi onis refe rred to a s a cu te p o is oni ng a nd is not ,

ordi n a rily c on sidered to be a dise a se B u t if the p oi s onIS

.

int roduced int o th e b ody a s p a r t o f supp o sedly g ood food ,

or is p roduced by a n y me a ns w ith i n th e b ody itself it

,

the nbec omes a c a use of dise a se .

D I S EA S E D U E T o M E CH A N I CA L I N ! UR Y — Th e e ff ect
o f mech a n ic a l i nj u ry or of a b u rn i s to dest roy s ome l iv
, ,

in g tissue Obvi ously th e m ore seve re the i nj u ry or the

“

.
,

bu rnthe m ore such tissue is d est royed D est royed tissue .

is i np a rt S l o ughed off a t th e su r f a ce of th e b ody a nd i n ,

p a rt t a kenup a nd a bs orbed withi nth e b ody One of the .

imp ort a nt medic a l disc ove ries o f th e Gre a t Wa r wa s th a t

the p roducts a bs orbed i nt o th e b ody from i nj u red tissues
a re highly p o is on ous . S u rge ons h a ve l ong k now no f a con
diti onof c oll a pse f oll owi ng seve re i nj u ry to wh ich th ey ,

h a ve give nthe na me w o u nd Sh ock I t is now k now n

.

th a t the symp t oms o f w ou nd S h ock a re due to th e p ois on

in g o f the b o dy by th e a bs orp ti on of th e dec omp o siti on
p roducts of th e i nj u red tissues It h a s a ls o bee n S h ow n
.

th a t th e p rol onged su ff eri ng a nd sl ow ness of rec ove ry i n

c a ses of seve re bu rns a re due to th e a bs orp ti oni nto the
b ody of p ois onous p roducts from the bu rned a re a s Gre a t .
1 90 DI S E A S E S D UE TO M ALAD J U S T M E N T

a dv a nces i n t re a tme nt p romptly f oll owed th ese di scov

e ri es s i nce a ll th a t is necess a ry to p reve nt the p ois oning
,

is to m a ke su re th a t none of these p ois ono us p rod ucts

ca nbe a bs orbed i n to the system . The su rge ond oes th is
by c a refully rem ovi ng a ll i njured tissue which migh t de
'

c om p ose a nd bec ome a s ou rce of p ois on .

 C HAPTER X VIII

A S S OCIATI O N O F O R GANI S M S

N ATU RE OF A S S O CI ATI O N . Th e numbe r of i ndividu a ls

c omp osi ng a S pecies rem a i ns p ra c tic a lly th e s a me from
ce ntu ry to centu ry The re ma y be a m a rked i nc re a se
.

whenc onditi ons a re u nusu a lly f a v ora ble a nd a g a i nthe re ,

ma y be le a nye a rs whe nthe re is a nequ a lly sh a rp decli ne ,

but ove r a l ong p e ri od of ye a rs th e a ve ra ge is f a i rly u ni

f orm Fu rthe rm ore th e numbe r of y ou ng p roduced
!

.
,

by the membe rs of th e species is g re a tly i nexcess of th ose

a rrivi n g a t m a t urity (see Ch a ps XX V I a nd XXX V ). .

S ome o f th e y o u ng a re d rive n int o u nf a v ora ble

situ a ti ons o the rs a re dest royed by e nemies a nd
, ,

a lw a ys th e re is a fie rce st ruggle f or f ood whe re

on ly a c omp a ra tive few eme rge a s c onque rors In .

this c omb a t th e i ndividu a ls of m ost S pecies o f

di ffe rent ki nds of a nim a ls a re wh olly self reli a nt depe nd -
,

in g up on thei r own st re n gth a nd kee nness of se nse to

esc a pe dest ructi on Bu t wh ile th e re a re m a ny species
”
.

whe re life is a c onti nu o us free figh t i t ca n not be

,

denied th a t a m ong highe r a nim a ls especi a lly the re a re

a ss oci a ti on s of i ndividu a ls whe re th e st rong p rotect the
we a k a nd displ a y a spi ri t of coOp era ti on a nd mutu a l a i d
a ki n to th a t i n ma n Ag a i n the re a re m a ny ki nds of
.
,

an im a ls a nd pl a nts i n which one i ndividu a l e nte rs i nt o

mutu a lly a dv a nt a ge ous rel a ti ons wi th a no th er a nim a l or
pl a nt of a t o t a lly diffe rent type And fi na lly the re a re
.
, ,

m a ny species which i n th e kee n st ruggle for life

h a ve t a ke n up p osi ti ons on or i n the b odies of o ther
org a n isms from which th ey de rive th ei r nou rishme nt .

The a ss oci a ti on of org a nisms is the refore a b roa d com ,

1 93
1 94: A S S OCIATI O N O F O R GANI S M S

p rehe nsive te rm includi ng the u ni ono f living thi ngs o f

,

th e s a me or diffe re nt s p ecies whe re the results a re of ,

m ost v a ried ch a ra c te r .

F A M I LY G ROUP S H ER DS —Th e simplest ex a m p le of

, .

mutu a l a i d a ppe a rs i n th e c a se of fa mi ly group s wh e re ,

a s i n the c a se o f sw an s geese ce rt a i n a pes a ntel opes

, , , ,

eleph a nts a nd m a ny oth e r species a kee nse nsed p a re nt

, ,
-

le a ds a nd p ro tec ts th e off sp ri ng of diffe re nt a ges .

I n th e he rd a few or m a n y f a mily g ro up s bec ome

me rged i nt o a l a rge r a ss oci a ti onof individu a ls which ma y ,

numbe r th ous a nd s E x a mples a re f o u nd a m ong wild

.

sheep dee r a nd seve ra l species of a ntel ope whe re a

, , ,

fem a le ma y le a d or a m ong seve ra l o the r ki nds o f a nte

,

l opes a nd bis onwhe re one or m ore p owe rful m a les a ct a s

le a de rs Inth is c a se a ls o th e st rong le a d th e we a k to th e
.

feedi ng grou nds a nd p rotect th em a g a i nst th e a tt a cks o f

e nemies .

B EE COLO N I E S — Bey ond this p o i nt seve ra l se ries o f

.

ex a mples c ould be ci ted which illust ra te a n i nc re a s ing

c oope ra ti ona nd divisi ono f l a b or culmi na ting i nc omplex ,

s oci a l org a niz a ti ons such a s th ose of the h oney bee a nt , ,

an d ma n Inthe bumble bee for i nst a nce we fi nd a typ e

.
,

o f c o mmu n a l l i fe a d v a nced a st a ge bey ond th a t o f th e

f a mily g roup or he rd In th e sp ri ng a fem a le th a t h a s
.
,

p a ssed th e wi nte r i ns ome shelte red nook eme rges sele cts , ,

a site for a n est c onst ructs cup s for a mixtu re of h oney

,

an d p olle n th a t sh e c ollects a n
, d on th is ra ises a b rood,

o f f ro m th ree to twe n ty y ou ng These bec ome imm a tu re .

fem a les or w orke rs th a t h e nceforth c a re for oth e r y ou ng ,

e nl a rge th e nest a nd le a ve th e quee nm o the r th e s ole du ty

o f p roduci n g th e f a mily M a les a ppe a r i n th e fa ll to
.

gethe r wi th ce rt a i nfem a les which bec ome quee ns th e fol ,

l owi ng ye a r wh ile a ll th e othe rs die with th e c omi ng of

,

wi nte r .

The h oney bee c ommu nity i na ge ne ra l wa y resembles , ,

th a t o f the bumble bee A quee nh e a ds th e c ol ony ,co m

.

p rising m a ny th ous a nds of imm a tu re fem a les or worke rs ,

 ANT S 1 95

a nd a ttends to th e egg l a yi ng p rocess th a t t a kes pl a ce a t

-

th e a nnu a l ra te of a milli oneggs for seve ra l ye a rs The .

w orke rs a tte nd to a ll of the rem a i ni ng d uties of the

c ol ony c a ri ng for th e y ou ng c onst ructi ng c omb g a the r
, , ,

in g h on ey ev a p o ra ti n
,
g i t to th e p rope r c onsiste ncy ven ,

til a ti ng th e h ive w a rdi ng o ff e

,
n emies a nd o the rwise ,

pl a yi ng a p a rt inone of th e ph a ses of a nim a l a ctivity

m ost a bs orbi ngly i nte resti ng from a h um a nst a ndp oint .

A N TS Am ong th e a nts th e s oci a l org a niz a ti onre a ches

.

th e h igh est devel opme nt i none respect for the re is not ,

on ly a divisi on of l a b or a m ong the i ndividu a ls of th e

c ol ony bu t th ey a re st ructu ra lly fitted to pe rf orm ce rt a i n
duties a nd only these AS i nthe be es a quee nhe a ds the
.
,

c ol ony i nwhich th e membe rs a re wi nged m a les a nd fe

,

m a les desti ned to le a ve th e nest a nd f o u nd new c ol onies

, ,

an d l a rge a nd S ma ll S ized w orke rs a nd s oldie rs The .

w orke rs c onst ruct th e nest c ollec t the food th a t v a ries

, ,

widely i nch a ra cte r a m ong th e h u nd reds of species wh ile ,

the s oldie rs a s is to be exp ected defe nd th e h ome a g a i nst

, ,

the a tt a cks of i nv a de rs ch iefly i nsects Ea ch cl a ss or

,
.

c a ste is thu s i ndispe nsible a nd coOp era teS with the others
to f orm a u n it of a ve ry high type .

A COM PARI S O N AN T AN D M AN ,
Am ong the species .

of a n im a ls th a t f orm mu tu a l be nefit a ss oci a ti ons the a nts ,

a re th e on ly ones th us f a r me nti oned wh o se speci a l

duties a re c orrel a ted with m a rked st ructu ra l p eculi a ri
ties Y et i t w ould be h a z a rd ou s to cl a im f or this re a s on
.

th a t these i nsects rep rese nt a h igh e r type o f c ommu na l

life th a nth e bees B o th a ppe a r to i ndulge i nequ a lly v a
.

ried a ctivities a n d it i s ce rt a i n th a t b o th le a d emi nently

,

successful lives Wh a teve r ma y be th e ve rdict i n this

.

c a se it is evide nt th a t th e s o ci a l org a niz a ti on o f ma n

, ,

with its c omplex divisi on of l a b or a nd coOp era ti on is ,

i ndepende nt of a ny st ructu ra l di ffe re nces a m ong its mem

be rs The plumbe r a nd the bl a cksmith a re physic a lly
.

not di ffe re nt from th e c a rpe nte r a nd b rickl a ye r Thei r .

a ctivities like th o se o f a l l th e o the r cl a ss es o f w orke rs ,

,
 A S S O CIATI O N OF O R GANI S M S

a re a m a tte r of t ra i ning ; a nd t ra ini ng res ts up on high

b ra i ndevel opme nt Th e re is h e re th e el ement of ch oice
'

.
,

o r th e p owe r to di rec t th e a c tivi ties of th e b ody a l on g

an y on e o f seve ra l li nes ; whe re a s th e a nt a nd o the r l o we r

an im a ls a re lit tle m ore th a n dumb m a ch i nes wh o se

st ructu re de termmes wi thin na rrow limi ts thei r l ifetim e
d u ties .

S Y M B I OS I S Th e ex a mples cited to illust ra te c oo pe ra

.

ti ona nd mutu al a i d h a ve thus f a r bee n c once rned with

i ndividu a ls o f a single species Th e re a re h oweve r m a n . y , ,

c a ses whe re one a nim a l e nte rs i nt o a p a rt ne rsh ip o f mu

tu a l be nefit (symbi osi s ) with a noth e r t ot a lly d iffe re nt
ki nd o f a nim a l or one pl a nt bec omes a ss oci a ted wi th
,

an o th e r pl a n t or a pl a nt a nd a nim a l ma y me rge i h
,

terests . Inthe lich e ns for ex a mple th a t i nc rust fe nces

, , ,

ol d roo fs a n d simil a r st ructu res two pl a nts a re i n i n ,

ti
m a te u ni on a g ree n one celled a lg a a nd a fu ngus I t
,
-
.

is a ssumed th a t nut ritive p roducts m a nu f a ctu red by th e

g ree npl a nt a re utilized by th e fu ngus wh ich rep a ys the ,

debt by fu rnishi ng m oistu re a nd offe ri ng p rotecti on .

U nice llul a r pl a nts a re a ls o p rese nt i n l a rge numbe rs i n

m a ny a nim a ls such a s th e g ree n hyd ra sea a nem ones
. , ,

an d ce rt a i n w orms whe re it is believed a be nefici a l

, , ,

i nte rch a nge o f food m a te ri a ls t a kes pl a ce .

M a ny species of a nts feed up on a swe et liquid th a t

issues f rom the b ody of ce rt a i na phids or pl a nt lice wh ich ,

a re sm a ll i n sects feedi ng up onpl a nt j uices I nretu rnth e .

an t p rotects th e a ph id f rom e n emies t ra nsp orts it to ne w ,

feedi ng g ro u nds o r c a rries it u nde rgrou nd whe re food is

, ,

p rovided S eve ra l species of h e rmit c ra bs livi ng i n the

.
,

empty sh ells of s na ils c a rry sea a nem ones a tt a ched to

,

thei r h ouses (Fig The c ra b is p ro tected by th e

.

sti ngi ng cells of th e a nem one wh ich e nj oys a ple nte ou s .

f ood supply i n c onseq ue nce o f bei ng c a rried a b out .

CO M M E N S A LI S M OR M ES S M AT I S M
,
— The re a re m a ny .

ex a mples of th e as s oci a ti on of two di ff e re nt species '

o f o rg a n isms wh e re o ne p a rt ne r only de rives a be nefit .

 CO MM E N S AL I S M 1 97

For ex a m p le , v a ri ous species sp onges a re p rovided with

of

ra th e r sp a ci o us c a na l systems th ro ugh wh ich a cu rre n t

of w a te r is c on tinu a lly p a ssi ng Th ese p a ss a ges a re fre
.

quently i nh a bited by seve ra l species of c ra bs a nd w orms

th a t s a fe from a tt a ck live up on fl oa ti ng nut ritive ma
, ,

teri a l . Th e sp onge g a ins nothi ng by th e p a rt ne rship ,

neithe r d oes it a ppe a r to su ffe r a ny i nc onvenie nce Th e .

s a m e one S ided be nefi t is e nj oyed by m a ny species of

-

c ra bs th a t a tt a ch fra gme nts of s p onges se a weeds a nd

.

, ,

FI G 4 7 . . Hermi t Cra b i nsna il S h ell co vere d wi t h sea a nem ones .

o the r org a nisms to th e b a cks of thei r S hells a nd a re th u s ,

so effectu a lly c on ce a led th a t th ey esc a pe th e a tt a cks o f

m a ny S h a rp eyed e nemies He re a g a i n th e a dv a nt a ge
-
.

a ppe a rs t o be O nth e S ide of the c ra b Th e sm a ll fishes .

th a t fi nd a s a fe ret re a t a m ong th e stingi ng te nt a cles of

j elly fi sh es the c ra bs a nd oth e r org a nisms th a t a re a t
-
,

ta ch ed to the b odi es o f tu rtles a n d th e sm al l a nim a ls ,

th a t live i n the shells of oyste rs s na ils a nd rel a ted , ,

s p ecies a re fu rth e r ex a mples o f this widesp rea d typ e of

,

p a rt ne rship .
1 98 A S S O CIATI O N OF O R GAN I S M S

P ARA S ITI S M I t is a ve ry sh ort step indeed f rom mess

.

ma ti sm to p a ras itism Let th e c ra b living a s a mess

.
,

m a te in th e sp onge begin to feed up onth e subst a nc e of

,

th e sp onge a nd it bec omes a p a ra site P a ra sitism th e re .

,

f ore i s a h a s s oci a ti on with one membe r th e p a ra site

, , , ,

g a ining its livelih ood f rom th e o th e r te rmed the h ost , .

Ass oci a ti ons of th is na tu re a re exceedi ngly c omm on .

With a nim a ls a t le ast i t h a s bee n a ffi rmed th a t every

, ,

species s a ve th e ve ry sm a llest se rves a s h o st for a t le a s t

, ,

on e species o f p a ra site .

M a ny p a ra sites such a s ticks fl e a s a nd leech es l ive

, , , ,

onth e o ute r su rf a ce o f th e h o st a n d a re te rmed exte rna l ,

p a ra si tes Othe rs livi ng i n th e digestive t ra ct muscles

.
, ,

or bl oo d st re a m a rec a lled i n te rna l p a ra sites Al s o th e re .

,

a re m a n y species such a s th e t a pew orm a nd t rich i na

, ,

wh ose org a niz a ti onrequi res th a t th ey spe nd th ei r e nti re

existence in one or a t m ost two or th ree h osts These .

a re p e rm a n e nt p a ra si tes Fi na lly , th e re a re m a ny o the rs

.
,

such a s th e flea a nd m osquit o th a t m ove f reely f rom one ,

h ost to a no th e r a nd a re cl a ssed a s temp ora ry p a ra sites .

P AR A S ITI C P LA N T S Th e re a re a l s o m a ny pl a nts th a t
.

a re p a ra sitic up on o th e r pl a n t s or a ni m a ls Among th e .

high e r types th e best k now n ex a mple is p rob a bly th e

mistlet oe species of wh ich a tt a c k f orest a nd culti
,

v a ted t rees I t is a bush y sh rub (Fig 4 8) with yell ow

. .

ish gree nle a ves a nd is a tt a ch ed to th e h ost by me a ns of

,

speci a lly modified root s wh ich a re believed me rely to a b

s orb w a te r a nd dis solved sa lts Th ese subst a nces a re .

th e n c ombi ned i nto m ore c omplex c omp ou nd s by the

a ctivity o f th e p a ra site itself .

I nth e d odde rs a group o f pl a n

,
ts rel a ted to th e m orn
i ng gl ory a n ,
d a tt a cki ng m a ny ki n ds o f h e rbs a nd l ow
sh rubs th e le a ves a re a bse nt a nd th e stem a ppe a rs like
,

a twi n i ng yell ow or red th re a d (Fig At f reque nt .

i nte rv a ls th e stem s a nd b ra nch es a re p rovided with roo t s

wh ich a bs orb f rom th e h ost pl a nt th e w a te r suga r a nd , ,

mine ra l s a l ts necess a ry f or life o f th e p a ra site Wh e n .

 ANI M AL P A R A S IT E S 1 99

st a rved th e d odde r ma y f orm chl orophyll a nd m a nu

fa ctu re s ome f ood but u nde r f a v ora ble Cond iti ons it i s
,

wh olly p a ra sitic Othe r ex a mples of p a ra sitism a m ong

.

th e l owe r pl a nts a re desc ribed i n l a te r ch a pte rs i n con

-

necti onwith th e subj ect of dise a se .

A N I M A L P ARA S I TES — P a ra sitic sp ecies h a ve bee n

f ound to occu r i neve ry one of th e l a rger a nima l g rou p s

FI G 48 . . M istletoe p a ra si ti c n oa k
o .
~

with th e excepti onof th e sp onges a nd th a t to wh ich th e

sta rfi sh es,
sea u rch i ns a nd th ei r rel a tives bel ong I n .

some of th ese g roup s th e list of such a n im a ls is com

p a ra ti v ely sm a ll wh ile i n o the rs a l a rge n
,
umbe r h a ve
a ssumed th is m od e o f existe n ce .

Am ong th e u nicellul a r species the re a re th ous a nds th a t ,

a re p a ra sites a n
,
d a lth ough th e gre a t m a j ority i nfest th e
,

l owe r a nim a ls th e re a re seve ra l th a t p roduce ve ry se ri

,

ou s hum a n dis orde rs V a ri ou s dise a ses of th e digestive

.

t ra ct rel a psing feve r sleeping sickness a nd m a la ri a a re

, , , ,

c a used by one celled p a ra sites Am ong th e fl a tw orms

-
.

a ls o the re a re seve ra l hu n d red p a ra sitic species fa mil

i a rl y k n ow n a s t a pew orms (Fig 50 ) a n d flukes Th e . .
20 0 A S S OCIATI O N O F OR GAN I S M S

flukes may be eith e r i nte rna l or exte rna l p a ra sites ; th e

t a pew orms i nv a ri a bly i nh a bit the d igestive t ra ct M a ny .

s p ecies of th re a d w orms (Fig 5 1 ) a re likewise p a ra sitic

.
,

FI G 4 9
. . D odd er pa ra si ti c on cock le b urr
-
.

and wh ile a few i nfest pl a nts th e g re a te r numbe r live In

an ima ls a nd a re i n seve ra l c a ses resp onsible for certain
h um a n dise a ses Lice fl e a s a nd ticks a re a ls o d re a d ed
.
, ,

pes ts i n ma ny i ns ta nces especi a lly owi ng to th e fact

,

th a t th ey c a rry dise a se p roduci ng org a nisms

-
.
 S T R U CT UR E OF P A R A S IT E S 20 1

STRU CTU R E P AR A S I TES I t is ve ry ce rt a i n th a t a l l

OF .

existing p a ra sites h a ve desce nded f rom free livi ng non -

,

p a ra sitic a ncest ors Wh e re th e p a ra sitic h a bi t h a s bee n

.

of c om p a ra tively S h ort du ra ti on or whe re th e a ctivities ,

M u lti cep s mu lti ceps f romth e intes

‘

FI G 50 . .
— Adu lt ta pe- w orm , ,

t ine of th e dog .
(N a tu ra l S iz e ; a fter Ha ll ) .

of th e p a ra si te
~

c onfi ned to th e S ki n o f th e h ost th e

a re ,

effects on the p a ra site of such a m ode of life are not es

p e c i a ll y m a rked Th e m o squit o f o r exa mp l e flies f rom
‘

h ost to h ost a nd p ossesses org a ns of l oc om o ti ona nd sen

, ,
.

,
sa
ti onof th e usu a l i nsect typ e On th e o the r h a nd th e re
‘

a re n ume ro u s typ es th a t a re w h olly dep ende nt u pon

on e h ost a n d th ese a re o fte ns o
,

highly m odified th a t thei r r el a

ti on sh i p s a re difficult to dete r

fn i ne In th e S a cculi n an
i

. a ,

an im a l rel a ted to th e b a rna cle s

an d p a ra sitic ona c ra b th e b ody ,

is sc a rcely m ore th a na b a g filled

with eggs a nd a tt a ch ed to a roo t Fh ma
ig En ca psul e d .

‘c
i ifefi;
k f
like a bs orbi ng system pe ne ig igfijfi nfig
tra ti n
. E
g th e t i ss ues o f th e h o st a f ter B ra u n) Fro m K e ll o gg
CO”
. .

an oa n
Inth e t a pew orms th a t cli n g to ,
d D e E Z O OL ,

th e w a ll o f th e digestive t ra ct a nd a bs orb digested

m a te ri a ls th ro ugh thei r b ody w a ll th e org a ns of l oc o ,

m oti ona nd digesti onh a ve e nti rely dis a ppea red In the .
202 A S S OCIATI O N O F O R GANI S M S

gre a te r numbe r of species the eff ects o f p a ra sitism a re not

so m a rked ye t i np ra ctic a lly eve ry c a se th e re a re speci a l
,

cli nging org a ns — h ooks sp ines cl a ws a nd sucke rs, , ,

an d a n u nusu a l devel op me nt of eggs a s the re a re g re a t ,

difficulties in th e wa y o f y ou ng re a ch i ng a new h ost .

On e species o f t a p ew orm for ex a mple l a ys , ,

eggs a ye a r a nd it i s p rob a bl e th a t not m ore th a n

,

on e or two off sp ri n g w ill g a i na nothe r h ost .

D EGEN ERAT I O N The te rm dege ne ra ti ono fte nimplies

.

i nthe p opul a r mi nd th a t a n im a ls which h a ve l ost one or

m ore org a ns a s a result o f p a ra si tism a re u nfit ; th a t a n
a ctive org a n ism with delic a te se nses rel a tively g re a t ,

st re ngth a nd p owerful org a ns of offe nse a nd defe nse is

,

bet te r fitted for life s b a t tle th a n a p a ra site Acc ording

’
.

to such a belief a h orn et or a li on is m ore successful i n

the st ruggle for existe nce th a n a t a pew orm or a h ook
w orm S uch h oweve r is no t th e c a se Th e ve ry f a c t
.
, , .

th a t these species exist is sufficie nt evide nce th a t e a ch is

w ell a d a pted for a p a rticul a r m ode o f life In one c a se .

sti ngs a nd cl a ws help to wi n th e b a ttle ; in th e o the r ,

sucke rs a nd a suit a ble a bs orbi ng system pl a y a nimp ort

an t p a rt. In b o th c a ses th e i ndividu a ls le a ve off sp ri n g
an d th e species pe rsists Bi ol ogic a lly spe a ki ng e ac h is
.
,

equ a l ly successful .

L I F E H I S T OR I ES Th e life h ist ories of p a ra si tes a re i m

.

p ort a nt f ro m a scie ntific st a ndp oi nt a nd p ra ctic a lly th e ,

successful t re a tme nt o f m a ny dise a ses d epends up on a

th orough k nowledge o f th e d evel opme nt o f th e c a us a tive
org a n ism Inthe c as e of m ost ex te rna l p a ra sites the p rob
.

lem is c om p a ra tively simple bu t wi th m a ny o f the i nte r ,

nal p a ra si tes one or two immedi a te h os ts must be

,

i nh a bited be fore th e a d ul t st a ge is re a ched a nd so th e ,

subj ec t bec omes c omplex T wo ex a mples will se rve to .

illu st ra te th is fa ct : A t a pew orm i nh a bits th e dog O ne .

o f its eggs th a t h a s p a sse d f rom th e h o st a nd bec ome

, ,

a tt a ched to g ra ss i s e a te nby a ra bbit

,
Th e ti ny w orm .

h a tch es i nthe i ntesti nes b ores its wa y i nt o a bl ood vessel

, ,
 E FFE C T S O F P A R A S IT E 20 3

a nd is c a rried to a p ositi on i n s ome org a n He re it re .

m a i ns u ntil th e ra bbit is e a te n by a dog a nd th e libe r

‘

a ted w orm a tt a ches it self t o th e i n testi na l c oa t g rows , ,

l a ys eggs a nd th us c ompletes th e cycle ; i nth e h oo kw orm ,

a sm a ll th re a dw orm i n h a biti ng th e i ntesti ne of ma n th e ,

eggs esc a pe from th e h o st devel op to a c onside ra bl e

, ,

deg ree i n th e g rou nd a nd th e yo u ng w orms the n b ore

,

th rough the ski non th e s ol es of th e feet of b a ref oot ed

. n
l a b ore rs Ente ri g th e bl ood th ey a re t ra nsp orted to ,

the lu ngs wh e re th ey b or e th eir wa y th rough th e wa l l

/

, ,

m a ke th ei r wa y to the th roa t a nd a re sw a ll o wed In , .

the i ntesti ne th ey a dhe re by h ooks a nd c om p let e th e ,

cycle of devel opme nt .

E FFE CT S O F P A R A S I TE O N H OS T —I n th e c a se o f .

anan im a l wh ose S ki n is p ie rced by th e p rob oscis of a

m osquit o the inj u ry a nd l oss of b ody subst a nce is u nim
p ort a nt E ve n whe n a p a ra s ite such a s a t a pew orm is
.
, ,

a pe rm a n e nt reside nt th e i nj u ry due to i ts a tt a chme nt

,

an d a bs orpti onof nut ritive m a te ri a ls is no t o f a ny g re a t

imp ort ance I t ma y h a ppe n h oweve r th a t th e S ligh t
.
, ,

b re a k i n th e S ki nor w a ll o f th e digestive tra ct or i n th e

oute r su rf a ce of pl a n ts ma y se rve a s a p oint of e nt ry for
othe r p a ra sites such a s b a cte ri a m olds a n
,
d v a ri ous one , ,

celled a nim a ls wh ose e ffects ma y be of a m ost se ri ou s

,

ch a ra cte r Ins ome c a ses th e sec ond a ry p a ra sites ma y be

.

c a rried in th e p rob oscis o f a fly flea or m osquit o a nd ,

‘

, ,

p a ssi ng th rough th e ski n wh e n th is is pie rced c a use ,

pl a gue S leepi ng sick ness m a l a ri a a nd m a ny othe r te rri

, , ,

ble sc ou rges Als o the re a re m a ny know nc a ses of p a ra

.

sites occu rri ng i n such gre a t numbers th a t th e bl oo d

ves sels of th e h ost h a ve bec ome cl ogged to a se ri ou s a nd
even f a t a l ex te nt Fi na lly th e re a re m a ny p a ra sites
.
, ,

b oth pl a nt a nd a nim a l wh ose w a stes libe ra ted i n th e

, ,

b ody of th e h o st c a use s ome of th e m ost seri ou s dise a ses

,

a flli cti n g m a na n d th e d o mestic a ted a nim a ls .

 C HAPT ER XI X

D ISEA S ES O F P L AN T S

T H E E CO N O M I C I M P ORTA N CE O F P LA N T D I S EA S E ca n
h a rdly be ove rst a ted for it is evide nt th a t th e lesse ning
,

o r p reve n ti on of c rop s is a ve ry se ri ous m a tte r The .

U nited S t a tes D ep a rt me nt of Ag ricultu re estim a tes the

l o ss by pl a nt dise a se i n th e ye a r 1 91 9 a s f oll ows : 1 90 ,

bushels of wh e a t ,
bush els of c orn ,

bush els of p ot a t oes ,

bu shels of
a pples ,
b a les of c ott on .

R ust a l one i ns ome ye a rs h a s e nt a iled the l oss of 200

, , ,

bushels o f wh e a t wh ile sc a b c a us ed i n1 91 9 the

,

l oss of ne a rly bush els of whe a t .

T H E CAU S ES O F D I S EA S E ma y be eithe r li feless or liv

in g; th ey m a y be exte r n a l to th e pl a nt or with i nit ; th ey
ma y c onsist i nu nf a v ora ble fe a tu res of th e e nvi ronme nt
exte rna lly or th e i nte rfe re nce o f othe r org a nisms Only .

th e s econd type f a lls p rope rly u nde r th e h e a d of a s soci

a t i ons o f org a n
”
isms ; bu t bec a u se th e re a re o th e r c a uses
o f dise a se th a np a ra sites we S h a ll a l so c o n
,
side r ex a mples
o f dise a se d ue eith e r (1 ) to u n suit a ble e nvi ronme nt or
( )
2 t o i nte rn a l c a u ses .

U NSUI TABLE ENVI R ON ME NT

T H E SOIL ma y be o f such c omp ositi onth a t it is s a id to

be a lk a li ne or a cid I nce rt a i na re a s i nth e fa r west the
.

p rese nce o f u nusu a l qu a nti ties o f s odium o r c a lcium s a l ts

m a kes th e s oil a lk a li ne ; wh e re a s on th e o th e r h a nd
, ,

espe ci a lly i nwet pl a ces th e s oil ma y be a cid bec a use of

,

the p rese nce i ni t o r i nits s oluti ons o f a cids formed i nthe

, ,

c ou rse of the dec a y o f org a nic w a s te E xce ssively h a rd.

20 4
 TILLAG E 20 5

s oil ma y c a use dise a se bec a use it i nte rfe res mech a nic a lly
with th e g rowth of roots I f the s oil is e xcessively wet
.
,

its p rope r a e ra ti onis p reve nted If it is too dry the pl a nt

.
,

suffe rs from l a ck of w a te r These p roduce a b norm a l or

.

dise a sed c onditi ons i n the pl a nt distu rbi ng norm a l

,

physi ol ogic a l a ctivi ties resulting i n u nusu a l a nd un

,

he a lthy a ppe a ra nce a nd lesse ned c rop p roducti on All .

of these di fficu l ti es a l ka li nity a cidity a s well a s me

, ,

ch a n — ma y be a llevi a ted by till a ge

.

i ca l defects .

T I LLA GE Inm a ny p a rts of our coun

.
try th e Ame ric a n
f a rme r while libe ra l i n th e Use Of w a t er and fe rtilize r
, ,

a tt n ti on to th e physic a l c on diti on of
e
n
l owi g for ex a mple which keep s d ow n
, ,

es f resh su rfa ces to the a cti onof a i r a nd

a lso b ri ngs t ow a rd th e su rf a ce f resh p a rts of th e
s oil from wh ich f ood m a te ri a ls may be th e ore re a dily m
ext ra cted .

P OLLU TED AIR — C e rt a i n a i r p ol hi ti ons ma y c a use

chemic a l i nj u ry a nd dise a se in pl a ts I n th e a i r of n .

cities or in th e v icinity o f i ndust ri a l pl a nts i n wh ich

,

c oa l or ore c ont a i ni ng sulph u r is bu rned or roa sted ,

sulphu rou s v a p ors a re i nv a ri a bly p rese nt Th ese c a use .

dise a se by i nte rfe ri ng with th e norm a l fu ncti ons of

th e pl a nt especi a lly th o se c onnected with f ood m a nu
n
,

fa ctu re (ph ot osy th esis ) (Fig P olluti onof th e a ir

.

by s olid p a rticles wh eth e r of s oot or dust f rom roa ds

,

or such w orks a s ceme n t m a nu fa ctu ri ng pl a nts ma y ,

c a use such imp a i rment of the cl a ri ty of the a i r a s g re a tly

to reduce th e a m o u nt of ligh t re a chi n g the su rf a ce o f th e
e a rth or th e s olid p a rticles ma y f orm a n Op a que c o a ti ng
,

ove r th e le a ves a n d so p reve nt th e a bs orp ti ono f ligh t by

the chl orophyll c ont a ini ng cells
-
In eithe r eve nt th e
.
,

a m ou n t of f oo d m a nuf a ctu red ma y be gre a tly reduced .

M OI S T AN D D RY AI R — Th e h umidity of th e a i r ma y
.

s ometimes by sudde n ch a nge c a use dise a se not a lw a ys ,

rec og n ized a s to its origi n S udde n d ryness of th e a i r

” ,
.

such a s s ometimes a cc omp a nies h igh wind ma y bu rn ,

20 6 DI S E A S E S OF PLAN T S
f oli a ge especi a lly wh e nit is y ou ng Onthe othe r h a nd
,
.
,

sudde ni nc re a s e of humidity ma y c a use suhh a ch a nge i n

the cells a nd tissues th a t e rup ti ons k nown a s eme r “

g en c es o r i ntumesce nces m a y devel o p These eme r .

g en c es a re o utb re a ks th ro ugh the epide,

rmis o r b a rk of ,

s oft u nde rlying tissues i n wh ich the p ressu res rise ove r
ra p idl y by re a s onof a sudde ncheck to ev a p ora ti on .

FI G 52 . .
— T he Eff ect of S ulphur D i o xide Fum es from a smel ter
u po n th e f ores t two miles a w a y

H ATE AN D L COLD ,
T AN D S H A I GH DE — P rol onged ex
p osu re to u n fa v ora ble tempe ra tu res ma y p roduce dise a se ,

sh owi ng itself m a i nly i n ch a nges i n c ol or of the f oli a ge ,

th ough ext reme c a ses ma y exh ibit i nj u ry a s se ri ous a s

u ntimely le a f fa ll I nsu ffi cie nt or excessive ligh t will a ls o
-
.

sh ow itself i na b norm a l c ol or o f th e foli a ge ; a nd ve ry i h

su ffi ci ent ligh t a s is well know n will be f oll owed by f a ll o f
, ,

th e le a ves .

INTERNAL C A U SES

I t must be a dmitted a t le a st for th e p rese nt th a t th e

, ,

so c a lled i n
-
te rna l or physi ol ogic a l c a uses of dise a se a re
, ,

a ccepted a t a l l o nly bec a u se o th e r a nd exte rna l c a use h a s

no t bee nrec og nized Th us the m a l a dy k now na s little
.
“

pe a ch a nd a no the r c a lled s ou r sa p a re u nd oubtedly

, ,
 FA S C IATI O N 20 7

due to physi ol ogic a l distu rb a nce ; bu t wh a t ma y be the

re a s onf or th e physi ol ogic a l distu rb a n ce is not yet cle a r .

FA S CI AT I O N which c onstitutes a def ormity is due to

, ,

pl a nts or p l a nt p a rts growi ng so ra pidly a nd so cl ose

t ogethe r th a t th ey bec ome m ore or less me rged or
fuse d In c onseque nce a wide or b a nd like st a lk of
.
,
-

seve ra l or m a ny d a ndeli on he a d s g row n t ogeth e r c a rries

a t the top a nov a l i n ste a d of a ci rcul a r h e a d of bl oss oms .

S imil a rly th e C a liforni a p oppy p a rticul a rly i n a se a s on

, ,

of a bu n d a nt ra ina nd m ode ra te tempe ra tu res f orms such ,

fusi ons seve ra l fl owe rs being b orne a t th e tip of b a nd like

,
-

merged st a lks The sucke rs which sp ri ng up a rou nd th e

.

b odies Of t rees or S h rubs ma y be S imil a rly m e rged i nt o

b a nd like st ructu re s These occ a si ona l fusi ons or f a sci a
-
.

ti ons a re simil a r to th ose wh ich t a ke pl a ce i n succeedi ng

gene ra ti ons of pl a nts u nde r cultiv a ti on Ina ll su ch c a ses
.

such fusi ons or me rgi ngs seem to be due to f ailu re of the

pl a nt i n one or m ore of its c orrel a ti ons g rowth i n one ,

di recti onbec omi ng unduly ra p id or u nduly S l ow so th a t ,

the symmet ry of th e pl a nt i s not m a i nt a i ned a nd deformi

ties a rise which reve a l th e dise a sed c onditi on .

T H E L I V I N G C AU S E S O F D I S EA S E : I N FE CTI O N S — In
c onside ri ng th e livi ng c a uses o f dise a se or wh a t ma y be
,

c a lled i nfecti ons one sh ould disti nguish a t once betwee n

,

the mech a nic a l i nj u ry c a used by th e i nv a di ng org a nism

an d the distu rb a nce o f the norm a l physi ol o gic a l a c tivity
of the h o st . For ex a mple one ma y p u nctu re a le a f or
,

twig with a ni nst rume nt of the s a me size a s th e ovip osit or

of a
”
g a ll fly a nd p roduc e a woimd wh ich i n ordi na ry
'

ci rcumst a nces will p romptly h e a l ; but if a S imil a r

w ou nd i s p rodu ced by th e fly which dep osits a n egg a t
'

the b ott om of the pu nctu re it will not p rom

,

,
p tly h e a l .

A g a ll will g row a rou nd th e devel opi ng g rub Th e dis .

tincti on betwee n mech a nic a l i nj u ry a nd th e subseque nt

re a cti on s of the h ost to c ontinui ng stimul a ti onof th e i n
v a di ng org a nism ma y be illust ra ted by the foll owi ng
ex a mples
20 8 DI S E A S E S OF PLANT S

ANI MAL PA RASI TES

W ORM S —Th e c omm onest w orms to c a use dise a se i n

.

pl a nts a re th re a d w orms o r nem a t odes Th ese a tt a ck .

th e roots or oth e r p a rts of thei r ch a ra cte ristic h ost pl a nts ,

e g
. . p ot a t oes fl owe r bulbs etc e nte ri ng th e a dj oi ni ng
, , ,
.
,

tissues a nd be ing c ove red by e nl a rgeme nt s o fte na t ta i n

in g c on side ra ble size I nth ese a b norm
. a l g ro wth s wh ich ,

a re s ometimes c a lled g a lls th ese a n im a ls re a ch m a tu rity

,

a t th e expe n se of th e h ost pl a nt which feed s a nd S h elte rs

them Th ese w orm s not only rob th e h o st of food but
.
,

ma y a ls o p ois on it to a gre a te r or l ess exte nt by thei r

exc reti ons .

FI G 5 3. . Le a f Ga l l C a li f ornia
K ell ogg , Ameri ca nI ns ects .

F LI E S of v a ri ous s orts p a rticul a rly th ose c a lled g a ll

,

flies a re k now nto c a use th e f orma ti ono f le a f a nd stem

,

g a lls th e st ructu re a nd c omp ositi ono f wh i ch a s well a s

, ,

th ei r a ppe a ra nce a re of te n ve ry s t riki ng Th us o n a

,
.

ce rt a i n species o f o a k th e l a rge b a lls wh ich form

 PI N E GALL S 20 9

onth e twigs s ometim es c a lled oa k b a ll s i nste a d of oa k

g a ll s — a re c omm on These sph e ric a l e nl a rgem ents or
.

outgr owth s f orm a ro u n d grubs wh ich h a tch o u t f rom eggs

dep osited i n th e y ou ng te nde r b a rk o f growing twigs i n
the sp ri ng C a ll s i n a dditi on to c onsisti ng of a l a rge
.
,

a m ou n t of ext remely ligh t sp ongy tissue c ont a in a ls o a,

l a rge r p rop orti on of t a nni nth a noth e r p a rts of th e oa k ,

an d for th is re a s onm a tu re g a lls h a ve b ee nu sed for cen

f u ries a s a s ou rce of i n k .

Othe r sp ecies o f i nsect s c a u se th e f orm a ti on o f

n
g a lls o le a ves Of oa k (Fig 53 ) a nd othe r pl a nts Th e
. .

s a me oa k le a f ma y f orm two or m ore di ffere nt ki nds

-

of g a ll. I t is obvi ou s from th is fa ct th a t th e dise a sed

st ructu res c a lled ga ll s ta ke on a f orm wh ich is due not
t

on ly to th e h ost but to th e species of i nsect dep ositi ng th e

egg which is to h a tch i nt o th e p a rasitic g rub We h a ve
. .

here , the n a ve ry i nteresti ng dem onst ra ti onof th e sp ecific

,

a cti on of th e di ff e re n t a ge nts of dise a se a cting on th e

sa me orga nism th e h ost re a c ting i nch a ra cte ristic th ough
,

di ffe rent f a sh i onto th e a tt a cks of th e differe nt i ndividu a l

org a n isms One ma y o ften see a s m a ny a s h a lf a d ozen
.

diffe rent kinds of g a ll onth e s a me white oa k le a f due to ,

th e i nv a si onof this le a f by a s m a ny diffe rent species of

g a l l fli es
-
.

form a ti on of g a lls with thei r ch a ra cteristic a lly

diverse sh a pes th e tissues of v a ri ou s s orts c omp osing
,

these g a lls th e subst a nces wh ich a ct a s stimuli resulting

,

i n these o ve rgrowth s th e na tu r e o f th e a ss oci a ti on a ll

, ,

these a re subj ects of ext reme i nte rest .

P I N E GA LLS — In this group ma y be me nti oned a n

o the r peculi a r g a ll f ormed with o ut mech a n
,
ic a l inj u ry but ,

due to th e dep osit of its eggs by a fly a t the b a se o f a nd

betweenthe needles o f th e M onte rey pi ne a s these c ome
o u t in the sp rin g Th e grubs h a tch a s the ve ry y ou ng
.

le a ves begi n to g row ou t i nt o th e ligh t from the S he a th

wh ich encl oses th em Th e re is no w o u nd the re is me re
n
.
,

c ont a ct betwee g rub a nd tende r le a f b a se Th e le a ves -

.
21 0 DI S E A S E S OF PL AN T S
g ro w nly h a l f or a th i rd th ei r norm a l le ngth but a re
to o
,

two o r th ree times the n orm a l thick n ess a t the b a se These .

sh ort th ick le a ves give to M onte rey pi ne twigs a ve ry

cu ri ous a ppe a ra nce whe nth e re is ex te nsive g a lli ng One .

o f the e ffects o f th is i nth e c a se of M on te rey p ines is the

f a il u re o f th e b ra nch a nd o fte nof th e wh ole t ree to m a ke
, ,

the u su a l a m o unt o f w ood ; he nce i t is see nth a t th e eff ect

o f this dise a se is n o t limi ted to th e immedi a te p o in t of
In j u ry .

T H RI P S R ED S P IDER
, Pe a r a nd o the r f ruit t rees
.

a re subj ect to a tt a ck by sm a ll i n sects such a s th rips a nd ,

by red spide r etc wh ich c a u se t rue dise a se a nd no t

“
, .
,

simple mech a nic a l inj u ry for th e t rees which these i h ,

sect s a tt a ck yield less th a n th e norm a l a m ou nt of f ruit .

PLANT PARASITES

F U N GI Th e fu ngi f orm a ve ry l a rge divisi onof the

.

veget a ble ki ngd om They h a ve i nc omm onth e one qu a l

.

i ty o f depe nde nce they c a nno t m a ke thei r own f ood

, .

He nce th ey mu st p rovid e th emselves with f ood ei the r ,

by robbi ng o the r org a nisms or by livi ng on th e p rod ucts

or rem a i n s of o the r org a nisms Th e fu ngi a re d ivided .

a cc ordi n g to th ei r f ood h a bits i nt o sa p rophytic a nd

p a ra sitic th e f orme r living o nde a d m a te ri a l th e l a tte r on
, ,

livi ng pl a nt s or a nim a ls C e rt a in sm a lle r fu ngi a re the

.

immedi a te c a u se o f dise a se i n pl a nt s wh ile th e l a rge r ,

fu ngi k now n a s mu sh room s a nd t oa dst ools a re m a i nly

, ,

sa p rophytic Fu ngi c a u si ng pl a nt dise a se a re ne a rly a l l

.

o f mi n ute size Th ey a re c omm only sp oke no f a s rusts

.
,

mildews ro ts a nd smu ts
, ,
.

N ATUR E O F D I S EA S ES D U E T o F U N GI — Fu ngi b ri ng .

a b o u t th ei r ch a ra cte ristic distu rb a n ces o f th e norm a l life

o f thei r h o st pl a n t s by a bst ra cti ng f rom th e h o st th e food
wh ich they requi re a nd by th e mech a nic a l o r ch emic a l
,

i nj u ry wh ich th ey do i na dditi on Thus th ey ma y g row .

supe rfici a lly i na nd up onthe t issues o f th e h ost or they ,

 THE P O TAT O FUNGU S 1 1

ma y pe net ra te ve ry deeply In o th e r w ords th ey ma y

.
,

c a use def ormities by p u rely mech a nic a l me a ns pushi ng ,

the tissues of the h ost a p a rt a lth ough i na lm ost eve ry i n

,

st a nce i t is evide nt th a t the exc reti ons a nd sec reti ons o f

the fu ngi exe rt a p ois onou s i nfl ue nce on the a dj a ce nt or
evenove r rem ote p a rts of th e pl a nt This ma y be ill us
.

tra ted i n th e f oll owi ng specific dise a ses :

T H E P OTAT O F U N GU S T H E V I N E M I LDEw T H E O N I O N
'

, ,

M OLD a ll grow i nth e tissues of th ei r h ost p ush th e cells ,

of the le a f a p a r t a n d i nj u re or kill th e cells p resently ,

eme rging by b re a ki ng th e epide rmis a nd se ndi ng ou t

th rough the cra ck th ei r rep roductive b odies .

TH E R U S TS .
— The rusts of whe a t a nd m a ny o the r
pl a nts a re due to mi nute fu ngi the sp ores of which either
,

infect the g ra in s or a re sc a tte red f rom th e p a re nt pl a nt

an d subseque ntly i nfec t the y o u ng pl a nts by e nte ring the
le a f th rough the st om a t a i neithe r c a se a tt a cki ng m a inly
,

the tissue of the le a f a nd the reby i nte rfe ring with th e

p rocesses of f ood m a nuf a ctu re up onwh ich th e p rofit a ble
cultiv a ti onof th e pl a nt must depe nd A noth e r rust is one
.

which forms th e ced a r b a lls onth e j u mp e r or ced a r t rees

of the e a ste rnh a lf o f th e c on tinent A simil a r one is to
.

be f ou nd onthe le a ves of th e wild m a ll ow of the P a cific

Coa st .

TH E M I LDEw s . A noth e r g roup of fu ng ous dise a ses is

know na s the mildews Th e one with which pe ople gen
.

era lly a re m o st f a mili a r o ccu rs on th e f oli a ge of th e ro se .

Th e v a rieties of rose di ff e r ve ry c onside ra bly a m ong

themselves i n thei r suscep tibility to mildew Th ey a re .

a ll m ore or less susceptible h oweve r a cc ordi ng to th e

, ,

c onditi on of th e we a th e r It is p e rfectly p ossibl e

.

to c on t rol th is disea se by th e S impl e exp edie nt of

sp ra ying Th a t th e mildews c a use ve ry re a l i nj u ry is
.

p roved by observ a ti onof th e beh a vi or of d a hli a s These .

ma y be bl oomi ng a bu nd a ntly i n th e g a rde n; bu t whe n

exte nsively a tt a cked by mildew onth ei r le a ves they ce a se ,

to bl oom a n d r em a i n ste rile for th e rest o f th e se a s on .

21 2 DISEASES O F P LAN T S

I t is no t e a sy to see why or h ow th e i nflue nce o f the p a ra

site onbl oomi ng ca nbe so di rect a s it a ppe a rs to be .

S M UT S o f which c ornsmu t is p rob a bly th e m ost f a mil

,
-

i a r c a use a g re a t de a l o f d a ma ge a m o ng cultiv a ted pl a nts

, .

Th e bl a ckness or smutti ness is due to the form a ti on o f

bl a ck one ce lled rep roductive b odies (sp ores ) which a re
-

th rust ou t th rough the epide rmis of th e i nfec ted p a rt .

B A CTERI A L I N FE CTI O N o f pl a nts is now rec og nized to be

th e c a use of ce rt a inc ommonm a l a dies s ome o f which a re ,

o f g re a t ec on omic imp ort a n ce Thu s pe a r blight a b a c

fecti on o f th e i n
,
.

teri a l in ne r b a rk c a uses th e fl owe rs ,

to a b ort so th a t th e qu a n tity of f ruit f ormed ma y be ve ry

se ri ously red uced Ag a i n so c a lled wilt i n cucumbe rs
.
,
-
,

etc ma y c ompletel y dest roy a v a lu a ble c rop Th is dis

.
, .

e a se i s due to th e i nfecti onof th e w a te r c onducti ng tissues -

by a ce rt a i nspecies of b a cte ri a Al f a lfa a nd oth e r pl a nt s ,

devel op a t th e top o f th e root or b a se of th e stem e nl a rge

me nts k now n a s c row ng a lls Th ese a ls o a re due to
-
.

b a cte ri a l infecti ona n d bec a u se o f th e na tu re o f th e ti ssu e

,

g rowth s f oll owi ng such infecti ons c omp a ris ons h a ve bee n ,

m a d e betwee n th ese g rowth s a nd th ei r i nfecti ng c a use

with c a nce r i n th e h um a n b ody I t c a nno t be de nied .

th a t th e re a re ce rt a i nresembl a nces i nth e beh a vi or of th e

cells o f i nfected tissue i n c rowng a lls with th e growth s -

t a ki ng pl a ce i n c a nce r .

R OOT T U B ER CLE S Th e roo ts o f legumi nous pl a nts

.
,

such a s pe a s be a ns cl ove rs a nd a lf a lf a ge ne ra lly f orm

, , , ,

e nl a rgeme nt s k now n a s tube rcles or nodules Ex a mi na .

ti onof th ese sh ows th a t they a ls o a re th e se a t of b a c te ri a l

i nfecti on Th e i nfecti on t a kes pl a ce th rough the roo t
.

h a i rs ; th e i nfecti ng b a cte ri a t ra ve rsi ng th e roo t h a i r a nd -

pe net ra ti ng i nt o th e oute r p orti on of th e roo t p roduce

such a ni rrit a ti onth a t a no utg rowth i ns ome c a ses resem ,

bli ng a l a te ra l root is f ormed Th is c onsists m a i nly o f

,
.

th i nw a lled cells i nwh ich th e i nfecti ng b a cte ri a multiply

-

ve ry ra pidly Th e form a ti on o f th ese tube rcles is see n

.
,

the refore to be th e result of i nfecti on I t is a dep a rtu re

, .
 T R E AT M E NT OF DI S E A S E 21 3

f rom the norm a l a nd so fa r a s th e h a bits of th e pl a nt a re

,

c once rned it is a di se a se
,
.

In this i nst a n ce h oweve r the f orm a ti onof the tube r

, ,

cles resulti ng from b a c te ri a l i nfecti on i n th ese pl a nts is

of g re a t be n efit f or th e b a cte ri a in these tube rcles fix the
,

f ree nit rog en of th e a i r thus a ddi ng to th e soil ni trog

,

en ou s c omp o u n ds of th e u tm ost v a lue to the h ost pl a nt

an d to othe r species wild or cultiv a t ed ,
While the .

tube rcles onthe roots of legumi nous pl a nts a re a dise a se ,

they benefit a g re a t v a riety of o the r org a nisms a nd p re ,

su ma bl y di rectly or i ndi rectly th e legumi nous pl a nts

themselves Inthis respect we h a ve a ve ry u nu su a l con
.

diti on for dise a se is na tu ra lly c onside red to be qui te the

,

reve rse of a dv a n t a ge ous b oth to th e i ndividu a l a nd to the

c ommu nity .

T R EAT M E N T OF D I S EA S E I N P LA N TS Th e meth ods of .

de a li ng w ith dise a se i npl a nts f a ll i nt o th e s a me ge ne ra l

cl a sses a s th e meth ods empl oyed for h um a n bei ngs
na mely p reve nti on medic a l a nd su rgic a l t re a tme nt Thus
, ,
.

the i nst ruc ti ono f the p ublic i nth e me a ns of m a int a i ning

its own h e a lth a nd the h e a lth of th e living org a nisms
,

empl oyed by ma n for h is p u rp oses a re b o th receiving ,

the m ost skillful a tte nti on J ust a s th e c once rn with.

public he a lth h a s resulted i n p reve nti ng or g re a tly re

du cing th e occu rre nce of typh o id feve r so th e i ntellige n t ,

study of th e c onditi ons of pl a nt dise a ses e na bles us to

c ont rol them th rou gh p reve ntive me a su res much m ore
cert a i nly th a nby t re a tme nt wh e nth ey b re a k ou t .

T H E P R EV E N TI O N OF D I S EA S E Th e me a ns of p reven
.

ti on i na dditi onto th ose which ma y be sp oke nof a s gen

,

e ra l hygie ne c onsist specific a lly i n th e a pplic a ti on of

,

sp ra ys a nd dusts to individu a l pl a nts th us a ttempti ng ,

to p reve n t i nfecti on S u ch t re a tme nt is obvi ously

.

limited to th ose p a rts o f the pl a nt a b ove g rou nd

an d so p rotected by w a te r p roo fed epide rmis th a t the
-

p ois onous sp ra ys or dusts will not i nj u re th e tissue

o f th e h ost while th ey p o is on th e b a cte ri a
,
fu ngi or in , ,
21 4 DI S E A S E S OF PLAN T S
sects which migh t o th e rwise infect a nd i nfest th e l a rge r
pl a nt T h e a pplic a ti ono f d usts by bl owi ng onf oli a ge a nd
.
-

b a rk is desig ned to i nte rfe re m ech a nic a lly a s well a s chem

i ca l ly wi th th e i nv a si on o f h o st by p a ra site S p ra ys .

diff e r f rom dus ts m a i nly i nth e m a nne r o f a pplic a ti on for ,

th ough they a ls o a re bl ow n up on th e f oli a ge a nd b a rk ,

they a re suspe nsi ons or s olu ti ons i nw a te r a nd th e p o is on ,

o u s m a te ri a l left by ev a p ora ti on a ct s b oth mech a n ic a lly

an d ch emic a lly up on th e i nv a di ng org a nisms .

T H E M EDI CA L T R EAT M E N T O F D I S EA S E with th e obj ect

o f cu ri n g a ls o c onsists i n the a pplic a ti on o f sp ra ys a nd
d usts desig ned to offe r mech a nic a l i nc onve nie nce to p a ra
sites or to exe rt a p ois onous i nflue nce up onthem Thus .

B orde a ux mixtu re a sp ra y ma y se rve not only to p reve nt

, ,

i nfecti ons bu t ma y a ls o be used i nth e t re a tme nt of mil

dews or othe r fu ngi S p ra yed up on th e le a ves of roses
.
,

vines etc a tt a cks of mild ew ma y be cu red e nti rely by

,
.
,

this me a ns .

T H E S U R GI CA L T R EAT M E N T O F D I S EA S E — In c a se o f
i nv a si onbel o w th e su rf a ce of th e h ost su rgic a l t re a tment,

must c ommonly be res orted to i n pl a nts a nd i n th e ,

m ore se ri ous c a ses a mput a ti on c a nno t be a v oided .

I n orde r to p reve nt the sp re a d o f i n fecti on f rom

on e p a rt to a n o the r th e s a me p rec a u ti ons must be
,

empl oyed i n ste rilizi ng or dest royi ng a l l i nfected tissues

an d i n ste rilizi ng th e i nst rume nts u sed i nope ra ti on In .

f a ct with th e m odific a ti ons requi red by th e d iff e re nces i n

,

th e org a nisms th e s a me p rinciples of hygie ne c ont rol

,

th e t re a tme nt o f pl a n ts a s of a nim a ls I t is th us evide nt

”
.

th a t th e p ra ctice o f t ree su rge ry i nv olves gr e a t c a re i n

the t re a tme nt o f l ivi ng tissue a nd i nth e h a ndli ng o f i h
s tru men ts (ste riliz a ti on) Whe re c a re a nd ski ll a re l a ck
.

in g the resul t c a n e n t a il o n ly pecu ni a ry a nd o the r l o ss .

S U M M AR Y Th e c a uses a nd t re a tme nt o f d ise a se a re

.

esse nti a lly simil a r i npl a nts a nd a nim a ls L oss from u h .

f a v ora ble we a th e r or o the r e nvi ronme nt a l c onditi ons l a ck ,

o f p rope r co brdi n a ti o ni nthe vit a l p ro cesses o f th e org a n

 S UM M A R Y 21 5

ism a tt a ck a nd i nv a si on by o the r org a nisms ca n be

, ,

g re a tly reduced if not e nti rely p reve nted by ge ne ra l h y

g l e ne a nd by medic a me nts wh ich ma y a ls o be empl oyed
i nt re a ting individu a l c a ses o f dise a se
. Inext reme c a ses
su rge ry must be res orted to for rem ovi ng a dise a se d p a rt
which migh t if neglected i nv olve the wh ole . I nsects ,
fu ngi a nd b a cte ri a a re th e p ri ncip a l c a uses of pl a nt a s
, ,

well a s of a nim a l dise a se

,
.
 C HAPTER XX

DI S E A S E S DUE TO HA R M FUL O R GANI S M S

S O M E K I N DS OF ORGA N I S M S P R OD U CE P OI S O N S i n th e
na tu ral c ou rse of their met a b olic a ctivities Am ong th e .

m inu te org a nisms th a t le a d p a ra s i tic l ives on or wi thi n

th e b odies of l a rge r org a nisms a re nume rous v a rieties of
,

which th is is t rue These a re inj u ri ou s to th ei r h osts

. .

O the rs seem not to a ff ec t thei r h o sts on e wa y or a nothe r .

Th e c a s e p rese nted by org a nisms p a ra si tic on a nd p ro

du ci n g dise a s e i npl a nts f o rms th e subj ect m a tte r o f much
of th e l a st ch a p te r. He re th e discussi onwi ll be rest ricted
to i nj u ry p roduced i na n i ma ls by mi croOrga n i sms .

T H E W A Y S I N W H I CH M I CR OORGA N I S M S C AU S E H AR M
I N A N I M A L S a re p ri n cip a lly th ree They ma y g row a nd
.

multiply i n food ge ne ra ti ng p ois ons a nd so m a ki ng it

,

d a nge rou s Th ey ma y c a use excessive p ut ref a c ti on of

.

th e i ntesti na l c onte nts with resulti ng ill effects to the

,

b ody Or th ey ma y g row on or with i n th e b ody itself

.
,

p ois oni ng it di rectly .

F OOD P O I S O N I N G. F rom time immem ori a l i nd ividu a ls ,

an d s ometimes wh ole c ommu nities h a ve bee n st ricke n ,

wi th dise a se a s th e result o f e a ti ng sp oiled food Among .

st riki ng i nst a nces a re th e wid e sp re a d p ois oni ngs which

-

h a ve occu rred i n E u rope f rom time to time d ue to a ,

fu ng ou s g rowth onry e which c o nt a i ns a p ois onk nowna s

e rg o t Wh e n this ry e wa s gro u nd i nt o fl ou r a nd m a d e
.

i nt o b re a d wh ole c ommu nities th a t c onsumed th e b re a d

,

bec a me ill a nd m a ny d ied .

D ec omp o siti on o f p ro tei n f oods by pu t ref a c tive b a c

n
te ri a inthe p rese ce o f too little oxyge ns ometimes gives
21 6
 P R E V E NTI O N OF B O TULI S M 21 7

r ise to subst a nces c omm only c a lled p t om a i ns which a re

p ois onous to ma nif t a ke ni n f ood A t th e p rese nt time .

c onside ra ble p romi ne nce is being give nto a ki nd of f ood

p ois oni ng known a s bo tu li sm i n which c a nned f ood b e ,

c omes p ois onous th rough the growth withi n it of mic ro .

org a n isms wh o se w as te p roducts a re inj u ri ous to th e

h um a nb ody .

P R EVE N TI ON OF B OTU LI S M Th e org a nisms of b otu

.

li sm p roduce sp ores (see C h a p XX V ) which a re ve ry .

resist a n t to h e a t If a ny of th ese sp ores h a pp en to be

.

p rese nt up onspi na ch or st ri ng be a ns or o the r veget a bles ,

s ome of them ma y esc a pe dest ructi ondu ri ng th e c a nni ng

p rocess a nd ma y the ref ore be a ble to devel op i nt o a ctive
org a n isms a nd multiply e norm ously with inth e ca n B oth .

th e a ctive org a nisms a nd th e p ois ons which th ey p ro

duce a re dest royed a t the tempe ra tu re of b o ili ng w a ter ,

so th a t i f th e f oo d is he a ted t o th e b oili n g p omt sh ortly

~

before bei ng e a ten it c a nno t p ossibly p roduce b otulism

p ois oning .

I N TE S TI N A L P UTR E FA CTI O N I t h a s l on
. g bee n know n
th a t th e i ntestina l t ra c t i n th e high er a nim a ls h a rb ors
n
m a ny org a isms For the m o st p a r t these a re mi nu te
.
,

being eith e r pl a nt forms b a cte ri a or one celled a nim a ls

, ,
-
,

p rot oz oa I ndigestible a nd u ndigested c onstituents of th e

.

f ood a ff ord h ighly f a v ora ble nu t ritive m a te ri a l for

them so th ey th rive a nd multiply For th e m ost p a rt
, .

thei r p rese nce m a kes no di fl erence so fa r a s we k now , ,

to the org a nism wh ich se rves a s h o st ; bu t on o cc a si on

thei r growth is a cc omp a nied by the p roducti ono f p ois on
ou s m a te ri a l which ca nbe a n d o fte nis a bs orbed fro m the
,

i ntesti ne i nt o th e bl ood a nd so is dist ributed th rough out

,

the b ody If th e i ntestine f a ils to be ev a cu a ted a t su ffi

.

ci en tl y f reque nt i nte rv a ls the p roducti ona n

,
d a bs orp ti on
of these p o is on s g oes onu ni nte rrup tedly a nd c onstitutes
a defin ite c a use o f dise a se This c on diti on is c omm only
.

r efe rred to a s a ut o i n t oxic a ti on signifyi ng th a t the b ody

-
,

is p ois oning its elf I t h a s received less a tte nti onth a n it

.
21 8 DI S E A S E S FR O M HA R MF UL O R GAN I S M S

sh ould bec a use o f i ts c omm on o ccu rre nce a nd c omp a ra

ti v ely mi ld ill e ff ects bu t the re ca n be n
-
,
o d o ubt th a t

h um a n efficie ncy is d e fi nitely l owe red a s a resul t o f this :

ki nd of p ois oni ng .

T H E M ETA B O LI C A CT I VI T I ES O F P AR A S IT I C ORGA N I S M S .

Inp revi o us p a ra g ra ph s it h a s bee nsh ow nth a t org a n

isms occu rri ng i nf oods o r wi thi nth e a lime nta ry t ra ct ma y
p roduce p ois ons i nc onnecti onwith thei r met a b olic a cti vi
t ies I t is a ls o t rue th a t ce rt a i n ki nds o f org a nisms ma y
.

live onor withi nth e b ody a nd p rod uce p ois ons by wh ich
th e b ody ma y be h a rmed Th ese m a ke up the g roup o f .

so c a lled dise a se p roduci n

- -
g org a nisms I t sh ould be no ted .

th a t th e dist i ncti onbetwee ndise a se p roduci ng org a nisms -

an d such org a nisms a s do h a rm th ro ugh c a usi ng f ood

sp oil a ge or i n testi na l p u t refa cti on depe nds on wh ethe r
they a ffec t th e b ody di rectly or indi rectly Th e t rue .

dise a se p roduci ng o rg a nism is one which is p a ra sitic up on

-

o r wi th in th e b ody a n d libe ra tes its p ois ons di rectly i n

the cells or b ody fl uids Th e p rese nce o f these or .

g a n i sm s up on o r i n the b ody i nn umbe rs su fli ci ent to

c a use h a rm x consti tu tes wh a t is tech nic a lly k now n a s
a ni n fe c ti on .

T H E R E L AT I O N O F H OS T AN D P AR A S I T E is a m a l a dj ust
me nt of a ss oci a ti on I nfecti on f rom the pu rely bi ol ogi
.

ca l p o i n t o f view is a ni ncide nt i nth e st ruggle for exist

e nce P a ra sitic o rg a nisms must es ta blish themselves i n
.

ce rt a i nrel a ti ons with h ost org a nisms or they will p e rish .

The ide a l rel a ti o no f p a ra site wi th h ost is obvi ously one

i nwhich b o th p a ra site a nd h ost fl o u rish i e symbi osis a ,
. .
,

fo r i f the p a ra site dest roys th e h o st i t p rese ntly fi n ds

itself dep rived o f its s o u rce o f nou rishme nt a nd must fi nd
a n e w h o st or die
,
I nfe cti ons th a t i nj u re the h o st a re
.
,

f ro m th is p oi nt of view m a l a dj ustme nts in the a ss oci a

,

ti o n s o f org a nisms .

C A S S S O F D I S E AS E P R ODU C N G OR GA N S S
L E I -
I M — Th ree
cl a sses o f org a nisms a re rec ognized a s c a p a ble o f p rod u c
ing i nfecti on; th o se whi ch a re u n d o ub tedly pl a nts th ose ,
 O R GANI S M S THAT AR E P LANT S 21 9

wh ose rel a ti onships a re u nce rt a in a nd fi na ll y th ose which , ,

a re u n d o ubtedly a nim a ls .

1 Orga n
. i sms tha t a re u n dou b tedly p la nts I t sh ould .

be noted th a t these c ont a i nno chl orophyll a nd the re fore

ca nnot an m
uf a ctu re thei r own c omplex foods bu t a re ,

t rue p a ra sites de rivi ng thei r no u rishme nt from th e

,

b ody up on or withi n which they fi nd l odgme nt Th ey .

bel ong b ot a nic a lly eith e r to th e b a cte ri a or to the high e r

fu ngi .

B a c teri a a re sm a ll one celled pl a nts (Fig

,
-
S ome .

o f them a re sphe ric a l a n d s ome o f them a re st ra igh t ,

be nt or c o iled rods Thei r le ngth ra rely exce ed s th ree

,
.

or f ou r th ous a ndth s of a millimete r a nd thei r ,

FI G 54 . .
—T he B a cter i a Th a t C a us e T eta nus or Lo ckj a w .

di a mete rs a re usu a lly not fa r f rom one th ou s a ndth ;

Thei r cell st ructu re is exceedi ngly simple so simple i n
-
, ,

f a ct th a t it is d oubtful wh eth e r th ey h a ve a re a l nucleu s

such a s is to be fou nd i na lm ost a ll o the r cells .

The fu ngi c a use c omp a ra tively fewe r dise a ses in a ni

m a ls th a n th ey we re S h ow n i n the l a st ch a p te r to do i n
pl a nts Th ose which a re c a uses of a nim a l dise a ses a re
.

ve ry S imple fil a me nt ous fu ngi .

2 Orga n
. i sms whos e rela ti onshi p s a re u n certa i n .

T h e re is a g roup of dise a se p roduci ng org a nisms know n

-

a s sp i roche tes wh ich c a nno t a t the p rese nt time be defi

ni tely a ssigned to eithe r the pl a nt or th e a nim a l ki ngd om ,
2 20 D I S E A S E S F R OM HA R M FUL O R GANI S M S

a nd th ey a re the ref ore cl a s sed a s u nce rt a in The re a re .

a ls o a g re a t m a n y org a nisms which ca np roduce dise a se ,

bu t a re so mi nute a s not to be visible eve n u nde r the

h ighest p owe rs o f th e mi c rosc ope M ore ove r ona cc ou nt .
,

o f th ei r sm a ll size they p a ss th rough ve ry fi n e filte rs a ,

cir cumst a nce which c a uses them to be g rouped u nde r

th e h e a d of fil tera ble vi ru ses a vi rus bei ng a fluid wh ich
,

is c a p a ble of p roduci ng i nfecti on a nd a fil tera bl e vi rus ,

on e which ca np roduce in fecti ona fte r bei ng fil te red This .

obvi o usly me a n s th a t th e org a nisms a re so sm a ll th a t

they ca np a s s th ro ugh th e filte r .

3 O rga n
. i s ms tha t a re u ndou b tedly a ni ma ls — U n de r .

this h e a d c ome b oth p ro t oz oa a nd met a z oa Inl a te r p a ra .

g ra ph s specific i llust ra ti ons of b o th cl as ses will be give n .

B A CTER I A L D I S EA S E S eve ra l c omm on dise a ses a re of

.

b a cte ri a l origi n; a m ong th em ma y be me nti oned tub ercu

l osi s a nd typh oid .

Tu bercu losi s i s due to th e g rowth withi n th e b ody o f

tube rcle b a cilli (see Fig 55 ) wh ich c omm only m a ke the ir
.

FI G . 55 .
—B a cte r a i Th a t C a u se Tubercul osis .

wa y th ro ugh th e respi ra t ory p a ss a ges i nt o th e lu ngs ,

whe re th ey est a blish themselves a nd f orm little h a rd

m a sses k now na s tube rcles Tube rcul osis is not c onfi ned
.

to th e lu n gs but ma y a ls o a ff ect a lm ost a ny o the r p a rt o f

the b ody Wh a t is now k now n to be tube rcul osis of th e
.

lym p h a tic system wa s fo rme rly c a lled sc roful a or K i ng s ,

’
 T Y P H OID 22 1

Evil
”this
,
l a tte r na me bei ng d ue to th e supe rsti ti on
th a t if th e su ffe re r f ro m sc roful a c ould get th e king to
t ouch him he w ould be mi ra cul ously cu red The org a . n
isms oi tube rcul o sis a re ve ry likely to be p resent i n th e
a i r a s a p a rt of the dust si n
,
ce eve ry su ffe re r f rom p ul
,

mona ry t ube rc ul osis expels g re a t numbe rs eve ry time he

expect ora tes Inthe m ore c omm onf orms of tu be rcul o sis
.

the p ois onp roduced by the org a nisms is eithe r mild or is

i n sm a ll qu a ntities so th a t the a ctu a l p o is oni ng is
,

gra du a l a nd th e p a tient p a sses th rough a l ong p e ri od of

sl ow decli ne Inpu l m ona ry tube rcul osis the imp a i rme nt
.

of he a lth is due i n c on ,
side ra ble p a rt to the i nte rfe re nce
,

with respi ra ti on wh ich results from the dest ructi on of

lu ng tissu e by the g rowth of th e tube rcle b a cilli i n the
lu ngs Occ a si ona l i nfecti ons of tube rcul osis a re seen i n
.

which the org a nisms a re highly virule nt a nd th e dise a se ,

ru n s a ra pid c o u rse to a f a t a l te rmina ti on V e ry m a ny .

lives h a ve bee n s a ved si nce th e disc o ve ry th a t su nshi ne

an d a i r f ree fro m dust a nd a bu nd a nt a nd no u rishi ng f ood
t , i nc om
,

a re sufficie n bi na ti on to cu re ordina ry tube rcul o

,

sis p rovided e a rly di a g nosis is obt a i ned

, .

Typ h oi d is due to the typh oid b a cillus wh ich is p resent

i n v a st numbe rs i n th e i ntesti nes of eve ry su ffe re r from
the dise a se a nd s ometimes i n th e Intesti nes of well p er
,

s ons wh o a re know na s typh oid c a rrie rs ” If th e exc ret a

.

of these i n dividu a ls fi nd their wa y int o d ri nki ng w a ter or

i nt o milk th e org a nisms p resent i n the w a te r or milk
,

will rem a i na live for a c onside ra ble pe ri o d of time so th a t ,

pe rs ons d rinki ng th e c ont a mi na te d fluids i nt roduce the

org a n isms int o their i ntestina l t ra cts Fro m the re the . .

typh oid ge rms m a ke th eir wa y i nto s ome of th e cells of

the muc o u s memb ra ne lini ng the i ntestine a nd b y ,

growth a nd multiplic a ti on bec ome ve ry nume ro us .

Ea ch on e p rod uces s ome p o is on which st a ys withi n it

so l on g a s it is a live but whe n it dies the p ois on is set

,

free to be c onveyed a b out th e b ody i n the bl ood a nd

to do i n j u ry .
222 DI S E A S E S F R O M HA R MFUL O R GANI S M S

P U S F OR M I N G OR GA N I S M S
-
The re a re seve ra l kinds of .

b a cte ri a (Fig 56 ) th a t es ta blish them selves ord ina rily

.

j us t bel ow th e o ute r su rf a ce o f th e ski nor i nc orre sp o nd

in gly exp o s ed p o siti ons wi th i n th e m o uth or othe r p a s
s a ges o f th e b ody Th es e o rg a nisms a re c a lled pus
.

f orme rs bec a u se the de a d b odies o f th e org a nisms bec om e

,

mixed with wh ite c orp uscles a nd a ce rt a i n a m o u nt o f

fl uid to form pus Ordi na ry pi mpl es a nd b oils a re f a mili a r
.

resul ts o f th e est a blishme n t o f p us f o rmi ng orga nisms -

j ust u nde r th e ski n M ost of th e pus f ormi ng o rg a nisms

.
-

p roduce p ois ons wh ich p a ss from th e i nfec ted sp o t into

FI G . 56 .
-
One Type of P u S -Fo rmi ng O rga nism (S tre p to coccus ) .

the b ody fl uids a nd so a re dist ributed a b ou t th e b od y .

C e rt a i no f th ese org a nisms p roduce ve ry vi ole n t p ois ons

an d give rise to the c on diti onf orme rly know na s bl ood
p ois o ni ng ” .

P U S P O C K ET S
-
Th e c omm ons ort s o f p us forme rs p ro
.
-

d u ce a rel a tively mild p o is o na nd th e b ody i nwh ich they

a re p rese n t is refe rred to a s h a vi ng a l ow g ra de i nfecti on -
.

I t h a s l o ng bee nk no w nth a t i n ce rt a i nregi ons p a rti cu ,

l a rly a ro u n d th e roo ts o f th e teeth a nd i nth e t onsils l ow ,

g ra de i nfec ti o ns o f th is ty p e ma y bec ome est a blished a nd

co n ti nue fo r ye a rs Th e typic a l c o u rse o f eve nts is for
.

th e o rg a n isms to live a l o ng m o nth i n a nd m onth o u t , ,

mul tiplyi ng o nly a s f a st a s they die a nd so no t i ncre a s ing ,

in numbe rs p ro duc i ng only sm a ll a m o u nts of p us a n

,
d
 COM M O N COLD S 223

nt
o much p ois on such p u s a nd p ois on a s a re p roduced
,

bei ng a bs orbed i nt o th e b ody with o u t d oi ng obvi ous h a rm .

At i nte rv a ls p rob a bly bec a use of s o me distu rbi ng i nflu

,

e nce th a t h a s a ff ect ed the b ody wh a t i s k now n a s a

,

fla re— u p o ccu rs wh e nth e p us f orme rs multiply with g re a t

,
-

ra pidity a n d p roduce l a rge qu a ntities o f pus a nd p ois on ,

a l on g with defi nite b odily sympt oms of m ore or less

seve ri ty Afte r a l onge r or S h orte r p e ri od of a ctivity
.
,

the a ff ected t onsil or ulce ra ted t ooth quiets d ow n a nd

gives little t ro uble for a nothe r pe ri od of time Of rece nt .

ye a rs defi nite evidence h a s bee n secu red th a t these l ow

,

gra de i nfecti ons a re resp onsible for a cute rh eum a tism ,

f orme rly c a lled i nfl a mm a t ory rheum a tism which i n tu rn ,

o fte n results i n v a lvul a r dise a se of the he a rt E ve n .

th ough such a n a cute t ro uble ma y not devel op b odily ,

h a rm is d one a lth ough th e symp t oms o fte n c ome on so

,

g ra du a lly th a t th e p a tie nt is u nc onsci ou s th a t a nything

is the m a tte r with him o the r th a nth a t h e seems to feel
c onti nu ously less fit a nd to be upset m ore re a dily th a n
f orme rly Th e meth od of t re a ti ng l ow g ra de i nfecti ons
.
-

is to get ri d of th e m a sses of org a nisms wh e re th e pu s a nd

p ois on a re bei ng p roduced Wh e re the se a t of i nfecti on
.

is th e t onsil this is a cc omplished by its simple rem ov a l

,
.

It is not too much to sa y th a t th e p ra ctice of de ntist ry

h a s bee n rev oluti onized withi n the l a st dec a de by the
rec og n iti onof th e e a se with which i nfecti onca nest a blish
itself a b o u t th e roo ts of th e teeth a nd of the freque ncy
,

with which such i nfecti on i s reflected i n dist a nt p a rts of

'

the b ody a s a result of th e p o u ri ng ou t of p o is onfrom th e

a re a of i n fecti oni nt o the bl ood .

CO M M O N CO LD S — P rob a bly the m o st c omm on a n d ,

ce rt a inly the m ost f reque ntly repe a ted infecti ons from
which i ndividu a ls su ff e r a re th o se which c a use c omm on
c olds The re a re v a ri ous re a s ons why c olds a re p rev a le nt
. .

In the fi rst p l a ce it is now believed th a t th e re a re

,

h a lf a d oze nor m ore di ffe re nt org a nisms wh ich ma y c a use

c olds Fu rthe rm ore a ll of them fl oa t a b out freely inthe
.
,
22 4 DI S E A S E S FR OM HA R MFUL OR GANI S M S

d ust of the a ir a nd so a re b re a thed inm ore or less con

,

ti n o u sl y The resp ira t ory p a ss a ges a re the refore ra rely if

.

eve r e ntirely f ree f rom org a nisms which a re c a p a ble o f

c a usi ng c old s if th ey get a ch a nce to est a blish themselves
withi nthe b ody Th ey a re lyi ng i nw a it so to S pe a k so
.
, ,

th a t a ny rel a x a ti ono f th e resist a nce o f th e b ody e na bles

them to g a i na foo th old .

C AT C H I N G C OLD I t h a s bee n l ong k now n th a t a

.

sudde nch ill ing of th e b ody a s from sitti ng ina d ra ft is

, ,

a p t to resul t i n a c old ! uite rece

. n tly the expl a na ti on
,

o f this f a ct h a s bee nfu rn ished by th e disc ove ry th a t wh e n

th e o ute r su rfa ce is chilled ra pidly the re is c a used by a ,

mech a nism which c a nno t be desc ribed h e re a p a rti a l sh u t ,

off o f th e bl ood supply to the muc o us memb ra n

-
es o f th e
resp i ra t ory p a ss a ges This i nterfe re nce wi th the p rope r
.

nut riti on of th e cells reduces their resisti ng p owe r

sufficie ntly to a ll ow the org a nisms o f c olds to est a blish
th emselves a nd to devel op I t sh ould be u nde rst ood th a t
f

a ra pid ch illi n g ra the r th a nsimply th e f a c t o f bei ng c old

, ,

i s th e c onditi onwh ich i s likely to st a rt the i nfec ti oni nt o

a ctivity .H oweve r one sh ould be a r i n mi nd th a t ind i
,

v i du a l s ma y a ls o be especi a lly susceptible to i n fecti ona s

a result o f u n de r nut riti on or of exh a usti on a nd so ma y
-
,

devel op c olds with ou t the re h a ving bee n a ny speci a l ex

p o s u re imm edi a tely bef or e th e a tt a ck O bvi o usly
. the ,

best p reve ntive o f c olds w ould be to h a ve th e respi ra t ory

p a ss a ges free f rom th e org a nisms which c a use th em ; since
th is is no t p ossible in p ra ctice th e a lte rna tives a re th e
,

m a i nte na nce o f th e b ody i n the best p ossible h e a l th a nd

th e a v oid a nce of such exp osu re a s is likely to c a use ra pid
chill i ng o f th e b ody su rf a ce The regul a r t a king o f c old
.

b a th s wh e re g ood re a cti ons a re secu red is a n excellent

, ,

me a ns o f keepi ng h igh the gene ra l resist a nce a g a inst

c olds .

D I S EA S E D U E T o H IGH ER F U N GI is c omp a ra tively ra re .

The re is i n p a rts of C a li fo rni a wh a t i s know n a s S a n

J oa q ui nD ise a se which resembles tube rc ul osis of the lu ngs
,
 DI S E A S E DUE TO S P I R OC H E T E S 22 5

i nm a ny its symp toms but is due to the p resence with

of ,

i nth e lu ngs o f m old like fu ngi i nste a d o f b a cte ri a

-
.

D I S EA S E DU E T o S P I ROCH ETES The spi ro chete is a n .

-

org a n ism sp i ra l i n f orm One ca n be im a gi ned on a .

l a rge sc a le by supp osing a bit of st ri ng to be w ou nd rou nd

th e finge r a nd th e fi nge r withd ra w n with o u t distu rbi ng
th e st ring The c omm onest spi roch ete dise a se i s syphili s
.

(Fig . The p oi son of this dise a se I s ve ry inj u ri ou s

to l ivi n g tissues so th a t b a d s ores a re e a rly c onsequ ences
,

of the in fecti on a nd wide sp re a d dest ructi on p a rti cu

,
-
,

—A
)

FI G 57
. . C o mm on Ki nd of S pi ro ch ete .

l a rl y of p a rt of the spi na l c ord a l a te r m a nifest a ti on

-

, .

Alth ough cu ra ble syphilis is ex t remely stubb orn so th a t

, ,
'

on ly skilful a nd th orough t re a tme nt e ra dic a te s it com !

p l ete l y whe n it on ce bec o mes est a blish ed withi nth e b od y .

D I S E A S E S D U E T o T H E F I LT E R A B L E V I R U S E S M a ny of .

the c omm onest dise a ses such a s me a sles a nd sc a rlet feve r

, ,

a re due to org a n isms too minute to be see n u nde r th e

mic rosc ope A ch a ra cteristic of m a ny of them is th a t
.

they a re di rectly c ont a gi ous th a t is to sa y simple c ont a ct, ,

with the p a tient often suffices to c a use one to bec ome

infected .

PR OTOZ OA N D I S EA S ES — M a l a ri a is a g ood ex a mple of

.

dise a se due to a one celled org a nism (Fig 58) which i s

-
.

knownto be a na nim a l Th is Org a nism est a blishe s itself

.

inthe red c orpuscles a n d by i nterfe ri ng with their fu n c

22 6 DI S E A S E S F R O M HA R M FUL OR GAN I S M S

ti oning hi nde rs th e norm a l t ra nsp ort a ti on o f oxyge nby

the bl ood M ore ove r th e p ois ons p rod uced by the
.
,

m a l a ri a l p a ra site b ring a b o ut defi ni te sympt oms of which

chills a nd feve r (Fig 59) a re th e m ost f a mili a r Th e
. .

d rug q u i ni ne f o u nd i n th e b a rk o f the ci nch ona t ree

, ,

dest roys m a l a ri a l org a nisms so th a t if qui ni ne ca n be ,

i nt roduced a t the tim e whe nth e org a nisms a re a ctive i n

th e bl ood th e b ody ca n be f reed f rom i nfecti on
,
.

FI G 58
.
— T h e P ro
. FI G 59
. . S u ccessiv e S ta ges i n th e Life
to z oa n Th a t C a us es His to ry o f th e M a l a ri a O rg a nism T he pe ri o d
.

M a l a ri a. o f h ig h f e ver is duri ng the tra nsi ti on fro m

s ta ge 3 to s ta g e 4 .

The re a re seve ra l species o f a m oeb a (Fig 60 ) th a t live .

i n th e h um a n a lime nt a ry t ra ct a nd by me a n s o f th e
p ois ons wh ich th ey p roduce b ri ng a b o ut a f orm of dysen
tery k n ow n a s t ropic a l o r a m oebic dyse nte ry This .

FI G 60 . .
—An Am b a from
oe th e Hu m a n D ige s tiv e Tra ct .

dise a se occu rs ch iefly i n th e t ropi cs a nd i s fou nd in

t empe ra te regi on s only i n pe rs ons wh o h a ve c ome f rom
t ropic a l clim a tes
—
.

D I S E AS E D U E T o M ETA Z OA Th e re a re v a ri ous highly

.

o rg a n ized p a ra sites wh ich ma y ca use dise as e Among

n
.

th em is th e h ookw orm a no rga ism wh ich h a s a tt ra cted

,
 VI R UL E N CE 227

much a tte nti onof rece nt ye a rs bec a use o f th e exte nt to

which it h a s l owe red th e efficie ncy o f th e p opul a ti on i n
th ose regi ons wh e re it is a bu nd a nt The org a nism is .

p rese nt i nl a rge numbe rs i nthe intesti na l t ra ct o f a ffected

pe rs ons a nd is expelled with thei r ex cret a Wh e re s a ni
,
.

t a ry p rec a uti ons a re not obse rved a nd th e exc ret a a re

a ll owed to bec ome mixed with the s o il th ese org a nisms ,

a re p rese n t i n a bu nd a nce a t th e su rf a ce of the grou nd .

Wh e n one w a lks with b a re feet ove r such a n i nfected

sp o t s ome o f th em a re likely to pe net ra te th e S ki no f th e
,

s ole whe nce they w ork th ei r wa y upw a rd a m ong the b ody

,

tissues a nd ultim a tely to th e a lime nt a ry t ra ct f rom which

,

they will be expelled i nc ou rse o f time to se rve a s p ossible ,

s ou rces of fu rth e r i nfe cti on F rom th e a b ove desc rip ti on

.

it is evide nt th a t th e h ookw orm ca n fl o u rish only i n re

gi ons whe re th e simplest s a nit a ry p rec a u ti ons a re neg
l ected a nd wh e re th e p opul a ti onis eith e r too p oor o r too
,

sl oth ful to we a r sh oes I t sh o uld be no ted h oweve r th a t

.
, ,

the chief m a nifest a ti onof h ookw orm i nfecti onis m a rked

b odil y l a ssitude so th a t th e l a ck of s a nit a ry p re ca uti ons
,

is i n p a rt due to th e l owe ri ng of vit a l ity c onseque nt on

'

the dise a se F ortu na tely for th e welf a re of th ose dis

.

tri cts i n cludi ng p a rts of the U nited S t a tes a nd l a rge a re a s

,

i nC ent ra l a nd S o uth Ame ric a wh e re h oo kw orm i nfest a ,

ti onoccu rs quite S imple t re a tme nt with th e d rug thym ol

, ,

c ombi ned with S imple s a nit a ry regul a ti on su fli ces to

.

st a mp ou t the p a ra site p romp tly a nd eff ecti v ely .

V I R U LE N CE A f a c t th a t must be t a ke n i nt o a cc ou nt
.
-

i n a ny a ttemp t to u nde rst a nd th e co u rse a nd S p re a d o f

i nfe cti o us dise a se is th a t its p roduci ng org a nisms ma y dif
fer g re a tly f rom time to time i nwh a t is c a lled vi ru len ce .

Unde r ce rt a i nci rcumst a nces the org a nisms a lth ough p res ,

en ,
t ma y p roduce on ly few c a ses o f dise a se a nd th ese
mild U nder ce rt a in o the r ci rcumst a nces ne a rly every
.

on e exp osed ma y be i n fected a nd th e i ndividu a l ca s es

ma y be ve ry se ri ous Th ese di ff e re nces a re u nd oubtedly
-
.

due to a lte ra ti ons in th e met a b olism oi the dise a se p ro -

228 DI S E A S E S FR O M HA R M FUL O R GAN I S M S

du cing or ag nisms themselves a nd whil e no t much i s,

k now n a b ou t th e c ondi ti o ns wh ich g ove rn these di ff e r

en ces i n vi rule n ce th e d iff e re nces th emselves a re o f u t
,

m ost p ra ctic a l imp orta nce for a pe ri od o f excepti ona l

,

vir ule nce is a lm ost su re to me a na nepidemic of the p a r

ti cul a r dise a se .

D EFE N S ES OF ORGA N I S M S E ve ry infecti o us dise a se is

.

a st rife betwee n th e i n divid u a l i nfected a nd th e dise a se

p roduc ing org a nism s by which th e i nfecti oni s est a blish ed .

S i nce no t a l l pe ople th a t a re exp osed to i nfecti onta ke it ,

th e re must be a me a ns of h oldi ng th e i nfecti on a t b a y .

Fu rth e rm ore if th e i nv a di ng org a nisms we re a ll owed to

,

d evel op a nd m ultiply with o ut rest ra int th ey w ould p ro

duce p ois ons i n sufficie nt qu a ntity to dest ro y thei r h o st -
.

Th e f a c t th a t a nim a ls rec ove r f ro m i nfecti on is p roof

th a t they h a ve s ome me a ns of c omb a tti ng th e org a nisms
of dise a se . Wh a t th ese a re is to be le a rned p rese ntly .

P R EVE N TI O N O F I N FE CT I O N D ise a se p roduci ng or

.
-

g a n i sm s a re ve ry widely sc a tte red a nd a re

q uite li kely to

be t a ke ni nt o the b ody with th e a i r th a t is b re a thed or

with the f ood or w a te r th a t e nte rs th e m o uth The p re .

v en ti ono f i n fecti ond epe nds i nth e fii s t pl a ce onkeeping

o rg a n isms which e nte r th e resp ira t ory or a lime nt a ry
t ra cts f rom getting i na m ong the b ody tissues whe re con
di ti ons for g rowth a re f a v o ra ble Th is is a cc omplish ed
.

by p rese nting u nb roke n su rf a ces which th e org a nisms

a re n o t a ble to pe n et ra te Th e re is no d o ub t th a t a ve ry
.

l a rge p a rt o f ou r f reed om f ro m infecti ondepe nds onint a c t

su rf a ces T o th e org a nisms o f d ise a se no t only the oute r
.
,

su rf a ce of the b ody i e th e ski n bu t a ls o th e muc ous

,
. .
, ,

li ning of th e respi ra t ory a nd digestive t ra cts a re p o ssible ,

a re a s th ro ugh which e n t ra nce i nt o th e b ody ma y be

g a ined An o rg a nism wh ich is b re a th ed in wi th th e a ir
.

is likely to l odge up o nth e muc ous memb ra ne of the lu ngs

o r b ro n ch i a l tubes a nd if it pe net ra tes th e memb ra ne
,

i t fin ds itself i nth e w a rm a n d m o ist b ody subst a nce whe re

g rowth ca np ro ceed re a dily S imil a rly org a nisms t a ke n
.
,
 O V ER CO M I NG I N FE C TI O N 229

i nwith f ood h a ve only th e lining of the a lime nt a ry t ra ct

betweenthem a nd the b ody subst a nce Ab ra si ons or i n .

fla mma ti ons of a ny of th ese su rfa ces ma y a ff ord j ust the

needed p ort a l of e nt ry for virule nt dise a se p roduci ng or -

n i sm s O n e o f the re a s ons why c o mm o n c o lds a re so

g a .

f requently f oll owed by m ore se ri ous i nfecti ons is th a t th e

i rrit a ti ono f th e muc ous memb ra nes b rough t a b out by the
c old m a kes the memb ra nes e a sily pe net ra ble by oth e r
m ore d a nge rous org a nisms .

T H E OV ER CO M I N G OF I N F E CTI O N
_
Afte r th e org a n
.
-

ism h a s p a ssed the b a rrie r of the b ody su rfa ce a nd est a b

l i sh ed itself i n a f a v ora ble l o c a ti on a m ong the tissues ,

the necess a ry c onditi ons for ra pid g rowth a nd multiplic a

ti onwith the a cc omp a nyi ng devel opme nt of dise a se w ould
seem to be p rese nt The re exist h oweve r ce rt a in
.
, ,

mech a nisms i nthe b ody wh ich opp o se th e a ctive gr owth

of the i n v a di ng mic robes I f th ese mech a nisms a re wh olly
.

successful th e org a nism s a re dest royed before a ny ch a nce

to multiply is a ff orded a nd so th e i nfecti on d oes n
,
ot

eve n get st a rted If th e resist a nce of th e b ody is less

.

p rompt a nd yet re a s ona bly e ffective th e org a nisms multi

, ,

ply for a time p roduci ng a nd p o u ri ng ou t th ei r p ois ons

, ,

an d so th e sympt oms of a cute dise a se a ppe a r P rese ntly .

,

h oweve r the b ody ra llies its mech a nisms of defe nse c ou n

, ,

tera cts the p o is on s a nd dest roys the org a nisms a nd thu s

, ,

r ec o ve rs c ompletely In still othe r c a ses eithe r on a o

.
,

c ou nt of excepti ona l vi rule nce Onth e p a rt o f th e inv a di ng

org a n isms or a n inefficient defe nsive mech a nism on th e
,

p a rt of th e i nfected individu a l the devel opme nt of th e

,

org a n isms a nd th e p roducti onof p ois ons by th em bec ome

m a rked enough to dest roy th e life of th e victim .

The b ody h a s two disti nct me a ns which it ra llies sep a

ra tely or i nco O era ti oni nthe e ffort to o ve rc ome i n
p fecti on .

Th e fi rst of th ese is ce rt a i n of th e wh ite bl ood c orpus -

cles to which on a cc ou nt of this ve ry a cti on th e na me

, , ,

p ha g yoc tes (sc a ve n ge r cells ) h a s bee n a pplied Th e .

sec ond is a meth od by which th e p ois ons or the org a n ,

23 0 DI S E A S E S F R OM HA R MFUL OR GANI S M S

isms wh ich p roduce th em or b o th a re a tt a cked by chemi

, ,

ca l me a n ,
s a nd to this is give n th e na me of the i mmu n
i ty rea cti on
-
.

T H E A CT I O N O F P H AGOCY T Es — P rob a bly not a ll of

.

th e white bl o od c orpuscles exe rt the so c a lled ph a g o cytic

- -

a cti on. I n th o se th a t do the a c ti on c o nsists i nw a n

,
de r
in g a b o ut by a m oeb o id m o ti on (see C h a p X I I I ) th e .
,

di recti on of the m o ti on p rob a bly bei ng i nflue nced by

ch emic a l subst a nces give noff by th e i nv a di ng org a nisms .

Whe neve r c ont a ct is m a de with a mi cro Orga ni sm a

b a cillus o f typh oid for inst a nce

,
the ph a go cyte p resses
cl osely a g a i nst th e b a cillus a nd by m o veme nts of i ts p ro
tOp l a sm e n gulfs it c ompletely Once withi n th e b ody o f
.

the ph a g ocyte th e o rg a nism is digested a nd dest royed .

It is a ni nte resti ng fa ct th a t if ph a g ocytes a re i nth e p res

e nce o f l a rge numbe rs o f a p a rticul a r ki nd o f org a nism ,

such a s th e typh o id b a cillus the numbe r o f orga nisms a

,

si ngle ph a g ocyte will e ngulf is quite defi nite so defi nite i n ,

fa ct th a t it ca nbe u sed a s a ni ndex o f th e efficie ncy o f th e

b ody i n defe ndi ng i tself a g a i nst typh oid si nce evide ntly ,

the m ore orga ni smS e a ch ph a g ocyte will e ngulf th e m ore

re a dily will th e b ody ri d itself o f a ll the org a n ism s wh ich
h a ve inv a ded it A fu rth e r cu ri o us f a ct a b o u t th e a c ti on
.

o f ph a g o cytes a n
,
d a f a ct o f ve ry g re a t imp ort a nce i nth e
wh ole p ro cess o f o ve rc oming i nfecti on is th a t if th e ,

typh o id b a cillus to u se th e s a me illust ra ti on bec omes

, ,

est a blished i n th e b ody a nd begi ns to multiply so th a t

p ois ons begi n to be p ou red ou t a nd the sympt oms of
dise a se begi n to S h o w th emselves a chemic a l re a c ti on
,

g oes on i n th e cells o f the b ody p roduci ng a m a te ri a l

,

which a ffec ts th e b a cte ri a i nsu ch a wa y a s to c a use e a ch

ph a g ocyte to e ngul f m a ny m ore o f th em th a n f orme rly .

Th e det a ils o f this a cti ona re not a t a ll cle a r but o f the ,

f a ct a nd i ts v a lue a s a me a ns o f o ve rc omi ng dise a se

the re ca n be n o d o ub t The p ro tective i nocul a ti on
.

a g a inst typh o id whi ch red u ced th e occu rrence o f th a t

,

di se a se to such a n exte nt th a t f rom being a f rightful

 T HE I M M UNIT Y R E A C TI O N 23 1

.
m
dest roye r o f life a t the ti e of th e S p a nish Wa r th e re ,

wa s only a h a n dful of well a uthe ntic a ted c a ses i n th e

-

Ame ric a n a rmy du ri ng the G re a t Wa r is a p ra ctic a l ,

a pplic a ti on o f this re a cti on by which the p o we r of th e

ph a g ocytes to e ngulf a nd des troy ty p h oid org a nisms is

gre a tly i nc re a sed
—
.

T H E I M M U N I T Y R EA CT O NI M a ny dise a ses a m ong

.
,

them sm a llp ox p neum oni a a nd i n f a ct th e m a j ority of

, ,

th e m ore p rev a le nt i nfecti ons stimul a te th e b ody i n

, ,

which the dise a se h a s g a i ned a f ooth old to p roduce sub ,

st a nces k now n a s a nti — bodi es which a re ca p a ble o f (a )

,

dest royi ng foreig n org a nisms or (b ) of neut ra lizi ng th e

.

p ois ons which they p ro du ce Th is dise a se resisti ng p rop

.
-

er ty of th e a n ti b odies is me rely a speci a l a nd ve ry

-

useful a pplic a ti on of a gene ra l p owe r th a t livi ng p ro

top l a sm h a s o f getti ng ri d o f f oreig n p ro tei ns I t will .

be rec a lled th a t f oreig n p ro tei ns a s such ra rely c ome

n
, ,

i nt o c o t a c t with the livi ng cells of the high e r a nim a ls ,

bec a use a ll p ro tei ns th a t a re t a ke ni n with th e f ood a re

digested i nt o th ei r c onstitue nt a mi no a cids before bei ng -

a bs orbed i n to the b ody fluids I t is p ossible h oweve r to

.
, ,

i nj ec t p ro tei ns di re ctly i nt o a vein or they ma y obt a in ,

a ccess th ro ugh i n fecti on th e b odies of dise a se p roduci ng

,
-

Org a n isms a nd th e p ois ons th ey p roduce b o th bei ng

l a rgely p ro tei ni n ch a ra cter .

It is ch a ra cte ristic o f the cells of th e highly org a nized

an im a ls th a t wh e n f oreig n p ro tei n c omes int o c ont a ct
with them they re a ct by p roducing a nti b odies Th e -
.

effect of the a nti b odies is to p recipit a te th e p rotei n or

-

disp ose of it i n s ome o the r wa y An imp ort a nt fe a tu re .

of the a n ti b odies is th a t the re is a p a rticul a r s ort for

-

e a ch ki nd of p ro tei n a nd wh en th e cells o f th e b ody

,

re a ct to th e p rese n ce o f a f oreign p ro tein th e a nti b ody ,

-

they p roduce is th e p a rticul a r one by which th a t p rotein

is disp osed of .

D I P H T H ER I A . Th e re a re a few ki nds of dise as e p ro -

duci n g orga n i sms , of which the di p hthe ri a b a cillus (Fig .

23 2 D I S E A S E S F R OM HA R M FUL O R GAN I S M S

61 ) is th e best know n ex a mple wh ich do not pe net ra te

-
,

i nt o the b ody a n d bec ome dist ributed th rough it but lo ,

c a te I nsome p a r ticul a r regi on i nthe ca se of d iph the ri a , ,

the regi on bei ng the muc o us li ning of th e th roa t The

“

o rg a n isms g row th e re a nd multiply a ctively a nd p ou r ou t

their p ois ons These a re a bs orbed i nt o the b ody di stri
.
,

buted th rough it by th e c ircul a ti on a nd p ois on th e ,

in d ividu a l cells Th e p ois on wh ich is know n a s th e

.
,

toxi n o f th e dise a se stim ul a tes th e cells to p rod uce th e

,

an ti b odies wh ich i n this c a se c onstitute a nti toxi n

- -
.

FI G 6 1
. .
—T he B a cte ri a Th a t C a u s e D iph th eri a .

Wh e the r th e p a tie nt su rvives or succumbs d epe nds on

wh ethe r th e a nti t ox i n is or is not p roduced in su fli ci ent
-

qu a ntities to neut ra li ze th e t oxin .

A RT I F I C I A L A N T T OX N
I I — I t is p ossible to g ro w
-
.

d iph th e ri a org a nisms i n test tubes i n a suit a ble cul tu re

medium a nd the reby p roduce l a rge qu a ntities of t oxi n
, .

This t o xi n is the n i nj ected int o h e a lthy h orses in qu a n

tities no t su fli ci ent to p ois on th em se ri ously bu t a de ,

qu a te to stimul a te th ei r cells to th e p roducti on of a nti

t oxi n Afte r a suit a ble i nte rv a l h a s el a psed d u ring
.
,

which the cells o f th e h orses b odies a re m a nuf a ctu ri ng ’

an ti t oxin bl ood wh ich will o f c ou rse c ont a i ni t is wi th

-
, , ,

d ra w n th e c orpus cles rem oved a nd th e cle a r liquid pu t

, ,

up i n suit a ble f orm f or i nt roducti on i nt o th e bo dies o f

diph the ri a p a tie nts Th us th e p ois on which is being
.
 I MM UN IT Y 23 3

p roduced withi nthe b ody of the p a tie nt ca nbe neut ra l

i z ed di rectly with o u t th e necessity o f w a iti ng f or th e
cells to re a ct a nd p roduce a nti t oxi n -
.

I M M U N IT Y M ost dise a se p roduci ng org a nisms re

.
- -

t a in withi n th ei r own b odies the p ois ons which they

p roduce a nd only give them up a s th ey a re dest royed .

Inorde r to c omb a t these successfully th e b o dy cells must

do m ore th a n p roduce the a nti t oxi n for the neut ra liz a -

ti on of p o is on Th ey must p roduce i n a dditi on a nti

.

b odies which a tt a ck a nd dest roy the org a nisms th em

selves Th is a ctu a lly h a ppe ns so th a t one wh o is ill
.
,

from such a dise a se a s p neum oni a is p roduci ng withi n

his tissue a nti b odies which a re a ctively dest ructive to
-

the p neum oni a org a nisms a s well a s a ble to neut ra lize

the p ois ons wh ich th e org a nisms p ro duce .

S M ALLP ox A N D VA C C N AT O N
I I — An a cc o u nt o f i m .

mu nity w ould be i nc omplete with ou t b rief refe re nce to

sm a l lp ox T wo h u nd red ye a rs a go this l oa th s ome dise a se
.

wa s so p rev a le nt th rough o u t the civilized w orld th a t

a dults with u np o ck m a rk ed f a c e s we re obj ect s of p ublic
- -

a tte n ti on A ce rt a i n fa m ou s be a uty La dy M a ry
.
,

M ont a gu went SO fa r a s to a d op t a p ra ctice well k now n

,

i nC onst a nti nople o f h a vi ng he rself i nfected with sm a ll

,

p o,
x i t h a vi ng been obse rved th a t pe rs ons wh o su rvived
delibe ra te i nfecti ons esc a ped with less p ock m a rki ng -

th a n th e ge ne ra l ru n of victims of the dise a se Wh e re .

p ock m a rked f a ces we re so p rev a le nt th e cu rre nt belief

-
,

th a t milkm a ids we re m ore likely th a n o the r p e rs ons to

h a ve sm oo th skins wa s b o u nd to a tt ra ct c onside ra ble
a tte n ti on J e nne r a y ou ng physici a n wh o l ooked int o
.
,

the m a tte r sh ortly before the end of th e E igh te e nth

Centu ry bec a me c onvi nced th a t the re wa s s ome r e a l re
,

l a ti on betwee n th e s ores wh ich milkm a ids freque ntly

g t onth e p a lms o f thei r h a n
o d s from th e udde rs of c ows
a fflicted with cowp o x a n d thei r rel a tive immu nity to
sm a llp ox F oll owi ng up this ide a h e p rev a iled up onth e
.
,

p a re nts of a y ou ng b oy to a ll ow thei r son to h a ve ma

23 4 DI S E A S E S F R OM HA R M FUL OR G ANI S M S

teri a l f rom such a cow p ressed i nt o a n a b ra si on on h is

S kin . Th is wa s th e beginni ng o f v a cci na ti on Al th ough .

bitte r h o stility wa s e nc o u nte red f rom a ll sides the p ra o ,

tice a lm ost immedi a tely vi ndi ca ted itsel f a nd J e nne r d u r , ,

i ng th e l a te r ye a rs o f h is life wa s h a iled a s a gre a t

,

be nef a ct or o f h is ra ce Th e p ossibility of v a cci na ti on

.

rests up on th e f a ct th a t th e org a n isms o f c owp o x a nd

vi rule nt sm a llp o x a re so ne a rly ide ntic a l i nth ei r p ro tei n
c omp ositi on th a t the immu ne b odies wh ich th e tissues
p rod uce for one se rve c ompletely to a tt a ck a nd dest roy
th e othe r Th is cl o se p ro tei n rel a ti onship o f a seve re
.

with a m ild i nfecti on h a s m a de i t p o ssible to m a ke a

ve ry s a tisf a ct ory begi nni ng t ow a rd riddi ng the civilized
w orld of one o f its m o st f a t a l a nd l oa th s ome dise a ses .

D E F E N S E S O F CO M M U N I T I E S — As civiliz a ti on a d
v a nces a nd men te nd m ore a nd m ore to cluste r i n com
mu ni ti es c onditi ons bec om e m ore a nd m ore f a v ora ble
,

for the org a nisms th a t p ro duce dise a se i n h um a nbei n gs .

These org a nisms a re p a ra sitic with in the b odies o f men

an d ce rt a i nki n d s of high e r a nim a ls so th a t thei r ch a nce
,

of existe n ce is g re a te r th e m ore pe rs ons a re withi nre a ch .

Fu rthe rm ore to m a int a i n th e p u rity of w a te r milk

, , ,

an d f ood supplies bec omes i nc re a singly difficult a s th e

p op ul a ti onbec omes de nse r Th e c o nt a mi na ti ono f th ese
.

subst a nces with org a nisms th a t p roduce dise a se is for

the m ost p a rt c ont a mi na ti on with hum a n exc ret a T o
.
.

p reve nt sew a ge f rom getti ng i nt o d ri nki ng w a te r a nd i nt o

the w a te r with wh ich milk c a ns a re w a shed seems like
-

a simple u n de rt a ki ng bu t i np ra ctice it p rese nts a g re a t

,

m a ny difficulties d ue eith e r to i ndi ffe re nce on the p a rt

,

o f th e educ a ted public o r to ig n o ra nce on the p a rt o f

o th e rs . C ommu ni ties ca nno t s a fely rely wh olly up on
th e i n tellige nt coOp era ti on o f a l l th ei r membe rs bu t ,

must o rg a nize d efi ni te s a fegu a rds a g a i nst c ont a mi na ti on

o f w a te r a n d fo od supplies e g w a te r systems p a steu ri
,
. .
, ,

z a ti o n o f milk s cree ni ng o f f o od f rom flies etc

, .
,

CO N TA M I N AT I O N O F T H E AIR f ro m d ried spu tum or ,

 CO N TA M INATI O N OF T H E AI R 23 5

f rom th e c oughi ng of pe rs ons wi th dise a se of th e respi ra

t ory t ra ct mu st a lso be reduced a s much a s p ossible
'

, ,

th ro ugh a c ombi na ti onof educ a ti ona nd a ctive me a su res

of rep ressi on by leg a lly c on stituted h e a lth a uth orities
”
.

n n rdi nan )
’

( He ce “
D o t spi t o .ces
D I R E CT I N FE CT I O N ca n be c ont rolled by qu a ra nti ne ,

th a t is by is ol a ti ng th e a ffected pe rs ons u ntil th e org a n

,

isms o f dise a se withi n thei r b odies h a ve bee n destroyed .

! u a ra n ti ne me a su res a re n o t di ffi cult to e nf or
,
ce p ro

vi ded the dis ea se is disc ove red i ntime by th e a uth orities .

The d ifficulty he re is th a t mild c a ses a re a p t not to c ome

to the a tte n ti ono f physici a ns a nd so f a il to be rep orted .

Th orough g oing m edic a l i nspecti on i n th e p ublic sch ools

-

i s a n imp ort a nt a i d t ow a rd th e c ont rol of c ont a gi ous

dise a se
.
 C HAPTER XX I

I N S E CT S AN D DI S E A S E

B EN EF I CI A L H AR M FU L I N S ECT S — M a ny i nsects
AN D .
,

such a s th e silkw orm m o th h oney bee l a c a nd c och ine a l

, ,

bugs a s well a s seve ra l species which p rey up on i nsect

,

p ests a re o f gre a t be nefit to ma n Th e g re a t m a j ority

,
.
,

a m ou n ti ng to pe rh a p s na med species , c onfe r no

speci a l be nefit a nd w ork no h a rm A c onside ra ble nu m
.

b er such a s bees a nd w a sp s a nd ce rt a i n species of a n

,
ts
an d t rue bugs ma y p roduce inj u ry ra nging f rom t rifli n
,
g
to se ri ou s by i n j ecting p ois on i nt o th e b ody of a no th e r
an ima l by me a ns of stings or pie rci ng m o u th p a rts And .

fi na lly th e re a re m osquit o es ce rt a i nfl ies a nd fle a s which

, ,

in t ru e p a ra si tic f a shi on subsist up on the bl ood o f a n

u nc omfort a ble th ough not se ri ously th re a te ned h ost .

S uch wa s a b ou t th e na tu re a nd exte nt o f ou r i nf orm a

ti on a l ong this li ne p ri or to 1 880 S i nce th a t d a te a
.

l a rge a m ou nt o f w ork h a s bee n d one with p a ras itic i n

sects a nd i t is now k now nth a t while th ey of themselves
,

a re ra rely m ore th a n a n noyi ng pests they a re ve ry i m

p ort a nt a ge nts i n the di s tri bu ti on of s ome of th e m ost
se ri ou s sco u rges a fflicti ng ma n a nd m a ny of the l owe r
an ima ls .

How I N S E CT S T R A N S M I T D I S EA S E — In a few i n
st a nces only a re d ise a ses a ctu a lly p roduced by i nse cts .

Th e bu rrowi ng flea o f t ropic a l c ou nt ries f or ex a mple , ,

tu nnel s its wa y be ne a th th e h um a n ski n a nd with its ,

p roge ny ma y d est roy a ce rt a i na m o u nt o f tissue Ce rt a i n .

fl ies i nthei r l a rv a l o r y ou ng st a ges ma y a ls o occ a si ona lly

i nfest th e h um a n b ody a nd d est roy th e cells T 0 th is .

ex te nt th ese i nsects do i nte rfe re with the p rope r fu nc

236
 FLI E S AND T YPH OID F E VE R 23 7

ti oning ofth e b ody a nd h e nce ma y be s a id to b e the

,

di rect c a use o f dise a se G e ne ra lly spe a ki ng h oweve r

.
, ,

the dise a ses with which insects h a ve to do a re a ctu a l ly

p roduced by b a cte ri a a nd u nicellul a r a nim a ls Th e i n .

sect me rely se rves a s a c a rrie r for these lesse r org a nisms .

Th ey ma y a tt a ch to th e h a i rs or th e oute r su rfa ce of the

i nsect s b ody a nd b e c a rried from pl a ce to pl a ce ; or th e
’
.

dise a se p roduci ng p a ra site ma y a ctu a lly live withi n the

-

b ody of th e i nsect a nd the re u nde rg o ce rt a i n st a ges i n

,

its devel opment bef ore being t ra nsmitted to s om e o the r

org a n ism .

F LI ES AN D TY P H O D FEVER
I — Th e fi rst of the two
meth ods whe reby insects t ra nsp or t ge rms is exemplified
by the h ist ory of the ordina ry h ouse fly In the c o u rse .

of i ts w a n de ri ngs a lw a ys on th e sea rch f or dec omp osing

,

org a n ic m a tte r it ma y settle up on th e exc ret a of a dis

,

e a sed p a tie nt pick up a milli onor two b a cte ri a (a s h a s

,

bee n repe a t edly p rove n) th a t a tt a ch to th e p a ds of its

feet or elsewhe re onth e b ody a nd the na ligh t up onf ood
,

or d ri n king a nd c ooki ng ut ensils a nd dislo dge th ous a nds

o f b a cte ri a i n th e p ro cess By this me a ns dise a se ge rms
.

a re widely distributed a nd u nfortu na tely find new

,

h osts i nwhich to live a nd mul tiply .

The h a bits of th e h ouse fly a re now so well k nown to

the physici a na nd s a nit a ri a n th a t h e is a ble to p resc ribe
e ffective me a ns of c ont rol He knows th a t by disi nfect
.

i ng the exc ret a of typh oid p a tie nts h e ca n p reve nt th e

fly from bec oming a c a rrier of the dise a se org a nism .

More ove r by sc ree ning a ll a nim a l exc ret a whe re the flies ,

b reed h e ca n g re a tly reduce th e number of flies By

, .

sw a t th e fly c a mp a igns h e ca n still fu rth e r reduce

thei r numbe rs especi a lly if this be d one l a te i n the
,

se a s on Then by insisti ng on h a vi ng a ll foods sc reened

.

or o th e rwise p ro tected f ro m fli es h e p reve n ts the few th a t

rem a i nf ro m d oi n g se ri ous h a rm .

F LI E S A N D S LE E P I N G S I C K N E S S — Th rough th e me
.

di um of the p ress much publicity h a s beengivento slee p

23 8 IN S E C T S A N D DI S E A S E
ing sick ness te rribly f a t a l d ise a se a fflicti ng wh ite s a nd
,
a

bl a cks a like i n ce rt a in dis tricts o f t ropic a l Afric a Th e .

fi rst sympt oms o f a n a tt a ck a re o fte n rel a tively sligh t

feve r a nd ski n dis orde rs but g ra du a lly these bec ome ,

m ore p rono u nced a nd a fte r m onths or eve n ye a rs th e

,

ne rv ou s system is a ffected , th e victim l a psi ng i nt o a »

st a te o f u nc onsci o us ness f rom wh ich the re ra rely is a

rec ove ry . Af te r a nimme nse a m ou nt o f l a b or it h as bee n
dete rmi ned th a t this sc ou rge is d ue to mi nute sle nde r ,

an d a cti v el y m ovi ng bl oo d p a ra sites wh ich a re c a rried

’

f rom ma n to ma n by the ski npie rci ng bl ood sucki ng - -

tse tse fly Th is i nsect is a dull b row n f a i ntly b a nded

-
.

species be a ri ng a ge ne ra l resembl a nce to th e ordi na ry

h ouse fly I t m a kes its h ome i n th icket s a l ong th e
.

b orde rs of s tre a ms a nd m a rsh y dist ricts a nd a s is , ,

to be expected sleepi n g si ck ness is c onfi ned to these

,

s a me m oist fly belts O th e r f orms o f this d ise a se a re

.

know n to a tt a ck c a ttle h orses m ules a nd c a mels i n the

, ,

E a ste rn Hemisph e re while a rel a ted a fflicti onis c omm on

,

a m on g h orses i n C e nt ra l a nd S outh Ame ric a I t ha s .

bee n dem onst ra ted th a t wh ile i t is not p o ssible to cu t

d ow na l l th e b rush i na ni nfested dist rict the regi onma y ,

neve rtheless be m a d e c omp a ra tively s a fe by rem ovi ng

th e th i ckets f ro m th e neighb o rh ood o f w a te r b odies ,

b ridges a nd f ords .

F LI ES AN D OT H ER D I S EA S ES Th e repu t a ti on o f the .
-

h o use fly or the typh oid fly a s s ome scie ntists h a ve sug

,

gested it Sh ould be c a lled is fu rthe r bl a cke ned by th e ,

f a ct th a t i t ma y a ls o be th e a ge nt resp o nsible for sp re a d

in g seve ra l o th e r dis orde rs a ffec ti n g ma n I t is ce rt a i nly .

k now nth a t these i nsects ma y fly a t le a st tenmiles f ro m

th ei r bi rthpl a ce a nd ma y c a rry a ny o ne o f no t less th a n
,

S ix ty di ffe re n t ki n ds o f o rg a nism a tt a ch ed to th ei r b o dies ,

a m on g wh ich a re the ge rm s o f diph th e ri a i nf a ntile p a r ,

a lysis ,m e a sles lep ro sy tube rcul o sis Asi a tic ch o le ra

, , , ,

sc a rlet f eve r a nd s eve ra l o th e r imp o rt a nt dise a ses .

H o use flies a re a ls o k now n to c a rry th e eggs o f seve ra l

 MOS ! UI T OES AND MALA R I A 239

species of p a ra sitic w orms occu rri ng i n ma n I t is a n .

est a blished f a ct th a t b o th eggs a nd ge rms a re disl odged

a s the i n sec t w a lks o ve r c ooki ng a nd d ri nking u te nsils ,

f ood a nd the h um a n b ody a nd it ma y i ndeed be t rue

, ,

a s ma n y stude nts of the subj ect believe th a t dise a se is ,

S p re a d o ve r wide a re a s by this p ro cess S w a tti ng th e

”
.

fly is a wis e p reve ntiv e me a su re a nd th e a dditi ona l

,

p rec a u ti onof sc ree ni ng m a nu re p iles a nd o th e r b reeding

pl a ces of this species is the ou nce of p reve nti on th a t
ma y remove the need for the p o u nd of cu re .

M OS Q U I TOE S AN D M ALAR I A — The sec ond m eth od

.

whe reby dise a se p roduci ng org a nisms a re t ra nsmitted

-

from one i ndividu a l to a no th e r is c omplic a ted by th e

f a ct th a t th e i nsec t c once rned is itself p a ra sitized by th e
mic robe In o th e r w ords the i nsec t a cts a s a so c a lled
.
,
-

sec ond a ry h ost i n which the p a ra sitic org a nism u nde r

g oes ce rt a i n st a ges bef ore it is su ited to a life i n th e
h um a n b ody Th e m a l a ri a l p a ra site is a n ex a mple of
.

this wh e re ce rt a i n spe cies o f m osqu t o es a re the dis i .

t ribu ti ng a ge nts .

The p a rasite resp onsible for th e m a l a ri a l dise a se is

a ve ry mi n ute u nicellul a r a nim a l which up on bei ng in
,

trodu ced by th e bite o f a m osquit o i nt o th e bl o od st re a m

o f ma n a tt a cks th e re d bl oo d c orp uscle e nte r s it con
, ,

sumes its subst a nc e a nd th e ndivides i nt o a l a rge number

o f sm a lle r c ells. Th ese esc a p e i nt p th e bl ood a tt a ck new ,

c orpuscles a nd repe a t th e p ro cess Ina few d a y s milli ons

.

of these p a ra sites ma y be p rese n t i nth e bl ood a nd thei r ,

dest ructi on of th e c orpuscles i s resp onsible f or the p a le

ness o f th e skin ch a ra cte rizing this dise a se With th e .

ruptu re o f th e bl oo d cell a n ,
d th e esc a pe o f th e Imp ri s
on ed p a ra sites w a ste p ro ducts a re libe ra ted
,
an d a cti ng ,

a s p o is on s p rod uce th e usu a l feve r a nd ch ills At th e

‘

tim e whe n th e y o u ng p a ra sites esc a p e i nt o the bl ood

they ma y be killed by giving p rope rly g ra du a ted a nd
timed d oses o f qui ni ne .

MA LAR I A L P A R A S I TE S I N T H E M OS Q U I TO — Th e re a re .
240 I N S E CT S AN D DI SEA SE

not le ss th a n 1 000 na med species of m osquit oes yet ,

p rob a bly not m ore th a na h a lf d oze na re c a p a ble of t ra ns

m itti ng th e m a l a ri a l dise a se (Fig Th ese few spec i es
.

a re ch a ra cte rized by sp o tted wi n gs a nd by tilting th e i r

FI G 62
.
— A nonma l a ria l m osqu i to Theoba ldi a
.
-
, inci dcns, m a le
,

s ta nding onthe w a ll (Ab ou t twi ce na tura l S iz c )

. .

K ello gg a nd D oa ne E conomi c Z oology

, .

b odies upw a rd a t a na ngle wh e na t rest wh ile th e b ody ,

i nothe r species is h eld p a ra llel to th e su rf a ce onwh ich i t

rests . I f a m a l a ri a l p a tie nt is bitte nby a mosquit o s ome
p a ra sites a re ce rt a into be withd ra w ni nth e p rocess One .

FI G 62A
. .
—
A ma l a ri a l m osqui to , Anop heles ma culi pcnni s, fema le ,

s ta ndi ng onth e w al l (Abo ut twi ce na t ra l

. u siz e )
.

K ell ogg a nd D oa ne, E con o mi c Z ool ogy

.

of two th i ngs now t a kes pl a ce : I f th e m osquit o is a non

m a l a ri a l one th e p a ra sites a re digested a l ong with th e
bl ood a nd th ei r hist ory is cl osed I f i t is a m a l a ri a l
.

s p ecies the p a ra si tes i nits digestive t ra c t c onti nue their

 Y E LL OW FE V E R AND MO S ! UIT O E S 24 1

a ctivities a nd s oon b ore th rough the s tom a ch w a ll a nd

,

e nte r th e tissues A c o mplic a ted se ries o f ch a nges the n

.

t a kes pl a ce resulting i nth e devel opme nt of a ne norm o us

,

numbe r of spindle sh a ped cells which m a ke thei r wa y

-
,

i nt o the s a liv a ry gl a nd of th e i nsec t Whe n a m os quit o .

a t th is st a ge bites a h um a n bei n g th e p a ra sites esc a pe

th rough th e w ound i nt o th e bl ood a nd once m ore e nte r ,

up on thei r so c a lled life cycle

-
.

Y E LLow F EVER AN D M OS Q U T O S I E — Th e sudde n on

'

S l a u gh t de a dly ch a ra cte r of th e dise a se and th e h elp ,

,

lessness o f th e p opul a ti onof th e st ricke ndistrict to w a rd

off a tt a cks h a ve m a de yell owfeve r one of the m ost
,

d re a ded o f hum a ndise a ses Efforts w ere m a de i nf orm er .

t imes to c omb a t th e sc o u rge by is ol a ti ng th e p a tie nt ,

fumig a ting eve ry a r ticle sent ou t a nd o th erwise t re a ti ng

the c a se a s th ough it we re sm a llp ox B u t these a ttemp ts .
,

e nt a ili ng a v a st a m ou nt of l a b or a nd exp ense p roved o f ,

no a v a il Pe ople wh o h a d neve r beenne a r a yell ow fever

.

victim we re smi tten a n d th e on ly s a fe c ou rse wa s to

,

le a ve the r egi on .

In th e ye a r 1 900 wh ile th e Ame ric a n t roop s were

,

i n C ub a a Y ell ow Feve r Commissi on wa s a pp o i nted to

,

m a ke a th orough i nvestig a ti on of th is dise a se Acti ng .

up on the suggesti on of D r Fi nl a y o f H a v a na th a t a .

ce rt a i n species of m osquit o i s th e resp onsible a ge nt l n

t ra nsmitti ng the dise as e D r C a rroll of the Commissi on
,
.

a ll o wed himself to be b itte n by a m o squit o th a t p revi

o u sl y h a d fed up onf o u r p a tie n ts su ffe ri ng from the feve r .

Ina S h ort time h e wa s st ricke na n d ultim a tely rec overed .

One of h is co w orke rs D r L a ze a r wa s a ccide nt a lly bi tte n

-
, .
,

by a ni nfected i nsec t a nd died O th e r mena ll owed them .

selves to be bitten a nd five d a ys l a te r devel op ed th e

,

ch a ra cte ristic sympt oms Afte r a l ong se ries of i nvesti

.

ga ti on s i t wa s p roved th a t (1 ) yell o w feve r is t ra nsmitted

i nn o o the r wa y th a nby on e ki n d o f m o squit o ; (2) th a t
to t ra ns mi t th e dise a se th e p a ra sites must be dev ou red
by the m osqui to du ri ng the fi rst th ree d a ys of the feve r ;
242 I N S E C T S A ND DI S E A S E
a nd (3 ) twelve d a ys mus t el a pse befo re the m osquit o
is a ble to infec t a no th e r pe rs on Th e p a ra site thus .

resembles th a t o f m a l a ri a i n th a t i t c ompletes ce rt a i n

st a ges o f its devel opme nt i n a sec ond a ry h ost th e ,

m osquit o .

R E M EDI A L M EA S URE S — I t wa s pe rfectly evide nt to

eve ry cle a r thi nki ng i ndividu a l th a t wh e n the m osqu it o
-

wa s sh ow n to be resp onsible for th e t ra nsmissi ono f ma

l a ri a a nd yell o w feve r th ese i nsects sh o uld be exte r
mi na ted Th e oretic a lly th is wa s a rel a tively simple
.

m a tte r All th a t a ppe a red to be necess a ry was to d ra i n

.

the m a rshes or p ou r oil on th e w a te r a nd th us sm o th e r

th e y o u ng a qu a tic w riggle rs ” I n re a lity th e t a sk
“
.

wa s a nd c on ti nues to be one of gig a ntic p rop orti ons .

M a ny dist ricts such a s th e v a lley o f th e Am a z on a re

, ,

too l a rge to be d ra i ned or oil c oa ted I n o the r regi ons

-
.

th e ig nora nce a nd indiffe re nce o f th e pe ople h a s p ro ved

to be a n a lm o s t i n su rm o u nt a ble obst a cle N eve rtheless .

th e re a re e nligh te ned c ommu nities whe re milli ons of

d oll a rs h a ve bee n spe nt for d ra i na ge h o use to h ouse ,

i nspecti on o f p ossible b reedi ng pl a ces th e i nt ro ducti on ,

o f m o squi to dev o u ri n g fishes a nd o the r remedi a l me a s

-

u res Th e results a mply j ustify a ll o f th e expe nse o f

.

time a nd l a b or T od a y yell ow feve r is p ra ctic a lly wiped

.

o u t f ro m th e l a rge r se a p o rts i n th e Wes te rn Hemisphe re

wh e re f orme rly its ra v a ges we re m ost seve re M a l a ri a .

is m ore ste a l thy i n i ts a tt a ck a nd the ge ne ra l public is

,

m ore i ncli ned to t a ke i t a s a m a tte r o f c o u rse a nd a ll ow

th e m osquit o to c a rry o n its de a dly w a rf a re I t is i m .

p ort a nt to no te h oweve r th a t a nnu a lly no t less th a n

, ,

pe ople die f ro m this dise a se i n the U nited S t a tes

a l on e a nd th e ye a rly m o ne ta ry l o ss d ue to th e redu ced
,

effi cie ncy of su fl eri ng w orke rs a m ou nts to m a ny milli o ns

o f d oll a rs Whe n these f a cts a re ge ne ra lly k now n a nd
.
,

th e p a rt th e m osquit o pl a ys i nthe g rim t ra gedy is u n de r

s tood i t is s a fe to sa y th a t m a l a ri a will bec o me a s ra re
,

a s ye l l o w feve r .
 P L AGU E 43

P LAGU E — Thi s dise a se s ometimes k now n a s bl a ck

,

de a th h a s for m a ny ce ntu ries bee n one of th e m ost

,

te rrible dise a ses k now n to ma n E a rly i n the C h risti a n .

Era it ra ged f or m ore th a n fifty ye a rs a n d swept a w a y

th e p e ople o f m a ny dist ricts Ag a i n in th e l a tte r p a r t
.

o f th e F o u rtee n th C entu ry a n outb re a k occu rr ed th a t

S p re a d o ve r the e n ti re w orld a nd dest royed i t is esti
, ,

m a ted ,
p e ople In m ore rece nt times othe r
.

epidemics h a ve o ccu rred a nd while of less ex te nt thei r

, , ,

e ff ects h a ve neve rth eless bee n f righ tful .

D u ri ng one of these rece nt o utb re a ks i t wa s disc ove red

th a t a mi nute ge rm i nv a ri a bly o ccu rs i n th e b odies
of pl a gue —a ff licted p a tie nts a nd it wa s a ls o p roved
,

th a t th is ge rm wh eni t g a i ns a ccess to th e bl oo d of o the r

i ndividu a ls p roduces th e dise a se It h a s bee n know n .

for ce ntu ries th a t wh e n the re is a n o u tb re a k o f pl a gue

the re is a ls o a high de a th ra te a m ong ra ts Th ese a nim a ls .

migh t the ref ore i n s ome wa y be r esp onsibl e for th e

dise a se but i nwh a t wa y wa s disc ove red les s th a ntwe nty
,

ye a rs a go .

F LEA S AN D P LA GU E . I t is now knownth a t d ogs c a ts , ,

mice a nd espe ci a lly ra ts a re subj ect to th e a tt a cks of

, ,

pl a gue a nd fu rth erm ore it h a s bee n S h ow n th a t wh en

,

such a nim a ls die of th e dise a se th e fle a s wh ich i nfest

t hei r b odies le a ve a nd seek new h o sts The new vi ctims .

ma y be a lm o st a ny o f th e w a rm bl ooded a nim a ls a nd -
,

whe na tt a cked by th e flea th ey devel op th e dise a se owing

to ge rms i nt roduced i nt o th e b ody Th is s a me i n sect ma y .

esc a pe a nd i nocul a te new h osts or th e s a me p a tient ma y

,

be bitte nby othe r fle a s wh ich then t ra nsmit th e dise a se

,

to o the r a n im a ls And th us th e dise a se sp re a ds due i n

.
,

eve ry i nst a nce to th e a ctivity o f this one type of insect .

These results a re now defi nitely est a blish ed a nd h a ve ,

led to th e int roducti onof i nte rna ti ona l s a nit a ry me a sures

which h a ve for thei r g oa l (1 ) the dest ructi on of ra ts
an d o th e r flea c a rryi ng a nim a ls i n pl a gue i nfested dis
- -

tri cts a n
, d (2) th e is ol a ti on of st ricke n hum a n beings .
24 4 I N S E C T S AND DI S E A S E
Th e v a lue of such p rec a u ti ona ry me a su res h a s bee n
a bu n d a ntly dem onst ra ted i nseve ra l i nst a nces a nd j us ti ,

fies th e belief th a t he nceforth i t will be p o ssible not only

gre a tly to reduce the numbe r o f c a ses bu t to p reve nt th e
,

S p re a d of th e dise a se .

OT H ER I N S E CT S AN D D S A S E
I E — Once m ore a tte nti on
i s di rected to th e f a ct th a t m ost insects a re h a rmless ,

an d th e c omp a ra tive few th a t a re k n ow n to t ra n smit

dise a se ca n be kep t withi n b ou nds whe n a c ommu nity
exe rcises a re a s ona ble a m ou nt of c a re Bu t u nfortu .

na tel y the re a re c ommu nities wh e re cle a nliness is not

a vi rtue a n,
d lice bedbugs a n
,
d c ock roa ch es fl ou rish An .

ex a mi na ti on of th ese i nsects a mid such su rrou nd ings

discl oses the f a c t th a t th ey like th e fl e a s ofte nc a rry on
, ,

thei r b odies l a rge numbe rs o f dise a se p roducing ge rms -

,

an d it is now th oroughly est a blished th a t typhu s a n d

th e rel a ted t re nch feve r a re c a rried by lice The re is .

a ls o c on side ra ble e vi dence th a t pl a gu e a nd inf a nt ile

p a ra lysi s a re b orne f rom one a nim a l to a no the r by th e
be dbug Tube rcul osis typh o id feve r a nd Asi a tic ch ole ra
.
,

a re believed by s ome scie n tists to be sp re a d by c ock

ro a ch es
. And th e re a re seve ra l o th e r dise as es a tt a ck ,

in g m a n a s we l l a s m a ny d o mestic a ted a nd wild a nim a ls ,

wh e re i nsects a re st rongly suspected of bei ng the me a ns

o f dist ribu ti on .

M ITES T I C K S AN D D I S EA S E — Al th o ugh th ese org a n

,

isms a re m ore cl o sely rel a ted to th e spide rs th a nthey a re

to the insects thei r h a bits a n
,
d meth ods o f t ra nsmit ti ng
dise a se a re so simil a r th a t th ey ma y be c onve nie ntly
t re a ted i n this ch a pte r Th e mi tes a re not ce rt a i nly
.

k now n to t ra nsmit dise a se i n ma n bu t th e f a ct th a t

,

th ey a re resp on sible for i tch sc a b m a nge a nd o th e r ski n

, ,

d is ord e rs b ri ngs th em u nd e r suspici on Th e ticks a re .

i nv a ri a bly v o ra ci ous p a ra sites wh ich pie rce th e ski n of

the victim with needle like m ou th p a rts a nd g orge them
-

selves wi th b lo od In this ope ra ti on they ma y suck up

.

minu te org a nisms which p roduce a c onside ra ble numbe r

 G O V E R N ME N T S AN ITATI O N 24 5

of dise a ses a ff ecti ng ma na nd th e l owe r a nima ls . R el a ps

i ng nd R ocky M ou nt a i n or Sp o tted feve rs i n ma n th e
a ,

de a dly a nd e norm ously c ostly Tex a s c a ttle feve r ce rt a i n ,

disorde rs a ff ecti ng p oult ry a nd m a ny o th e r dise a ses a re

due to tick c a rried p a ra sites
-
.

G OVER N M E N T SA N I TATI O N Th e f oreg o ing p a ra

.

g ra ph s p resent only the b a res t o utli ne of th e imp ort a nt

subj ect of i nsects inrel a ti on to dise a se a nd of necessity
,

f a il to give a nythi ng like a n a dequ a te c oncep ti on of th e

enorm ous a m ou nt of w ork th a t h a s bee n d one a nd is
now bei ng c a rried oni n this a nd othe r c ou nt ries The .

U S Public He a lth S ervice with b ra nch offices i np ra o

. .
,

tic a lly eve ry st a te a nd city of th e U ni on not only en ,

f orces qu a ra ntine regul a ti ons a nd p revents th e sp re a d

of dise a se bu t i ts t ra i ned offici a ls a re i nvestig a ting
,

dise a se p roducing org a nisms a nd their me a ns of disp os a l

-
.

Th e U nited S t a t es D ep a rtme nt of Agricultu re st a te ,

a g ricultu ra l sch ools ex p e riment a nd field st a ti ons

,

th rough out th e c ou nt ry a re l ikewIse d ev o ting much of

th ei r time a nd m oney to th e study of dise a se i n bo th
an im a ls a nd pl a nts Colleges public sch ools a nd physi
.
, ,

ci a n s a re a ls o pl a yi ng a p romine nt p a rt , a nd t ogether
with bulletins rep orts m a g a zine a rticles a nd th e p ress
, ,

a re educ a ti n g a nd a ro usi ng th e gene ra l p ublic to t a ke

a na ctive pa rt i nthis w a rf a re a g a i nst i nsects a nd dise a se .
 T I ON 6

D E AT H
 C HAPTER XX II

D E ATH AND T HE DU R ATI O N OF LIFE

NATURE OF D EAT H D e a th is de fined a s the cess a ti on

.

of life.Anorg a nism th a t for weeks or ye a rs h a s m oved ,

u nde rg one w a ste rep a i r a nd g rowth a nd h a s r esp onde d

, , ,

to stimuli fi n a lly ce a ses these a ctivities

,
It dies a nd the .
,

p rotopl a sm c omp osi ng its b ody neve r reg a i ns its c a p a city

i n this respect b u t dec omp oses i nt o S imple r m ore ele
, ,

me nt a l a nd lifeless c omp ou nds Th e a dvent of de a th is .

p receded a t le a st i nhigh e r a nim a ls by ol d a ge ch a nges

, ,

inwhich the o pe ra ti ons of the cells gra du a lly S l ow d ow n .

Eve nwhe nde a th occu rs its e ff ects a re no t i nst a nt a ne ous

,
.

Th e ne rv ous system f a ils i nits fu ncti ons the h e a rt st ops ,

be a ti ng b re a th i ng ce a ses ; but th e cells of th e S ki n a nd

,

h a i r ma y c ontinu e to devel op for h o u rs a nd th e white ,

c orpuscles exhibit thei r ch a ra cte ristic m oveme nts .

Am ong m a ny o f th e l ower a nim a ls these di ffe re nces i n

cell vit a lity a re even m ore m a rked seve ra l fu ncti ons ,

of the b ody c on tinuing for a c onside ra ble pe ri od eve n

th ough th e h e a d be seve red A numbe r of a nim a ls such
.
,

a s the op o ssum a n d ce rt a i ni nsects ma y feign de a th a nd , ,

the c a ses of so c a lled suspended a nim a ti on i n wh ich a

-
,

hum a nbeing for ex a mple S i nks i nt o a t orp id c onditi on

, , ,

a re o fte n such p e rfec t c o un te rfeits of de a th th a t th e

cl osest sc ru ti ny is necess a ry to detec t the di ffe re nce .

H I BER N ATI O N OR W I N TER S LEE P A typ e o f p a r .

ti a l ly suspe nded a n im a ti on de a th like i n i ts ge nera l

,
-

fe a tu res a nd know n a s h ibe rna ti on or wi nte r Sleep ,

norm a lly occu rs i n nume ro us a nim a ls du ring th e c old

se a s on whe n f ood is sc a rce In th e f a ll seve ra l species
.

of be a rs hedgeh ogs p ra i rie d ogs squi rrels b a ts a n

, ,
d o ther
, , ,

249
25 0 D E AT H AN D THE DU R ATI O N OF LI FE
h igh e r a nim a ls c onst ruc t nests a nd eve n before the , ,

c oming o f th e wi nte r withd ra w i nt o these ret re a ts a nd

, ,

e nte r up on a p eri od of sleep l a st ing f rom two to f ou r

m onth s N o f ood is t a ke ndu ri ng th is time th e p ulse is
.
,

ve ry feeble b re a th ing m o veme nts a re sligh t a nd occu r a t

,

l o ng inte rv a ls a nd th e he a t ge ne ra ted by the oxid a ti on

,

o f th e f a tty tissues especi a lly whe nit is ret a i n ,

ed by the
w a lls o f th e nes t is sufficie nt to keep th e a nim a l f ro m
,

f reezi ng D u ri ng h ibe rna ti on the refore a ll o f th e ch ief

.
, ,

f u ncti o ns o f th e b o dy a re bei ng c a rried on it is believed

‘

, ,

bu t th ey a re sl owed d ow n a lm o st to the p oint o f com

p l e te i na ctivity .

S U M M ER S LEE P — In t ropic a l a nd dese r t c o u nt ries :

.

whe re d rough t p rev a il s for m onth s a t a time m a ny species ,

of a n im a ls a nd pl a nts u nde rg o the so c a lled summe r S leep -

th a t resembles h ibe rna ti on i n ce rt a i n of its fe a tu res .

M a ny s na kes beetles c roc odiles a nd a llig a t ors up onth e

, , , ,

a pp roa ch of th e dry se a s on bu ry themselves i nth e mud , ,

an d with a ctivities a t l ow ebb exist i n a d orm a nt con

, ,

d iti on for m onths Am ong th e l owe r a nim a ls th e re a re

.

ce rt a i ns na ils a nd slugs th a t su rrou nd th e exp osed p orti ons

o f th ei r b o dies with a c oa t o f slime a n d i nth is st a te ma y ,

exist with o u t feedi ng for a t le a st two ye a rs P ra ctic a lly .

a ll o f th e f reshw a te r u n icellul a r a nim a ls ma y be d ried

i nt o i ne rt sh rivelled m a sses exp o sed to th e bu rni ng sun , ,

an d bl ow n a b ou t by th e wi nd a n d c onti nu e to exist i n ,

a n a pp a re n tly lifeless c onditi on u ntil a m ore f a v ora ble

time N ume ro us pl a nts likewise dry up a nd sh ri nk to
.

sm a ll size wh ile the seeds like the eggs o f m a ny a nim a ls

, , ,

a re p ro tected by h e a vy c oa t s to e n a ble th em to withst a n d

excessive d ryi ng for seve ra l ye a rs .

E FF E CT S O F FR EE Z I N G I t h a s l o ng bee nk now n th a t
.

nume rous species o f a nim a ls a nd pl a nts ma y be f ro ze n

s olid yet reg a i n th ei r cust om a ry a c tivity whe n a g a i n
b rough t i nt o f a v o ra ble c onditi o ns A s a m a tte r of f a ct .

this no rm a lly occu rs eve ry wi nte r i nc old clim a tes For .

ex a mple i n th e Ar ctic regi on o f N orth Ame ric a th e

,
 '

L E NGTH OF LI FE 1

g rou nd freezes to a depth of seve ra l feet a nd th e pl a nts ,

an d m ost of th e a nim a ls th a t live a t or nea r the su rf a ce

a re c ompletely f ro ze n f or seve ra l m on ths In this c a te .

o ry a re t rees sh rubs an d h e rb a ce o us pl a nts w rms

g , , o , ,

sna ils i nsects a nd l owe r f orms of life Als o i nthe sh a ll ow

,
.

p onds a nd st re a ms of Al a sk a the re is a S pecies of fish

( D a lli a ) th a t eve ry wi n te r is f ro ze n s olid I nd ividu a ls .

c a ught i n N ovembe r h a ve bee n kep t i n thi s c onditio n

u ntil the f oll owi ng M a rch a nd up on bei ng th a wed ou t
,

a ppe a red n on e the w orse for the expe rie nce The re is .

s ome evide nce th a t be a r a nim a lcules a l ow species of ,

an im a l fou nd fro ze n i ngl a ci a l ice ma y h a ve existed in

, ,

this imp ris oned c onditi on for m a ny p ossibly sc ores of , ,

ye a rs I t is c onceiv a ble th o ugh not p rob a ble th a t s ome

.

o f the a ctivities o f th e b o dy ma y be c a rried on i n these

f ro ze n org a nisms but the re i s n

,
o visible S ig n o f life

an d th e de a th limit is ce rt a i nly cl ose a t h a nd .

LE N GT H OF LI F E — Acc ording to a n ol d s a yi ng a ,

w re nlives th ree ye a rs a dog lives th ree times a s l ong a s

,

a w re n a h orse lives th ree times a s l on g a s a dog a nd a

,

ma n lives th ree times a s l ong a s a h orse ” Alth ough .

,

these figu res a re s omewh a t i na ccu ra te wh e n the a v era ge

le ngth of li fe is c onsidered they c a ll a tte nti onto the f a ct
,

th a t th e life of eve ry species of a nim a l a nd pl a nt is

limited A w orke r h oney bee lives six weeks a nd b e
.
-
,

tween this limit a nd th e h u nd red ye a r existe nce of th e

-

eleph a nt a nd e a gle o th e r species ca nbe f ou nd with a lm ost

an y in termedi a te p e ri od of a ctivity .

Th e expl a na ti ons offe red by bi ol ogists to a cc o u nt for

these f a cts h a ve bee n o f th e m ost v a ried ch a ra cte r a nd ,

yet none a t th e p resent ti me meets with ge ne ra l a pp rov a l .

Some scie ntists h a ve a sse rted th a t size de te rmi nes th e

a ge limit ; th a t l a rge a n im a ls live l onge r th a nsm a ll ones .

This is u nd o ubtedly t rue withi n limits An eleph a nt is .

twe nty fi v e ye a rs o f a ge bef ore it a tt a i ns full size a nd a

-
,

series ca nbe a rra ng ed a s f oll ows : eleph a nt 1 00 ye a rs ; ,

h orse , 4 0 ye a rs ; bl a ckbi rd 1 8 ye a rs ; m ouse 6 ye a rs

,
25 2 D E ATH AND T HE D UR ATI O N OF LI FE
butte rfly 1 ye a r N e ve rth eless wh e nth is exp l a na ti onis
, .
,

c ritic a lly ex a m ined m a ny excep ti ons a ppe a r A h orse .

lives 4 0 ye a rs bu t so d oes a sea a nem one or a t oa d ; a pig

,

lives 1 8 ye a rs a nd so do s ome of the c ra bs M e re b odi ly

,
.

size is the refore no t th e s ole dete rm ini ng f a ct or .

W EI S M A N N S T H EORY ’
T ow a rd th e cl ose of th e l as t
.

ce ntu ry Weism a nn a f a m ous G e rm a nz oOlo gi st formu

, , ,

l a ted a the ory th a t a ppe a rs to be m ore in a cc ord with

wh a t is know nof th e subj ect He emph a sized th e f a c t .

th a t i n m a ny species of a nim a ls a ll of th e sex cell s a re ,

libe ra ted a t one t ime ; a nd if th e oflsp ri ng a re not sub se ,

q ue ntly c a red f or by th e p a re nts th e l a tte r die a t once ,

o r exist f o r a sh ort time on ly Wh e re th e eggs a re not .

l a id i none se a s on bu t d uri ng a pe ri od of week s or ye a rs

, ,

i t is obvi ous th a t th e Sp a nof life is m ore exte nded , bu t

h e re a ls o i t l a sts but li ttle l onge r th a nth e tim e of sexu a l
m a tu rity Wh e re th e y ou ng a re c a red for th e lives o f
.

s uch p a re n ts a re c onti nued for a rel a tively much l onge r

time a fte r th e cl ose of the rep roductive p e ri od th a ni nth e
o th e r c a ses cited .

Th e numbe r of individu a ls of a species rem a ins a p proxi

m a tely th e s a me f rom ce ntu ry to ce ntu ry Th e re ma y be .

fl uctu a ti ons one ye a r bei ng rep rese nted by m a ny m ore

,

i ndividu a ls th a na no th e r bu t i nth e l ong runth e a ve ra ge

,

is p ra ctic a lly u niform This must me a nthe ref ore th a t for

.

eve ry p a re nt th a t dies th e re is a nothe r i nd ividu a l to t a ke

its pl a ce wheth e r th e rep roductive pe ri od be l ong or
,

sh ort .

A PP LI CAT I O N O F T H E T H EOR Y — Th e g re a t le ngth of .

life of th e eleph a nt i s thu s c orrel a ted with th e f a ct th a t it

is a sl o w b re ede r I t d oes not p rod uce o ff sp ri ng u ntil
.

twe nty fi ve ye a rs of a ge a nd a t a m a ximum p rob a bly

!

-
,

d o es not give bi rth to o ve r fi fty Th e e a gle d oes not .

build i ts nest u ntil it is seve ra l ye a rs o f a ge a nd it h a s ,

bee nestim a ted th a t i t d oes no t l a y o ve r one hu nd red a nd

fi fty eggs I nb oth of th ese i nst a nces th e numbe r of d e
.

scen da nts is sm a ll bu t th ey receive p a re nt a l c a re On .

,
 OL D A GE M O DI FI C ATI O N S 25 3

th e othe r h a nd the V i rg i ni a oy ster l a y s f orty milli ons of

'

eggs bu t these a re not c a red for Acc ord ingly th e numbe r

,
.

of y o u n g st a nd s in cl ose rel a ti on to thei r ch a nces of

c omi ng to m a tu ri ty a nd (a cc ordi ng to this th e ory ) th e
,

le ngth of time requi red to p roduce a sufficie nt numbe r o f

O ffsp ri n g to repl a ce the p a re nts is cl osely c orrel a ted with
the a ve ra ge le ngth of life of th e m a tu re i ndividu a l I t is .

difficult to see h oweve r h ow th e c orrel a ti ona ctu a lly fixes

, ,

the limit of the life of a ny i ndividu a l .

OLD AGE M OD I F I CAT I ON S I nf a ncy y ou th , middle

.
,

life a nd old a ge a re me rely st a ges i nth e p rocess of grow

,

ing ol d D u ring i nf a ncy th e chemic a l ch a nges i nthe cells

.

g o o n a t a r a pid ra te cell divisi

,
o n s o ccu r a t f reque n t i n
terv a l s the p rot opl a sm i n
,
c re a ses to a g re a t exte nt a nd ,

the b ody grows to a m a rked deg ree Beyond th is p oint .

th e i nc re a se i n weigh t i n th e h um a n species is much

sl owe r a nd a fte r gra du a lly sl owi ng d ow n st ops a t a b out
, ,

twenty ye a rs i n the c a se of the fem a le a nd twe nty f ou r -

n ri n fi rm th e p rop orti on f
i the m a le D u . g this s o th e
b ody Unde rg o ch a nges wh ich c onti nue th rough ou t life ,

an d a re usu a lly s o definite th a t th ey a ccu ra tely i ndi c a te

the a ge of th e individu a l .

I nne r ch a nges a ls o a re ch a ra cteristic of th e b ody a t

v a ri ous st a ges of its growth Inol d a ge th e b ones con
. _

t a ina rel a tively l a rge a m o unt of m inera l s a lts a nd bec ome

b rittle Th e c a rtil a ge a ls o is imp regna ted with mi nera l
.

s a lts a nd in c onseque nce bec omes less el a stic The .

muscles l ose their vigor a nd th e h a ir its p igme nt Th e

,
.

le ns of th e eye l oses its el a sticity a nd th e a bility to dis

tin gu i sh n e a r obj ect s cle a rly Th e met a b olism of th e
.

b ody bec omes m ore impe rfect a nd wh a t a ppe a r to be ,

dege ne ra tive ch a nges a ff ect th e n e rv ous system .

I M M EDI ATE CAU S ES OF D EAT H P opul a rly it is b e

”
.

l i ev ed th a t the vit a l m a ch i ne “
we a rs ou t or runs “

d ow n bu t bey ond this p oi nt no det a i ls o f th e p rocess a re

,

forthc omi ng Th e physici a n is i ncli ned to st a te th a t a

.

ma nis a s ol d a s h is a rte ries me a ning th a t if th e ch a ra c

,
254 D E ATH AND T HE D U R ATI O N OF LI FE
teri sti c ol d a ge thicke n
-
i ng of th e w a lls of th e bl ood vessels -

d o es no t o ccu r th e a tte nd a nt dis orde rs a re a bse nt a nd life

is likely to be p rol onged Bu t he re a g a i nno s a tisf a ct ory
.

re a s onis give nfor th e so c a lled h a rde n

-
i ng o f th e a rte ries .

Acc ording to o the r stude nts de a th is due to ch a nges i n

th e ne rv ous system or i n th e ra te o f met a b olism or i n
, ,

th e rel a ti on o f th e nucleus to th e cell p ro t opl a sm or to ,

th e m odific a ti onof th e physic a l or chem ic a l c onstituti on

o f th e cell ; a n d once m ore det a ils a re l a cki ng .

S till o th e r i nvestig a t ors h a ve suggested th a t th e b ody ,

a s a resul t o f its a ctivities ge n e ra tes p ois ons o the r th a n

,

th ose elimi na ted by th e S ki n a nd kid neys a nd thes e ,

a ccumul a tin g i n th e cells p reve nt the i r ope ra ti on .

M etch niko ff a ro used gre a t p op ul a r i nte rest i nthis subj ec t

by th e suggesti onth a t i nma nth e l a rge i ntesti ne h a rb or s
gre a t numbe rs of b a cte ri a wh ose exc reto ry p o is ons a b ,

s orbed by th e b ody p roduce distu rb ances ultim a tely re

,

su l ti ng i n de a th . Bu t a ttemp ts to dest ro y th e b a cte r i a

h a ve not bee n p rod uctive of m a rked res ults so f a r a s

th e le ngth e ni ng of life is c once rned .

I t a ppe a rs to be t ru e th a t ve ry o fte n if no t inv a ri a bly , ,

s ome one org a nce a ses to fu ncti ona nd he nce th rows th e

o th e rs o u t o f a dj ustme n t Bey ond this p oi nt no scie ntist
.

h a s give na ve ry s a tisf a ct ory expl a na ti onof the i mmedi

a te c a uses o f de a th .

I M M ORTA LI TY O F T H E U N I CE LL U LAR ORGA N I S M S I t is .

a st a rtli n g fa c t th a t de a th is not a ch a ra cte ristic o f th e

l o west forms of life An a moeb a or p a ra m oecium for
.
,

ex a mple multiplies by th e d ivisi ono f the cell int o equ a l

,

h a lves P a re nt A i nth e p rocess bec omes off sp ring B a nd

.

C The re is no d e a th ; no c orpse Th ese simple org a nisms

. .

a re thus p o te n ti a lly imm ort a l S ome i ndividu a ls to be

.
,

su re ma y be b oiled c rushed st a rved o r c a rried i nt o un

, , , ,

fa v ora ble situ a ti ons whe re they pe rish ; but these a re

c a s es of a c cide nt a l de a th only a nd u nde r no ci rcum ,

s ta nces do such cells p rod uce d esce nd a nts In o th e r .

w ord s eve ry one o f these p rimi tive c re a tu res is th e de

,
 D E ATH I N O R GANI S M S 255

sce nda nt of p re existi ng p a re nt a l cells These st a teme nts

-
.

h a ve bee n c riticized from v a ri o us a ngles by di ffe re nt

scie ntists bu t th e f a ct nevertheless rem a i ns th a t no one “
,

celled org a nism eve r l o st a n a ncest or th ro ugh de a th ” .

D EAT H I N M U LT C I ELL U AR
L O R GAN SI M S — Th e m a ny .

celled org a nism s a re likewise desce nd a nts of p re existi ng -

p a re nts yet i n th is g roup de a th is a regul a rly recu rri ng

,

p ro cess Whe rei n lie s th e di ffe re nce in this p a rticul a r

.

betwee n the multicellul a r a nd u nicellul a r species ? Th e

an swe r is forthc omi ng whe nth e devel opme nt of the fi rst
na med cl a ss is c ritic a lly ex a mi ned Th e egg cell of th e.
-

h en for ex a mple by repe a ted divisi ons gives rise to a n

, ,

i ndividu a l i n wh ich th e cells bec ome di ffe re nti a ted i nt o

two ve ry disti nct types th ose of th e b ody p rope r a n
,
d th e
ge rm cells Th e b ody is the m ore obvi ous a nd imp ress i ve
.
,

an d a ppe a rs to be th e m ore imp ort a nt I t i s fitted f or a .

defi nite set of c onditi ons a nd displ a ys a se ries of com

plex a ctivities of g a the ri ng food a v o idi ng e nemies a nd
, ,

begetti ng a nd c a ri ng for th e y o u ng N ev ertheless it is .

a me re she a th f or the r ep ro ductive eleme n ts th a t it

nou rishes a nd p rotects a nd which a l one a re c a p a ble of
,

p roduci ng the succeedi ng ge ne ra ti on .

Inth e l owest multicellul a r org a nisms such a s Gonium , ,

e a ch cell rep roduces its kind while i nh ighe r typ es where

, ,

the divisi onof l a b or is m ore c omplete only th e ge rm cells ,

-

h a ve ret a i ned this p owe r Th e b ody wh e n emptied of

.

its rep roductive cells th us bec omes a w orthless h usk a nd ,

while th e p rot opl a smic st re a m fl ows f rom ge rm cell t o -

ge rm cell de a th cl a im s th e b ody a t e a ch ge ne ra ti on
-
.

—
,

D EATH AN AD V A N TA GE T o T H E S P E CIE S Th e f ood

supply of eve ry sp ecies of pl a nt a nd a nim a l is limited ;
m ore i ndividu a ls a re p roduced t ha n ca p o ssibly exist n
th rough ou t th e norm a l lifetime a nd it the ref ore is evi
,

dent th a t i n th e u nce a si ng st ruggle for existe nce th e fi t

test individu a ls sh ould pe rsist i f th e species is to be kept
up to a high st a nd a rd M a nis no excepti onto this rule
'

.
.

Wh a tever our se ntime nts ma y be i n th e m a tte r i t must ,

25 6 D E ATH AND T HE D U R ATI O N OF LI FE
be evide nt th a t pe rmitti ng th e physic a lly a nd me nt a lly
unfit to rep rod uce th ei r ki nd is a se ri ou s me na ce to the
M ore ove r it is a ls o true th a t th e m ost pe rfec t i n
'

ra ce .
,

di v i du a l s o f a n y species o f a nim a l o r pl a nt bec ome i h

crea sin gyl c rippled a n d defective th e l on ge r they l ive ;

an d a cc ordingly the re i s a p ress ing need for the d evel op
me nt of new a nd m ore vig orous oflsp ri ng to t a ke their
p l a ces Whe n th ese desce nd a nts h a ve bee n p roduced
.
,

n
a d a fte r th ey a re c a red f or — i f th is is ch a ra cte ristic of
th e sp ecies — th e life w ork of th e p a rents is a t a nend .

Th e i nte rest of the ra ce i nv a ri a bly a b ove th a t of the i n

,

dividu a l i s best se rved by th e rem ov a l of th e ol d a nd

,

dec repi t D e a th th e ref ore is a na dv a nt a ge to the species

.
,
.

S U M M AR Y — D e a th wh ich be a rs a cl ose resembl a nce

,

.
,

to suspe n ded a nim a ti on is th e c omplete a nd p e rm a ne nt

,

st opp a ge of th e vit a l fu ncti ons I t occu rs norm a lly a t a

.

tim e a fte r th e pe ri od of sexu a l m a tu rity a nd th e c a re of

the y ou ng h a s c ome to a cl ose Am ong th e u nicellul a r
.

org a n ism s a nd th e ge rm cells o f multicellul a r species ,

de a th is a na ccide nt a l p rocess It me rely a flects th e b ody

.

th a t e ncl oses a nd nou rish es th e rep roductive eleme nts .

D e a th th e ref ore is a be nefit to th e ra ce i n rem ovi ng

, ,

i ndividu a l s wh ich othe rwise w o uld te nd to c rowd out th e

m ore vig orous a nd u se ful off sp ri ng .
 S E CTI ON, 7

T HE R OLE OF M I CR OOR GA N ISM S

 C HA PTER XX III

D E CO MP OS ITI O N

T H E D I S P OS A L O F OR GA N I C W A S TE S — The sy ntheses
which a re c a rried on by pl a nts in th ei r own b odies a nd ,

the met a b olic ch a nges in th e b odies of a nim a ls p roduce , ,

a m on g Othe r thi ngs i ns oluble subst a nces which bec a use

, ,

they a re ins oluble a re of no imme di a te use to the pl a nt

,

or a n im a l a fte r they ce a se to be p a rts of th e livi ng

mech a nism th a t is a fte r they a re exc reted or a fter the
, ,

org a n ism dies These m a tte rs w o uld a ccumul a te a nd

.

cumbe r th e e a rth i f th ey were not rem oved R em ov a l

”
. .

is a cc omplish ed by th e sl ow p rocesses of we a thering “

an d by dec omp ositi on chiefly fe rme nt a ti on, a nd p ut re

,

f a ct i on We a the ri ng c onsists i nth e ope ra ti on of i na ni

n
.

m a te i nflue ces th e a cti on of the oxygen o f th e a i r of

, ,

light he a t fro st w a te r wi nd d ryi ng etc Th e o the r

, , , , , ,
.

p rocesses a re c a rried on by livi ng th ings m a inly mic ro ,

org a n isms which ope ra te by me a ns of the enzyms a l re a dy

,

refe rred to i n C h a p te r V I I .

D E CO M P OS T O N
I I — Fe rme nt a ti ona nd p ut ref a cti ona re
.

p a rticul a r f orms of dec omp ositi on th e p roducts of which ,

o fte n disti n guish them Th us a lc oh ol a nd th e offe nsive

.

od ors of dec a y m a rk r espectively a ce rt a i n fe rme n t a ti on

n
,

na mely a lc oh olic a nd ce rt a i p ut ref a cti ons Ina ll these

,
.

c a ses th e re a re e ne rgy ch a nges a nd ch a nges i n chemic a l

cbmp osi ti on D ec a y which t a kes pl a ce with offe nsive
.

o d ors is c a lle d p ut ref a cti on S ome f orms o f dise a se .

ga s g a n gre ne ) a re a ctu a lly fe rme nt a ti ons or p ut ref a cti ons

-
.

Th e ch a nges wh ich a re te rmed de ca y fe rme nt a ti on p u , ,

trefa cti on dec omp ositi on a n

,
d th ose which a re th e a ctu a l
,

c a use of dise a se a re fu nd a me nt a lly a like S ome of th e

,
.

259
2 60 D E COMP OS ITI O N
chemic a l a nd e ne rgy ch a nges t a ki ng pl a ce i n dec omp osi
ti ona re th e s a me a s th ose which t a k e pl a ce inthe b od ies
o f the livi n g org a nisms wh ich p rod uced th e m a te ri a l s
a cted up on Thus digesti on a nd dec omp ositi on the
.
,

met a b olic p rocesses i nv olved i n respi ra ti on a nd ce rt a in

fe rme nt a ti ons a nd th e p ost m ort em ch a nges i mmedi
,

a tely f o ll owin g de a th h a ve m a ny fe a tu res in c omm on ,

th e m ost sig nific a nt o f which i s e nzymic a cti on (S ee .

Ch a p V I I ) . .

K I N DS OF D E CO M P OS I TI O N (a ) D iges ti on Th i s
. .

t opic me nti oned he re bec a use it is a p a rt of th e ge ne ra l

,

subj ect h a s bee n discussed elsewhe re (Ch a p V I I ) a t

,
.

su fli ci en t le n gth I t is a cc omplish ed by e nzyms which a ct

.

up onv a ri ous s ort s of org a nic m a tte r .

(b ) A u t o ly s i s This is
. th e na me give n to dec o mp o si
ti ont a ki ng pl a ce i nb odies immedi a tely a fte r de a th a nd ,

i s e nti rely i ndepe nde nt of inv a d ing org a nisms I t is .

a cc omplish ed by e n zyms formed by the org a nism p revi ous

to its de a th I t ma y be m ore th a n suspected th a t some
.

a t le a st o f these ch a n ges t a ki ng pl a ce a fte r de a th a re

th e s a me a s th ose t a ki ng pl a ce before de a th A ut olytic .

ch a nges u nd o ubtedly t a ke pl a ce i n th e b odies of pl a nts

a s well a s a n im a ls a lth ough eve n less is know n a b o u t
,

a ut olysis i npl a n ts th a ni na nim a ls .

( c ) M e ta b o l i s m f o r E nerg y R e lea s e I n b o th pl.a n ts

an d a nim a ls the use of f ood a s fuel results i nits decom
p ositi ona nd the libe ra ti onor rele a se o f e ne rgy Th e ca r .

b oh y dra tes a n d o th e r nonnit roge no us c omp ou n

-
ds a re
oxidized di rectly or i n di rectly i nth e met a b olic p rocesses ,

th e oxid a ti on wh e n c ompleted resulti ng i n th e p rod uc

, ,

ti onof CO a nd H 0 with th e libe ra ti onof e ne rgy Th e

Z 2 ,
.

e ne rgy libe ra ted ma y be immedi a tely used for w ork in

th e f orm of muscul a r or o th e r m oveme nt l oc om oti on , ,

etc ; but if not used for w ork it will esc a pe a s h e a t

.
,
.

( )
d P r o t e i n M e ta b o l is m Th e .c o mplex n it ro ge n o us

c omp o unds o f c a rb on know n c ollectively a s p ro tei ns

c onsti tue nts o f livi ng p rot opl a sm — a re c onve rted i n
 T HE AG E NT S OF D E CO MP O S ITI O N 26 1

th e c ou rse of thei r b a sic met a b olism i nt o simple r com

p o u nds such a s u re a a nd u ric a cid exc reted by a nim a l s i n
,

the u ri ne but with no c orresp ondi ng exc reti onby pl a nts

, .

Th is di ff e rence betweenthe beh a vi or of pl a nts a nd th e b e

h a vi or of a nim a ls ma y be m ore a pp a re nt th a nre a l but ,

p rese nt knowledge th rows no ligh t on th e m a tte r Th e .

nit rogenous c omp ou nds p roduced i nth e met a b olic a cti v i

ties of pl a nt cells seem to be u sed a ga i nby them i n th e ‘

rec on st ructi onof p rotei ns (See C h a p V )

. . .

(e ) C a r b o h y d ra te F e rmenta ti on S ug a r st a rch cell u

.
, ,

l ose a nd its de riv a tives (w ood ,

an d simil a r com
p ou nds a re b roke nd ow ni nthe p rocess c a lled fe rme nt a
,

ti on with the f orm a ti on of a l c oh ols a cids a nd o ther

, , ,

subst a nces a nd fi na lly CO a nd H 0

, Z Th is p rocess is
2 .

a lw a ys du e to th e a ctivities of mi c roorg a n ism s a nd is

b rought a b out by me a ns of e nzyms wh ich they m a nu
fa ctu re In the v a ri o us st a ges of the oxid a ti ons wh ich
.

c onstitute esse nti a l p a rts of fe rme nt a ti on c orresp ondi ng ,

a moun ts of ene rgy a re libe ra ted which c omm only S h ow

a s h ea t .

(f) P u t re fa c ti on In p
. ut r ef a cti o n or g a nic c o mp o ,
u nds
—
c ont a ini ng nit roge n m a ny of them with sulphu r or
ph osph oru s i n a dditi on— a re b ro ke n d ow n c omm only ,

with the p roducti onof off ensive od ors ultim a tely to a m ,

m oni a a nd ca rb on di oxid
’

T H E A GE N T S O F D E CO M P OS I TI O N Anim a ls b a cteri a
.
, ,

an d fu ngi a re ordi na rily the immedi a te a ge nts of decom

p ositi on In m a ny c a ses th e e nzyms p roduced by th e
.

livi ng org a nisms a re know n to f a cilit a te or a ccele ra te if

not to a cc om p lish th e dec omp ositi on He nce th e i nfer .

ence i se a sy th a t a l l dec omp ositi ons wh ethe r a ut olytic or ,

c a rri ed on by i nv a ding org a nisms a re a cc omplish ed by ,

e nzyms .

T H E P R I N CI P LE S OF D E CO M P OS I TI O N Th e p ri nciples .

c ont rolli ng dec omp ositi onw ill bec ome cle a re r if we study
ce rt a i nex a mples be a ri ng Inmind th a t we must ex a mine
,

( )
a the subst a n ce t o be dec o mp o sed ( )
b the o rg a n ism
,
26 2 D E COM P O S ITI O N
p roducing th e dec omp o siti on (c) th e e nzym by me a ns,

of which the org a n ism b rings a b ou t th e dec omp ositi on ,

( d ) th e m a te ri a l p r o ducts a nd (e ),
th e e ne rgy ch a nges .

Th e rel a ti onof th e d ec omp ositi on wh ethe r insid e or out ,

side th e b ody of th e org a nism p rod ucing i t to th e rest o f ,

th e met a b olic a ctivities of th e org a nism sh ould a ls o be

c onside red .

A L CO H O LI C F ER M EN TATI O N so c a lled f rom one o f its

,
-

p rincip a l p oducts
r — n
th e oth e bei g CO
r — is used by z

ma n i n ra isi ng b re a d i n the m a nu f a ctu re o f fe rme nted

, ,

liqu ors a nd i nthe m a nuf a ctu re o f a lc oh ol for use in th e

,

a rts a s fuel etc

, , Owi ng to its imp ort a nc e it h a s been
.

studied with g re a t c a re a nd i nc re a si ng ski ll Am ong th e .

m os t imp ort a nt c ont ributi ons to ou r k nowl edge of th e

wh ole subj ect a re th ose m a de by th e emi ne nt F re nchm a n ,

P a steu r wh ose studies of th e so c a lled dise a ses of wine

,
-

led to defi nite knowledge of th e p rocesses i nv olved i nthe

m a ki ng of a lc oh olic liqu ors I n a lc oh olic fe rm
. e nt a ti on
sug a r is th e subst a nce a ct ed up on Inb re a d d ough a nd .

i n bee rw ort m a lt sug a r is d e rived f rom st a rch i n fl ou r

an d i nge rmi na t ing g ra i n The s a me is t rue o f c orna n
. d
p ot a t o m a sh f rom wh ich B ou rb on a nd I rish whiskies
we re obt a ined In the must o f g ra pes th e fe rme nt a ble
.

suga r f ruct ose a l re a d y exists D i ff e re nt species o f ye a st s

, ,
.

a re the org a n isms p roduci ng the d ec omp ositi on The .

ye a st used in b re a d m a ki ng a nd in bee r m a ki ng is one

- -

f ou nd c omm only i nth e a i r a nd cultiv a ted ona c omme r

ci a l sc a le with th e resul t th a t supe ri o r st ra i n s o r v a rieties
o f ye a st a re n ow o b t a i n a ble j ust a s the fl orist fu rn
,
ishes
supe ri or st ra i ns or v a rieties o f roses a nd ch rys a nth emums .

The fe rme nta ti on of wi ne is a cc omplish ed by me a ns o f

th e wild ye a sts fou nd on the ski ns o f th e g ra pes Th e .

supe ri or qu a lity o f wi nes o f ce rt a i ndist ri cts a nd o f ce rt a i n

se a s ons is d ue i n p a rt to the supe ri ority of th e l oc a l
.

v a rieties o f ye a st th o ugh due a ls o i np a rt to th e f a v ora ble

,

c onditi ons a t th e time of wi ne m a ki ng a nd to the supe -

,

ri or subst a n ces cont a ined i n the gra pes Th e a lc oh ol .

 T HE E N ZY M S CO N CE R N E D 263

bt a i ned from c orn a nd p ot a t o m a sh es ma y equ a l in

”

o .

pu ri ty the a lc oh ol obt a ined from o the r fe rme nt a ble m a te

ri a ls ; bu t th e illicit m a n uf a c tu re o f a lc oh ol f rom such
m a sh es ofte nresults i nthe p roducti onof liqu ors th e b a d
u a lities f which due t the p rese n ce f the sub
q o a re o o o r

st a nces th a nthe p a rticul a r a lc oh ol for wh ich the te rmen

ta ti onwa s c a rried on Th us j a ck a ss b ra n dy m oonshi ne
.
, ,

whiskey a nd th e o the r li qu ors i l leg a lly p roduced owe

,

thei r obj ecti ona ble qu a lities to th e p roducts of o the r

org a n isms such a s th e m olds ra the r th a nto p u re ye a sts
, , .

They c ont a i n so c a lled fusel oils a na me give n to o the r

”
-
,

a lc oh ols th a n th a t obt a i ned from th e cle a n fe rme nt a “

ti on of g ra i n or g ra pes The e ffect of th e fusel oils is

.

much m ore vi ole nt th a nth a t of ordi na ry ethyl a lc oh ol .

T H E E N ZYMS CO N CER N ED Th e hyd rolytic e nz yms .

-

p roduced by th e cell s of th eb a rley g ra i na re (1 ) a my la s e

which digests st a rch c onve r ting it in
.

t o m a lt sug a r (see
“

m
,

Ch a p V I ) a nd (2) ma lta se wh ich splits

.
,
a lt S ug a r with ,

the a dditi onof a m olecule of wa te r i nt o the fe rment a ble

sug a r glu cose Th e chemic a l rel a ti ons of these sug a rs
ma y be i n
,
.

d ic a ted by thei r respective f ormul a s Thu s .

m a lt sug a r is C l O while gluc ose is C l O

zz 1 1 Th e e e.

a cti on of m a lt a se is rep rese n t ed by th e equ a ti on

C1 :2}1 2 2 0 1 1 j Hz o “ “
2C6 H1 2 0 6
g 1u cose

(3 ) Th e oxidizi ng e nzym z yma se c onve rt s t h e su ga rs

i nt o a lc oh ol a nd CO rele a si ng a cert a in a m ou nt of2,

e ne rgy s ome of wh ich ca nbe me a su red a s h e a t Thus it

,
.

is seenth a t a p a rt of the sug a r which wa s ori g i na lly m a de

from CO a nd H O by me a ns of e ne rgy a bs orbed by th e
Z Z

—
chl orophyll pigments i ngree nle a ves is re c onve rted int o ,

CO a n
2 d p a rt o f th e e ne rgy is rele a sed Th us .

C6 H1 2 0 6 2C 2Hs OH 2COz
g l u co s e a l coho l

COM M ER CI A L U S E S P R ODU CT S In ra ising OF TH E .

b re a d COZ i s the esse nti a l p roduct of th e fe rment a ti on .

26 4 D E CO M P O S ITI O N
In wine nd bee m a ki ng a s in th e m a nu f a ctur e of
a r- ,

a lc oh ol the p roducti on of a lc oh ol is the esse n

,
ti a l result .

Th e CO is v a l u a ble i n th e m a nu f a c tu re of bee r a nd i n
”
2

th e p roducti on of ch a mp ag ne a nd oth e r sp a rkl ing

wines Th e a lc oh ol itself p roduced being a p a rtly oxidized
.

c omp ou nd ca nbe fu rthe r oxidized by bu rni ng ; th is ma y

,

be d one in the b ody i n a l a mp o r i n a n e ngi ne or by

, , ,

b a cte ri a In th e fi rst th ree c a ses th e resul t is c omplete

'

oxid a ti ona nd the p ro ducti onof C O a nd H O a nd e ne rgy 2 Z .

Cs OH 30 2 2COZ 3 Hz O
a l coho l
A T FCE I C ER M E N TAT I ON .
—I n th e l a s t c a se ,
o xid a ti on
by b a cte ri a ,
o xid a ti on is i nc omplet e ,

C s OH 02 CH3 COOH + HZ O
‘

a l co ho l a ceti c a ci d

th e p roducts bei ng a cetic a cid f a mili a r to us invineg a r

an d a ce rt a i na m ou nt of e ne rgy s ome o f it rec o gniz a bl e
'

a s he a t I f th e a cti onof th e b a cte ri a is a ll owed to con

.

ti n u e th e r esult will be th e o xid a ti on o f th e a cetic ac id

to CO a n 2 d H 0 an d th e libe ra ti on o f th e rest of th e
2

e ne rgy .

T H E S O UR I N G O F M I L K This p rocess i nv olved in ,

the m a nuf a ct ure of m a ny food s a nd d ri nks h a s a lso been ,

c a refully studied on a cc ou nt o f its tech nic a l a nd com

merci a l imp ort a nce Cow s milk a s well a s th e m ilk of
.
’

o th e r a n im a ls c o nt a i ns a sug a r o f the s a me p roporti ona l

,

f ormul a a s c a ne a nd m a lt sug a rs c a lled milk sug a r sug a r , ,

o f m ilk o r l a ct o se
,
Cms On wh ich ma y be a cted up on ,

by m a ny di ffe re nt org a nisms b o th l a rge a nd sm a ll Wh e n ,

.

milk is digested or a cted up on by ce rt a i nb a cte ri a or by

ce rt a i nye a sts it s ou rs a cu rd f orms a nd th e bu tte r fa t
, , ,

ma y be sep a ra ted off Th e re a cti on so fa r a s th e sug a r

.
,

is c once rned c onsists i nth e splitti ng o f the l a c t ose m ole

,

cul e i nt o fou r m olecules of l a ctic a cid bec a use of th e

a c ti ono f th e b a cte ri a l e nzym k now na s ba c teri a l z yma se .

C HM O“ H2 0 3 5: 40 31 1 60 3
-
 T HE R O TTI NG OF W OO D 265

B ut i f othe r b a cte ri a which f orm a ne nzym c a lled la ctose

a re p rese n t milk su ga r will be split a fte r the a dditi onof
, ,

a m olecule o f w a te r i n t o gluc o se a nd a nothe r sug a r c a lled

,

g a l a ct ose Th is ca nbe c onve rted i nt o a lc oh ol a nd l a ctic

.

a cid by fu rthe r b a cte ri a l e n zym a cti on K oumiss kephi r .

, ,

y o gu rt etc a re milk de riv a t ives used a s d ri nks by ce rt a i n

,
.
,

ra ces ,
wh ich a re now o cc a si ona lly p resc ribed a nd con
sumed i n limited qu a ntities i n this c o u nt ry a nd which ,

a re obt a i n ed by the a cti on of b a cte ri a a nd ye a st on th e

milk of c ows g oa ts etc These v a ri ous p roducts a re
, ,
.

f ormed a t the s a me time th a t a ce rt a i na m ou nt of e ne rgy

is rele a s ed a nd a s they ca nbe still fu rth e r oxidized u ntil
,

CO a n
Z d H 0 a re th e en
2 d p roducts still fu rthe r a m o u nt s ,

of e n e rgy ma y be obt a i ned f rom them Fi na lly a l l the .

,

e ne rgy used i nthe m a nuf a ctu re of sug a r i nthe fi rst pl a ce

will be rele a sed the org a nisms c once rned bei ng duly
,

nou rish ed by th e v a ri ous i ng redie nts of th e milk .

T H E R OTTI N G OF W OOD Th e cell w a lls of pl a nts con

.
-

sti tu te th e bulk of thei r r em a i n s c onsisting Of cellul ose

,

o r s ome de riv a tive o f it a s i n th e w oo d of t rees

, S ome .

an im a ls a re a ble to digest th ese subst a nces by me a ns of

e nzyms which a ct up onthem a nd c onve rt them i nt o sug a r
which they p ro ceed to use i nthe usu a l wa y Th us g oa ts .
,

ca n digest cellul o se a lth o ugh we c a n

,
no t B a rk a nd .

w ood b ore rs digest w ood a nd p roduce sug a r Fu ng i a re

-
.

the chief a gents in cle a ri ng a w a y th e rem a i ns o f pl a nts .

M a ny fu ngi a re a ble to diss olve th ei r wa y th rough w ood

o f b o th livi n g a nd de a d pl a nts by me a ns of p rope r e nzym s .

Whe nfu ngi a tt a ck a nd th us digest the cell w a lls of livi ng -

pl a nts th ey c a use dise a se Whe nthe s a me m a te ri a ls a re

.

a cted up onby org a n isms livi ng up on the lifeless rem a i ns

o f these s a me species the result is be n ,
efici a l The a cti on .

i nb o th c a ses is by me a ns o f a pp rop ri a te e nzyms Th us .

cy ta s e diss o lves p u re cellul o se while ha dro ma s e diss olves

,

w ood Th e fi na l p roducts i na ll c a ses a re CO a nd H O

.
Z Z .

CO M M ERCI A L F I BER P R OD U CTI O N Pectina se diss olves

.

the subst a nce which ceme nts the w a lls of the fib rous cells
2 66 D E COMP O S ITI ON
of fla x nd othe r fi b er p roduci ng pl a nt s P ecti na se by its
a -
.

a ctivities rele a ses th e fibe rs for we a vin g or o th e r me ch a n

i ca l t re a tme nt In p ra c tice th e fibe rs o f fla x a nd o the r
.
,

tex tile m a te ri a l s a re Obt a ined ona c omme rci a l sc a le by a

p ro cess o f ro tti ng i n w a te r know n a s ret ti ng This is .

a cc o mplish ed by o n e o r m o re species o f b a cte ri a livi ng in

w a te r th e desi red a cti on o f which is the p roducti on o f
,

pecti na se which diss olves th e fi b er ceme nting m a te ri a l -

.

H oweve r a s ge ne ra l dec a y a cc omp a nies th is specific a c

,

ti on the st re ngth a nd col or o f the fibe r a re imp a i red a nd

,

th e p rocess is to this exte nt obj e cti ona ble Attempts h a ve .

b eenma de to c a rry onretti ng by me a ns of pu re cult ure

'

an d it is c onceiv a ble th a t u nde r ce rt a i n c on diti ons a t

le a st the p rod uc t ma y b e imp roved i n this wa y I t is .

a l s o c on ceiv a ble th a t a ut olysis ma y be m a de to yield th e

desi red result with ou t c orresp ondi ng inj u ry In these .

fe rme nt a ti ons a s i n th e o the rs a b ove desc ribed the cel l

, ,

w a ll m a te ri a ls a re digested by bei ng c onve rted int o sug a rs

wh ich fu rnish th e f ungi b a cte ri a or othe r org a nisms with
, ,

the f ood which th ey requi re .

D E CO M P OS I T I O N O F F AT S AN D OI LS is a cc omplish ed by
th e a cti on o f org a nisms Th us for ex a mple i n bu tte r
.
, ,

wh ich we ea t a nd bu tte r wh ich we keep too l ong th e -

,

e nzyms a re li p a s es These whethe r f ormed i nth e i ntes

.
,

ti ne or by b a c te ri a b re a k d ow nth e f a ts to glyce rine a nd

,

fa t ty a cids Th ese i n tu rnma y be fu rth e r b roke nd own

.

o r oxidized i nthe cells u n til fi na lly CO a nd H 0 a re th e

, 2 2

en d p ro ducts Corresp on
. di ng a m ou nts of e ne rgy a re
libe ra ted i nth e successive st a ges of dec omp o siti on .

D ECO M P OS IT I O N O F N IT ROGE N O U S W A S TE I t ha s .

bee n sh ow n (C h a p V I I I ) th a t s o me o f th e nit roge nous

.

f ood a nd some o f the nit roge no us m a te ri a l o f th e a nimal

b ody a re b roke n d own in b a sic met a b o l i sm a nd th row n
o ff bei n
, g exc reted m a i nly by th e kid neys i nthe f orm of
u re a Th is p a rtly oxidized c omp ou nd se rves a s food a nd
.
'

fuel for ce rt a i nspecies o f b a cte ri a wh ich fu rthe r oxidize ,

it to a mm oni a a nd 0 0 Th e liquid o rg a nic nit roge nous

2 .
 T HE .
S OLID NIT R O G E N O U S WA S T E S 26 7

w a stes a re th us b roke n d ow n to thei r origi na l st a tus of

ra w m a te ri a ls wh ich a re rest ored to th e s o il an d th e ,

ene rgy i nv olved i n thei r m a nuf a ctu re is once m ore

rele a sed .

T H E S OLI D N I TR O GE N OU S W A S TE S c onsisti ng of p ro ,

tei ns a mi no a cids a nd the i nte rmed i a te p roducts u nde r

,
-
, ,

g o successive dec o mp o siti o ns by th e c orresp ondi ng ba c

te ri a whi ch a re nou rish ed the reby a mm oni a a nd 0 0
n
, 2

bei ng the end p roducts I the c o u rse of such dec omp osi k
.
-

ti ons off e nsive od ors ma y be p roduced He nce the na me .

put refa cti onis a pplied to them B u t whethe r th e p rocess .

r esults i nthe p ro ducti onof evil smells or n ot th e results ,

a re the s a me the rele a se o f e n

,
e rgy a nd th e p roducti onof
simple r a nd S imple r c omp ou nds u ntil the residue fi na lly
a ppe a r s a s a mm on i a a nd v a ri o us sulph a tes a nd ph o s
p h a te s .

P UTR E FA CTI O N I s BROU GH T AB O UT chiefly by b a cte ri a , ,

a ided to s ome exte n t by fu ngi Th e number of ki nds of .

b a cte ri a a cc omplish ing these ch a nges is ve ry l a rge a nd ,

th e ex a ct chemic a l re a cti ons p roduced by e a ch a re a s yet

on ly impe rfectly know n While b a c te ri a requi ri ng f ree
.

oxyge n (a e ro bes ) a re a ble to dec omp o se p ro tei n s this ,

a bility is sh a red by ce rt a i nspecies o f b a cte ri a B a cil

lus b o tul i nus ) which ca nbe a ctive only i nth e a bse nce of
free oxyge n (a na erobes ) M a ny of the f orme r ca nc a rry
.

the p rocess to completi on gIvmg ri se to C O a nd a m , Z

m oni a (N B ) Th e a na e robic b a cte ri a c a nno t usu a lly

S .

c a rry dec omp ositi ons so fa r a nd th e two s orts form ,

i nte rmedi a te p roducts which a re m ore or less di ffe re nt .

P UTREFY I N G S U B S TA N CES a re o f m a ny s orts a nd from

m a ny s o u rces — me a t milk eggs seeds a re f a mili a r
, , , ,

m a te ri a ls wh ich pu t refy Comp o u nds which u nde rg o

’

a ctu a l put ref a cti on a re p ro tei ns a nd thei r p rim a ry de

ri va ti v es F rom th ese p roteins m a i nly by b a cte ri a l
.
,

a ctivities th o ugh fu n,
gi ma y a ls o t a ke p a rt the a mi no ,

a cids a re f ormed which ma y i n tu rn be b ro ke nd ow n

,
to
still simple r c omp ou nds .
26 8 D E COMP O S ITI O N
TH E P R ODU CT S P UT OF Wh ile the end RE FA T C ION .
-

p roduct s C O2 a nd H2 0 a nd a mm on i a a re re a dily e nough

rec og n ized the re a re a g re a t m a ny i nte rmedi a te p roducts
, ,

m a ny o f which a re Only temp ora ry The first p roducts .

a re simil a r to th o se p roduced i n digesti o n a n d in clude ,

pept ones a nd a lbum oses wh ich a re d ec omp osed i nto ,

th e a nim o a cids A numbe r o f g a se ous c omp ou nds ma y

-
.

a l s o be f o rmed a m on ,
g which a re meth a ne (C H ) a nd 4

hyd rogen sulph ide (H S ) Bubbles o f m eth a ne a nd

z .

sulph u retted hyd roge nma y ofte nbe see nrisi ng f rom th e
muddy b ott oms o f sw a mps a nd sulph u retted hyd roge nis ,

give noff by rotte n eggs a nd by th e dec a yi ng rem a i ns of

pl a nts a nd a nim a ls a ccumul a ted i nth e mud a t th e b o tt om
o f S l o w ru n
-
ni ng p olluted st re a ms a nd i n p onds S ome .

times H a nd N a re a ls o give noff Th e v ol a tile c omp ou nds .

which ch a ra cte rize put ref a cti oni nclude me rc a pt a n S k a t ol , ,

an d in d ol subst a nces th e c omp ositi onof which h a s bee n

,

f a i rly w orked ou t ; bu t u nd oubtedly in a dditi onto these ,

an d a ss oci a ted with p ut rid od ors a re o the rs no t yet k n o wn ,

which give th e ch a ra cte ristic smells to th e p ut ref a cti ons

o f diffe re n t m a te ri a ls .

P TO M A I N ES f orm one o f th e m ost i nte resting groups

o f deg ra d a ti o n p roducts f ro m p ro tei n They a re s i mple .

c omp ou nds of C H O a nd N p roduced by b a cte ri a l

, , , ,

dec a y i n f ood Th ey resemble th e a lk a l oid s u sed in

.

medici ne a nd include s ome of th e m os t vi ole nt p ois ons

k now n B a c te ri a l t oxi ns on th e o th e r h a nd a re ch a ra c
.
, ,

teri sti c met a b olic p rod ucts o f ce rt a in species o f b a cte ri a ,

an d a re l a rgely indepe nde nt o f th e f ood up onwh ich they

a re livi n g .

T H E P UTR E FA CT I O N P ROCES S ma y be best illust ra ted

by a specific i nst a nce Thus we ma y disc ove r th e b ody of
.

a co w rece n tly d e a d lyi ng onth e g ro u nd This b ody con

,
.

f a in ed a t th e time o f de a t h m a ny b a cte ri a withi nits d iges

tive t ra ct a nd it be comes i nfe cted wi th o the r d est ructive
,

b a cte ri a none of wh ich ca nbe held i nch eck by the de a d

,

o rg a n ism a s they w o uld h a ve bee nh a d the b ody c ontinued

'
 S U MM AR Y 26 9

to live Th e a ctivities of these b a cte ri a resul t i nth e p ro

.

ducti onof e nzyms wh ich c a rry onhyd rolysis th e split

ting of p ro teins i nt o simple r org a nic nit roge nous com
p o u nds like th e a lbum oses a nd pep t ones — which a re
,

themselves b roke nd ow nby o th e r e nzyms to a nim o a cids -

.

The put refyi ng c a rc a s s fu rth e rm ore bl oa ts f rom the p ro

, ,

ducti on of g a se ous subst a nces withi n the c a vities o f the

b ody a nd th e se ries of dec omp ositi ons i ncludes th e p ro
,

ducti onof th ose offe nsive od ors wh ich i ndic a te to us th e

p resence of a de a d a nim a l As th e b ody of th e a nim a l
.

b re a ks d ow nmech a nic a lly a s well a s ch emic a lly th e fi na l ,

p roducts of dec a y a re libe ra ted esc a p ing i n t o the a i r or

,

w a sh ing int o the s oil thus b e comi ng once m ore a v a il a ble

,
‘

a s ra w m a te ri a l s f or th e m a n ufa ctu re of f ood by pl a nt s .

S U M M AR Y — .F rom the illust ra ti ons which h a ve been

empl oyed it sh o uld be cle a r th a t the dec omp ositi on of
ca rbo h ydra tes a n d fa tty su bs ta nces c onsists of successive
ph a ses Th ese c onsist i np rocesses resembli ng th ose which
.

go on i n the pl a n t or a nim a l b ody a nd p rope rly te rmed

digesti on Th e p roducts of di gesti on wh eth e r i nside the
.
,

living b ody or i n exc ret a or i n l ifeless r em a i ns ma y ,

be oxidiz ed th u s libera ti ng e ne rgy wh ich ma y be

,

used by the a ctive org a nisms The fi na l result i n eve ry .

c a s e so fa r i nvestig a ted is the libe ra ti on o f the e ne rgy

origi n a lly st ored by ph o t o sy n thesis or by othe r met a b olic
p rocesses a nd th e setti ng f ree of the C i n th e form
,

of CO a n Z d H i nthe f orm o f H O Th us th rough a l l the Z .

multifa ri ous t ra nsf orm a ti ons of subst a nce the re h a s bee n

no l o ss of m a te ri a l or o f e ne rgy The free e ne rgy of th e
.

su nligh t h a s bee nst ored up i norg a nic c omp ou nds a s p o

tenti a l e ne rgy to be libe ra ted eithe r by the org a n
,
ism
which st ored the e ne rgy or by o the rs i n thei r v a ri ous
a ctivities .

Th e p roteins a re b roke n d ow n i n th e a nim a l b ody

itself by digesti on etc a nd i nth e lifeless rem a ins of a ni
,
.
,

m a ls a nd pl a nts by the a i d of othe r org a nisms m a inly

, ,

b a cte ri a a nd fu ngi Here a ls o th e subst a nces a nd th e

.
27 0 D E COMP O S ITI O N

e ne rgy used i n f ood m a nu fa c tu re a re retu rned i nuseful

f orm a g a i nto the a i r a nd the s oil f rom which th ey we re
t a ke n Inth e ord i na ry c ou rse of na tu re the re is little or
.

no distu rb a nce of th e b a l a nce i n a give n a re a . Only

wh e re ma nor fi re or s ome o the r a ccide nt (v olc a nic ) in
terv en es is the re a n y c on side ra ble de p a rtu re f rom th e
regul a r cycle.
 C HAPTER XX IV

T HE FE R TILI T Y OF T H E S O IL

TH E EX H AU S T I O N OF T H E S OI L
P LA N T S w o uld o ccu r
BY

i n a c omp a ra tively sh ort time u nless the re we re s ome

wa y to repl a ce the subst a nces wh ich they rem o ve Ina n .

e a rlie r ch a pte r i t wa s S h ow nby che mic a l a na lysis th a t th e

b odies of pl a nts c ont a i n the eleme nts c a rb on hyd roge n , ,

oxyge n n,
it roge n ph o sph orus sulphu r p ot a ssium ma g
, , , ,

nesi um c a lcium i ron silic on s odium a nd chl ori ne It

, , , , ,
.

wa s fu rthe r dem onst ra ted by th e w a te r cultu re meth o d

th a t a ll of these excep t th e l a st th ree a re i ndispe ns a ble to
g reenpl a nts All of th em excep t c a rb ona nd oxy ge na re
.

de rived enti rely from th e s oil i nw a te r s oluti on S uit a ble .

s oluble s a lts o f nit roge n (nit ra tes )? ph osph orus (ph os

p a tes ) a n
h ,
d p ot a ssium a re sc a rce i n m ost s oils Hence .

thei r p romp t r etu rnto i t ina f orm suit a ble for use is a
p rime requisite i nma i nt a i ning the fe rtility of the s oil .

N ATUR A L F ERTI LI T Y — In a st a te of na tu re th e com

.

p ou nds rem oved from th e s o il a re a fte r a l onge r or S h orte r

,

s o j ou rn i n the b odies o f pl a nts or a nim a ls usu a lly re ,

tu rned to it i nth e s a me vici nity whe re they a re a cted on

,

by a se ries of m i croorg a ni sms a nd fitted for use a g a i n .

U nde r such c onditi ons the re is no exh a usti on of fe rtility ,

but the re ma y a c tu a lly be a ni nc re a se of it by me a ns to ,

be expl a i ned p rese ntly .

FERTI LI Z ER s — U nde r cultiv a ti on it usu a lly h a ppe ns

.

th a t th e c rop s a re rem o ved f rom th e fields a nd little or

none of th ei r subst a nce retu rned The l oss thus sust a ined
.

must be m a de g ood by the a pplic a ti ono f fe rtilize rs By

m
.

this te r th e a gricultu ra list me a ns Comp ou nds or mix

tu res which c ont a in the eleme nts i n which th e s oil is
27 1
27 2 THE F E R TILI T Y OF T H E S O IL

deficie nt Th e rem a ins of pl a nts a nd a nim a ls a nd th ei r

.

w a stes h a ve bee nused a s fe rtilize rs f rom time immem ori a l

an d a re na tu ra l ly a m ong the best since they c ont a i nth e ,

ve ry elements which we re rem oved f rom th e s oil .

C e rt a i n dep osits o f rock a re rich i nph osph a tes wh ich

ca nre a dily be m a d e a v a il a ble for pl a n t use Cy a na mid .
,

a w a s te p rod uct o f ce rt a i nm a n uf a ct ories c ont a ins a v a i l ,

a ble n it roge n G u a no is the exc reme nt o f ce rt a i n sea

.

bi rds which h a s bee ndep osited for a ges on s ome isl a nds
off th e P a cific sh ore o f the ra i n less regi on of South
Am e ric a It is rich i n nit roge n c omp ou nds S odium
. .

nit ra te or Chi li s a ltpete r i s mi ned i nnorth e rnCh ili a nd

, ,

sh ipped to E u rope a nd th e U nited S t a tes P o t a sh com .

,

p osed of p ot a ssium c omp ou nds is exte nsively mi ned i n ,

G e rm a ny a nd is obt a i ned to a less ex te nt i n th e weste rn

U nited S t a tes D u ri ng th e wa r it wa s secu red f rom sea
.

weeds Nit roge n comp ou nds a re now bei ng ma nu fa c

.

f u red f rom the f ree nit roge n o f the a i r by th e use o f

elect ricity .

A N A L W A S TES
I M — In p revi o us ch a p te rs i t h a s bee n
S h o w nth a t s ome o f th e c a rb ohyd ra te f oods a re used i nth e

f o rm a ti onof p roteinby pl a nts Th e l a tte r subst a nces a re .

used chiefly to build new p rot opl a sm or to rep a i r th e

w a st a g e of m a tu re cells Once a p a r t o f the livi ng ma
.

ch ine i t is rel a tively fixed bu t no t a bs olutely so Inth e

, ,
.

ve ry p rocess o f livi ng p ro to pl a s m p roduces ce rt a inw a ste

,

c omp o u nd s c ont a i ning nit roge n which a re exc reted f rom ,

th e b ody i n th e u ri ne Th e l a s t ch a pte r desc ribed th e

.

ch a n ges i nduced i n these by b a c te ri a wh e reby th ey a re ,

b ro ke n d ow n to a mm oni a (N H ) c omp ou nd s , .

M I CROORGA N I S M S Th e re a re m a ny ki nds o f b a cte ri a

.

(Fig . fu ngi a nd sm a ll
,
a nim a ls i n th e s o il S ome .

b a cte ri a a re c once rned wi th fe rme nt a ti ons a nd p u trefa c

ti ons but seve ra l a re di rectly c once rned wi th ch a nges i n
,

th e c o m p o u n ds o f nit ro ge na nd sulph u r wh ich h a ve to do

wi th s oil fe rtility For p rese nt p u rp o ses th e f oll owi ng
.
,

will be disc ussed : (1 ) Th e ni trifyi ng b a cte ri a ; (2) Th e

 T HE NIT R IF Y I NG B A CT E R IA 27 3

nit roge nfixi ng b a cte ri a ; (3 ) Th e de nit rifyi ng b a cte ri a ;

-

an d (4 ) Th e sulph u r b a cte ri a .

T H E N I TR I FY I N G B A CTER I A The re a re two sep a ra te

an d disti nct s orts of b a cte ri a c once rned i n ch a ngi ng a m
m oni a c omp o u nds i nt o us a ble nit ra tes Th e fi rst ki nd ca n .

utilize a mm oni a a nd oxidize it to nit ro us a cid c omp o u nds

c a lled nit rites The sec ond set ca n m a ke no use of a m
.

m oni a comp ounds7 b u t must h a ve nit ri tes wh ich th ey

’

F FI G 6 3 . . S o il B a cte ri a . Fi g 1 , A z oto b a cter, a m o til e b a cterium

.

li vi n g i nth e s oil whi ch is a bl e to fix th e f ree ni tro genof th e a tm os

p h ere a n d l ea v e i t b e hi nd a f te r i ts d ea t h in su c h a f o r m th a t i t ca n
ev en t u a lly b e us e d by hi gh er pl a n ts Fi g 2, Tubercl e b a cter ia , whi ch
. .

inh a bi t the s o il a nd inva de th e roots of cl over a nd o th er l e gumi nous

pla nts , ca using th e pl a nt to f orm tu bercl es inwhi ch th e b a cteri a live .

Th ey a ls o fix free ni tro gen a nd th ereby enri ch th e s oil Fi g 3 , . .

S ulphu r b a cteri a living i ns o il w a ter Th ey a re a bl e t o u tiliz e th e ga s ,

.

hydr ogensulphi de, a s a s ou rce of energy by o xi di zi ng fi rs t th e hydro gen

to f orm w a ter a n d l a ter th e s ul ph ur to f orm co mp o u nds of sulph ur i c
ac id .

o xidize to nit ra tes being thus depe ndent

, n th e fi rst
o

s ort .

T H E N I TR I TE B A CTER IA The re a re seve ra l di ff e re nt

.

S pecies o f s o il b a cte ri a wh ich ca n ch a nge a mm oni a com

p ou nds i nt o c omp o u nds of nit rous a cid c a lled nit rites .

Th ey a ll seem to h a ve e sse nti a lly th e s a me m ode of l ife .

This is i ndic a ted i n a s imple wa y by th e f ormul a :

N H OH (a mm oni a )
4 20 (o xyge n) HN O (nit rous
2 z

a cid 2H O (w a te r ) z Of c ou rse th e a ctu a l ch emic a l

.

p roces ses a re m ore c omplic a ted bec a us e th e a mm oni a ma y

‘

occu r i n th e s o il i n o th e r c omp ou n ds e g a mm onium ,

. .
,

c a rb ona te (N H ) C O The re a cti on i s p resum a bly

4 2 g.

b rough t a b out by th e a cti on of a n oxidizi ng enzym

27 4 T HE FE R TI LI T Y OF THE S O IL

f ormed by th e b a cte ri a I t is a l s o one wh ich yield s e nergy

.
,

which th e o rg a nisms u se to ch a nge C0 or s ome c a rb ona te

2

Ca COa c a lciu m c a rb ona te ) int o such c omp o u n

, ds a s
a re useful to th em i nc a rry i n g onth eir life a ctivities .

T H E N ITR AT E B A CTER I A c a nn o t use a mm o n i a or its com

p ou nds bu t a re rest ricted to the nit ri tes f orm ed by th e
,

o rg a n isms d iscussed i n the l a st p a ra gra ph P resum a bly .

by me a ns o f a ne nzym th ey ox idize nit rous a cid a nd its

s a lt s to nit ric a cid a nd i ts s a lts s omewh a t a s f oll ows :
2HN O (n 2 it ro us a cid ) 0 (oxy ge n)
2 2HN O (nit ric s

a cid )

N I TR I FY I N G ORGA N I S M S AN D H I GH ER P LAN T s Th e .
-

c ont ra st betwee n these two sorts o f nit rifyi ng b a c te ri a

’

onth e on e ha n d a nd th e h ighe r pl a nts a nd o the r b a cte ri a

o n th e o the r is a st riki n g a nd imp o r t a nt one Highe r .

pl a nts de pe nd on c omplex c a rb on c omp ou nds for th e

e ne rgy to c a rry o nth ei r li fe p rocesses
~

met a b olism )
m
.

They u se s o eo f this e ne rgy to t ra nsf orm simple nit roge n

c omp o u nds like nit ra tes i nt o th e v a stly c omplex o n es
wh ich c onstitute a n imp ort a nt p a rt o f th ei r p ro t opl a sm .

Th ese b a cte ri a do j ust th e opp osite Th ey ca n use only

.

si m ple c omp o u nds o f c a rb on like CO z This th ey build

.

int o such c omplex f o rms a s th ey requi re by th e e ne rgy

d e rived f rom o xidizi ng nit roge n c omp o u nds F urth er .

m ore th ey le a ve th e nit roge n i h j ust th o se c omp ou nds

,

wh ich a re requi red by th e highe r pl a nts .

E E C S OF C
FF T U LT I V AT I ON — .B o th s orts of nit rifyi ng
b a cte ri a w ork best i ns oils with a n a lk a li ne re a cti ona nd
a re i n j u red by we t a cid s oils For th is re a s onthey usu a lly
.

th rive u nde r cultiv a ti on bec a use this te nd s to secu re

p rope r a e ra ti ona nd suit a ble dist ributi ono f m oistu re a nd ,

p reve nts the f orm a ti on a nd a ccumul a ti on of inj u ri ous

a cids .

T H E N ITR OGE N F I X I N G B A CTE R I A a re so c a lled bec a use

-

th ey t a ke f ree a tm osph e ric nit roge n a nd u nite i t w ith

hyd ro ge na nd oxyge n f ro m w a te r to f orm comp lex com
p o u nds Th i s rea cti on is one i n wh ich e ne rgy is st ored
. .
 FR E E —LIVI NG S O IL B A C T E R IA 27 5

up Th e b a cte ri a de rive th ei r e ne rgy to do this a nd to

.

c a rry ona ll thei r o the r met a b olism from the oxid a ti ono f
c a rb onc omp ou nds such a s sug a r whic h o c cu r i n the s oil .

These a re de rived chiefly fro m the rem a i ns o f pl a nts con

sisti ng l a rgely of cellul o se a nd simil a r m a te ri a ls .

Th ro ugh the a ctivities of these b a cte ri a a ctu a l a dditi ons

of a v a il a ble n it roge n c omp ou nds a re bei ng m a d e to th e
s oil I t is i nte resti ng to note h ow these nit roge nfi x i n
.
g
-

org a n isms suppleme nt th e w ork of th e nit ri fyi ng b a cte ri a

‘
.

Th e l a tte r a re e nga ged i n ch a ngi ng a l re a dy existi ng

c omp ou nds of nit roge n to S imple nit ra tes a nd ca n
not do thei r w ork i nthe p rese nce of s oluble c a rb ohyd ra tes .

Th e f orme r a l s o f orm nit ra tes but th ey do i t by fixi n

, g
free nit rogena nd must h a ve suit a ble org a nic f oo d to do so .

And s o they a re m ost efli ci ent j ust whe n the nit rifyi ng
b a cte ri a a re le a st efficie nt .

A c onside ra ble v a riety of s oil org a nisms ca n fix nit ro

g e n : ( )
1 F r ee livi n
-
g s o il b a cte ri a (2) S oil b a cte ri a livi ng
,

S ymbi o tic a lly with a lg ae (3 ) B a cte ri a livi ng symb i oti

,

c a lly with ce rt a i n legumi nous pl a nts i n roo t tube rcles ,

(4 ) C e rt a i nfu ngi c a lled M yc orrhiz a , livi ng symbi otic a lly

onth e roo ts o f a n umbe r of species of t rees a nd sh rubs .

FREE LI V I N G S OI L B A CTER I A a ble to fix nit rogen a re

-

nume rous a nd o f seve ra l d iffere nt ki nds (Az ot ob a cte r ,

Gra nu l ob a cter Amyl ob a cte r C l ost ridium )

, ,
Alth ough
.

they a re not equ a lly nume rous i na ll s oils they seem to be ,

a lm o st u n ive rs a l i nth ei r dist ribu ti on P rovided by culti

.

v a ti onwith suit a ble c onditi ons of g rowth , they will mul

ti p l y a n d e nrich th e s o il i na lm o st a n y p a r t o f th e w o rld .

The b odies of th ese b a cteri a a re s a id to be rich e r in

‘

nit roge nth a na ny o the r livi ng thi ngs so f a r i nvestig a ted .

B A CTER I A S Y M B I OTI C W I T H AL GZE In th e su rf a ce

.

l a ye rs of t h e s o il th e re is e no ugh ligh t for s ome l owly

a l gm They o f c o u rse fix c a rb onby ph o t osynth esis a nd
‘

.
, ,

need nit ra tes It is s a id th a t s ome s orts of nit roge n

fix i ng b a cte ri a f orm a symbi o ti c a rra nge ment with them
.

i nwh ich the a lg ae receive nit ra tes i nretu rnf or the sug a rs
27 6 T HE F ER TILI T Y OF T H E S O IL

or o the r s oluble c a rb on c omp ou nd s wh ich the b a cte ri a

a b sorb f ro m th em Th e ex a ct f a ct s a ppe a r no t yet
.

to be d ete rm i n ed i n respect to th e rel a ti on exist i ng b e

twee n these cl a sses Of org a nisms I t h a s bee n rece ntly .

rep orted th a t s ome o f th ese a lg ae ca n fix th ei r o wn

nit roge n .

T U BER CLE B A CTER IA I t is a widely know nf a ct th a t

.

cl ove r a nd s ome o the r legumi nous pl a nts e nrich th e s oil .

Th i s b i t of knowledge h a d p a ssed i nt o c omm on sl a n g in

”
~

th e ph ra se too p o or to grow be a n
,
s l ong before its ,

scie ntific expl a na ti onh a d bee nw orked ou t Onth e roo ts .

o f such pl a n ts the re occu r sm a ll swellings v a ri ously c a lled

nodules or tube rcles I nside th em a re f o u nd nume rous
’

b a cte ri a of a speci a l s ort Wh e n th e pl a nt is y o u ng its

.

roo ts a re a tt a cked by th e b a cte ri a which pe n et ra te i nt o

them th ro ugh the delic a te root h a i rs a nd est a blish them
selves i nth e oute r l a ye r of root cells Th e roo t re a cts by
-
.

multiplyi ng th ese cells i nnumbe r the reby f orming a s ort ,

o f tum o r like o u tg ro w th i n wh ich th e b a cte ri a a re con

-

fi ned He re th ey fix nit roge n de riving th e e ne rgy to do

m
.
,

so f rom th e c a rb on c omp o u n ds obt a i ned f ro th e root

cells Inretu rnthe roo t uses thei r nit roge nc omp ou nds
. .

M Y CORR H I Z A a re m a sses of fi ne hyph ae or fil a me nts

o f ce rt a i n s orts o f fu n gi Wh ich a tt a ck th e roots o f s ome
k inds of t rees a nd sh rub s Th ey p e net ra te th e roo t a nd
.

t a ke f ro m i t org a nic c a rb on c omp ou nds O utside th ey .

,

ra mi fy th rough the s o il Th ey h a ve th e p owe r to fix nit ro

.

g e n
,
wh ich they sh a re with th e roo ts This a rra n ge .

me nt a ppe a rs to be so s a tisfa ct ory to b o th p a rt ne rs th a t

s ome o f th ese t rees no l onge r eve n f orm root h a i rs but
a bs orb a ll th ei r w a te r a n d sa lts th ro ugh th e fu n gus .

Th is a rra ngeme nt illust ra tes a no the r i nte resti ng a d a pt a

ti onof org a nisms to th ei r e nvi ronme nt ; fo r m o st o f these
t rees g row i n s oils wh ich a re p oor i n nit ra tes a nd a ls o
u nfit for th e g rowth o f th e nit rifyi ng b a cte ri a .

D EN ITR I FY I N G B A CT ER I A a re th ose which reduce

nit ra tes i e b re a k up th e c omp o u nd a nd libe ra te the
,
. .
,
 S UL P H U R B A C T E R IA 77

nit roge na s a ga s Obvi ously from th e p oi nt of V iew of

.
,

c onse rving s oil fe rtility th ey a re u ndesi ra ble Fortu , .

na tely they th rive best i n wet a nd p oorly a e ra ted s oils

an d do no t do much d a m a ge i np rope rly cultiv a ted s oils .

D ra i na ge a e rati on a nd a ddi ti onof lime to neut ra lize th e

, ,

a cids a re th e usu a l me a n s of h oldi ng them i n check a s

well a s p rom o ti ng th e a ctivity of th e nit rifyi ng org a nisms .

S U L P H U R B A CTER I A S ulph u r is t a ke n f rom th e s oil

.

i ns oluti ona s a c omp o u nd of sulphu ric a cid (H S O ) a n d Z 4

built int o th e p ro teins a nd s ome o the r c omp ou nds of

pl a nts a nd a nim a ls I npu t ref a cti onit is c omm only libe r
.
-

a ted a s hyd roge n sulph ide (H S ) ga s f a mili a r a s the Z ,

o d or of ove r rip e eggs Inthis f orm it is us a ble by types

-
.

of s oil a n d w a te r b a cte ri a wh ich oxidize the hyd rogen to

form w a te r a nd st ore the sulph u r inthei r b odies Whe n .

the s upply of hyd roge nsulphide ru ns S h ort they oxidize

this st ored sulph u r to form sulphu ric a cid a nd its
sulph a tes .

T H E CARB O N CY CLE I t is cle a r from wh a t h a s been

.

s a id i n this a nd p recedi ng ch a p te rs th a t a lm ost th e only

s ou rce of th e c a rb onof org a nic c a rb onc omp o u nds is th e
CO of th e a i r
2 This is c ombi ned with H O by g reen
.
z

pl a nts to f orm f oods a nd fuels Th e se i n tu rn a re used .

by ne a rly a ll living th i ngs a s thei r s o u rce of e ne rgy I n .

thei r met a b olism they oxidize c a rb ohyd ra tes a nd f a tty

foods to C O a nd H O Th u s th e c a rb onwithd ra w nfrom
Z Z .

th e a i r a s C O by th e g ree npl a nts i nth e p rocess of ph o t o

2

synthes i s i s eve ntu a lly retu rned to i t whe n the f ood is

o xidized by th e p ro t opl a sm o f s ome a n im a l or pl a nt .

Fuels a re a ls o ox idized i nst oves a nd fu rna ces a nd yield ,

CO a n d H 0 Th e s a me c a rb onh a s bee n i n this ma n

“

Z 2 .

ner used ove r a nd ove r a g a i nby successive ge ne ra ti ons of

org a n isms (Fig .

TH E N I TR OGE N C CLE like th a t o f c a rb on p e rmits th e

Y , ,

s a me nit rogento be used repe a tedly S t a rting with the .

a bu n d a nt supply i nth e a tm osphe re it i s fi rst m a d e a vail ,

a ble by n it rogenfi xing org a nisms Accumul a ti ng i n the

-
.
 ‘

27 8 T HE F E R TILI T Y OF T H E s oi L

s oil by the a ctivities o f th ese org a nisms the nit ra tes a re ,

t a ke nup by gree npl a nts a nd c ombined with sug a rs a nd

sul ph u ric a nd ph os p h oric a cid c omp ou nds to f orm a mino ,

a cids a n d p ro tei ns These a re used to f orm livi n

. g p rot o
pl a sm or to rep a ir its w a stes Whe n de a d p rot op l a sm
.
,

re e nPl a n
ts Ani ma ls

u
P tre fa ctio nof

b
De a d od ie s of

a nts

Ni tra te s +
p h a te s e
Ph o s p h a te s

The S oil
FI G 64
. .
—A
D i a gra m to sh ow s om e o f th e tra n s f o rm a ti o n
s of

ca rb o n a n d ni tro ge n i n th e cou rs e o f th e ir u ti liz a ti o n by livi ng or

ga n i s ms . N o te t h a t th e a i r co n s ti tu tes the grea t so ur ce o r rese rvo ir
f rom whi ch bo th ca rb ona nd ni tro gena re dra w n by th e m e ta bo lis m
o f pl a n ts N o te fur th er th a t o n
. ly gree n pla nts ca n wi th dra w the
ca rbo nf ro m the a i r to m a n uf a ctu re suga r a nd o nly certa i ns o rts o f
b a cte ri a a nd o th er low f orms of pl a nts ca n u tili z e f ree a tm osph eri c
ni tro ge n All a n
. i m a ls d e pe nd o nth ese pla nts di re ctly o r i ndi re ctl y
for th e ir us a bl e co mpo unds o f ca rbo na n d ni tro gen T h e su nis the
.

origin a l so u rce o f t h e e n e rgy u se d to sy n th esi ze suga r a n d o th e r

o rga n i c co m po u n ds of ca rb on a nd ni tro gen In the v a ri o us t ra ns
.

forma ti o ns whi ch th ese compo u nds u nd ergo u ntil fi na lly relea s ed a s

ca r bo ndi o xi d , w a ter, n i tra te s , e tc , a ll the energy whi ch wa s ori gi na lly
.

l ock ed u p i nth em is a ga i n f reed i nthe f orm of w ork or h ea t .

and it s w a stes a re bo th dec omp osed by p ut refa cti ona nd

yield a mm oni a which i nits tu rn is oxidiz ed to nit ra te
,
.

These ma y be eith e r used by g ree n p l a nts a g a i nor re

 P H O S P H OR U S C Y CL E S 27 9

duced to g a se ous nit roge n by the de nit rifyi ng b a cte ri a

an
,

d so c omplete the cycle .

S U LP H U R AN D P H OS P H OR U S CY CLE S I t h a s a l re a dy
.

bee n expl a i ned th a t sulph u r t a ke n up by pl a nts a s sul

p h u ri c a cid c o mp o u nds is built int o p ro t o pl a sm an d re
a ppe a rs du ri n g p ut ref a cti on of de a d pl a nts a nd a nim a ls
a s th e m a l od orou s ga s hyd roge nsulphide Th is i ntu rn
, .

is eve ntu a lly rest ored to t h e so il a s sulph a tes by th e

a cti on of th e sulph u r b a cte ri a Ph osph orus ente rs the
.

pl a nt a s a ph osph a te a nd bec omes a p a rt of the ce lls On .

thei r de a th a nd dec omp ositi oni t rea p p ea rs a s ph osph oric

a cid a n d u nites with lime or o th e r e a rthy m a te ri a l to
f orm ph osph a te a g a in .

T H E E N ER GY C Y CLE di ffe rs from the m a te ri a l cycle

'

j ust discussed i n th a t it is not a cl osed one so fa r

a s th e e a rth a l on e is c once rned Th e e ne rgy f or ph o t o
.

synthesis c omes from th e su n i n th e f orm of ligh t It .

is l ocked up inf ood a nd fuel Wh e nlibe ra ted by meta b o

.

l i sm or c ombusti oni t r ea pp ea rs i n th e f orm o f ch emic a l

e ne rgy he a t or mech a nic a l w ork In these f orms it is
, ,
.

eventu a lly dissip a ted i nt o s p a ce or st ored up i n o the r

chemic a l c omp o u nds or i n e ne rgy of p ositi on wh e n ex
,
,

pe nded i nra isi ng a weigh t for ex a mple In no c a se is

,
.

it a g a i nu s a ble di rectly by g ree npl a nts In th e c a se of .

fuel bu rned to runa nelect ric ligh t pl a nt s ome of i t c ould ,

be used a g a i n Wheth e r th a t wh ich esc a pes i nt o sp a ce

.

eve r retu rns a s ligh t is a q uesti onnot a nswe ra ble i nthe

p rese nt st a te of knowledge .
 PAR T II

S ECTI ON 8

GR OWT H AN D R E P R OD UC TI ON
 C HAPTER XXV

CE LL—DIVI S I O N

T H E GR O WT H OF OR GA N I S M S . In e a rlie r
ch a p te rs i t
wa s sh ownth a t livi ng thi ngs g row by the t ra nsf orm a ti on
o f ce rt a i n s orts of f oo d in t o thei r livi ng subst a nce a nd ,

th a t c omplex org a nisms a re c omp o sed of ve ry nume ro us

cells h ighly speci a lized to pe rf orm thei r p a rticul a r fu nc
m
ti ons And fu rth e r ore it wa s i nd i c a t ed th a t a ll the dif
.

ferent s orts of cells i n a c omplex org a nism a rise fro m

a si n gle orl gi na l one by divisi on Growth of a high e r a ni
.

ma l or pl a nt thus c onsists of th ree th ings : (a ) th e ma k

i ng of new p ro t opl a sm wh e reby th e i ndividu a l cell be
c omes l a rge r (b ) the divisi onOf th e cell whe nit a tt a i ns
,

its a pp rop ri a te size a nd (c) th e g ra du a l speci a liz a ti on

,

o f diff e re n t cells to pe rf orm th ei r i ndividu a l fu ncti ons .

CE L L G R O WT H N D V S O N
A D I I I — Assumi ng th e a bility
.

of p ro to pl a sm to t a ke f oo d of a suit a ble s or t a n d a ss u m
l a te it i nt o its own subst a nce a nd t he reby i nc re a se th e
a m ou n t of livi ng cell c ontents ,it is now desi red to di rect
a tte n ti onto th e f a ct th a t cells h a ve a defi nite limit a ti on
to th ei r in cre a se in size .Whe n a cell re a ch es th e size
ch a ra c te ristic of th e p a rticul a r pl a nt or a nim a l of which
it i s a p a rt it eithe r ce a ses to grow or it divides V e ry
,
.

little is k now n resp ecti ng th e mech a nism which dete r

mines wh a t th is size must be On th e c ont ra ry the a d
.
,

v a nt a ges O f size lim it a ti on a re obvi ous S i nce cells t a ke

.

in f ood a n d exc rete thei r w a ste th ro ugh thei r su rf a ces ,

it f oll ows th a t the ra te of this exch a nge w ill be dete r

mi ned by th e exte nt of the su rf a ce The su rf a ce of a cube
.

or b a ll in c re a ses a s the squ a re of th e di a mete r (Fig .

The v olume i nc re a ses with th e cube Thus d o ubli ng the

.
284 CE LL -
DI VI S I O N
di a mete r of a sphe ric a l cell w o uld multiply its a re a fou r
times th e sq u a re of 2) a nd its v olume eigh t times
th e cube of Th e result w ould be th a t th e v olum e
or a m o u n t of p rotopl as m to be supplied th ro ugh th e sur
f a ce o f the cell w o uld h a ve i nc re as ed twice a s much as th e
su rfa ce th ro ugh wh ich it is to be supplied Cle a rly fur .
,

the r inc re a s es in size w o uld fina lly re a ch a p o i nt whe re

the v olume of p rotopl a sm in th e cell c o uld not be sup

FI G 6 5
. .T h e C omp a ra tiv e S urfa ces of S o lids T h e edge o f the
.

la r ger s oli d is 2 i n onea ch sid e I t t h e re ore conta i n

. . f s 8 cu in E a c h
. .

side is 2 x 2 i n a nd h ence 4 sq ; in in a rea T he wh ol e a rea is thus

. . .

24 sq in A cube o f 1 i n onth e sid e , a s e g , th e o n

. . . . e i n th e uppe r
.

ri gh t corn e r o f t he l a r g e r o ne , a , B , c, d , e, f , g, it , con ta i ns 1 cu in . .

an d h a s a su rf a ce of 6 sq i n Fro m th es e fi gu res it is cle a r th a t

. .

d oub ling the sid e i ncrea s es the a rea 4 times bu t a t t he sa m e tim e

i ncrea ses the so lid conte nts 8 tim es Ino th er w ords the conte nts in
.

cre a se twi ce a s f a s t a s th e a rea Al th o ugh no t q ui te so o bvi o u s t hi s

.

f a ct is eq a ll y tru e o f a sph ere or a ny o th er sha pe of s o lid

u .

p lied th rough its su rf a c e i n a n a d equ a te m a nne r I t is .

not imp rob a ble th a t this s ta te o f a ff a i rs ma y a ct a s th e

stimulu s to divisi on Pl a nt cells usu a lly meet this situ a
.

ti on by formi ng a l a rge v a cu ole which reduces th e

v olume of p ro t opl a sm a cc ordi ngly As we sh o uld expec t .

f rom this f a c t pl a nt cell s a re onthe a ve ra ge l a rge r th a n

,

th ose of a nim a ls .
 M E CHANI S M OF CE LL D I VI S I O N 2 85

TH E M E CH A N I S M OF CE LL D I VI S I O N is ve ry much bet
ter u n de rst ood th a n th e stimulus which b ri ngs it a b out .

In the g re a t m a j ority o f cells divisi on begi ns with the

nucleus which u nde rg oes a c omplic ated p rocess known
,

u nde r a v a riety of na mes nucle a r divisi on mi t osis k a ry o

, , ,

kinesis This p ro cess ca nbe best studied i nth e g rowi ng

.

an im a l emb ryo or i n th e g rowi ng regi ons a t th e tips o f

roo ts (Fig 66 ) or stems. B ef ore we ca nu nde rst a nd th e
. .

FI G 66
. . from the Growing R e gi onof th e R oot T i p of a n
Cells
Oni on N o te ca ref ull y th e di fferent siz es a nd sh a pes o f the cell s
.

an d th a t th e ch ro m a t i n is i n a di ff erent s ta ge o f d ev el o pm ent i n
d iff erent cells .

p rocess it will be necess a ry to c onside r the st ruc t u re of

the nucleus .

T H E R E S TI N G N U CLE U S i nm ost c a ses c onsists O f a su r

ro u n di ng envel op e k now n a s th e nu cle a r memb ra ne a ,

semi flu i d sa p a netw ork of ch rom a ti n a nd a nucle olu s

-
, ,

(Fig. The m ost imp ort a nt p a rt a t le a st the m ost

,

pe rm a nent p a rt of it is th e ch rom a ti n Inthe cell which

,
.

is not i n a st a te O f divisi on th e ch rom a ti n a ppe a rs i n

st a i ned specimens to c onsist of i rregul a r th re a ds which
freque ntly i nte rsect Th ey a re of ve ry i rregul a r size a nd
.

s h a pe a n d a lm ost a lw a ys much th icke ned whe reve r th ey

286 C E LL—DI VI S I O N
j oi n one a nothe r C h rom a ti n is c a lled by th i s na me
.

( de rived f ro m th e G r eek w o rd f or c ol or ) bec a use its sub

st a nce c ol ors e a sily a nd be a utifully whe n v a ri o us dyes

o

a re a pplied to it The stu ff o f whi ch it is c omp osed is

.

a ve ry c o mplex n ucle o p rotei n or m ore likely a na ggreg a

-
,

t i on o f m a ny sligh tly di ff e re nt c omplex p rotei ns The .

nucle olus is a ls o a p roteinb ody I t a ppe a rs to be s ome .

FI G 6 7
. . A Typi ca l Pl a nt Cell from th e ro o t tip o f th e S pid er
,

w ort (T ra des ca nti a vir gi ni ca ) c w cell w a ll ; ogz cy to pl a s m ; n

. . .
, , ,

nu cl eus ; chr chr o m a ti n; ns nu cl ea r sa p ; n nu cl e olus ; v v a cu o l e

, . .
,
’
, ,

fill e d wi th w a ter a nd di s s olv e d subs ta nces .

wh a t less c omplex i nits ch emic a l st ructu re a nd is not a

pe rm a ne nt org a nof the nucleus I t i s th ough t by m a ny .

to be ra the r o f th e n a tu re o f a supply o f f ood for th e

g rowth o f th e ch rom a ti nth a na na ctu a l livi ng org a n I t .

is ce rt a i nly not a na bs olute ne cessity si nce m a ny nuclei ,

do no t p ossess nucle oli S till less is k nown a b ou t th e

.

ch emic a l n a tu re o f th e n ucle a r sa p It seems likely th a t .

it is a c oll oid a l s oluti ono f p ro tei ns bu t wh eth e r i t is liv ,

in g o r n o t is no t yet dete rmi n ed The nu cle a r memb ra ne .

is now know nnot to be a p a rt of the nucleu s p rope r but

me rely a th i nm emb ra ne p roduced by th e a cti on of the
nucle a r sa p onth e su rro u nd i ng p ro t opl a sm .

T H E B EH A V I O R O F T H E C H R O M AT I N i np rep a ra ti onfo r
an d du ri n g nucle a r divisi onis m ost i nte resting a nd imp or
 THE B E HAVI OR OF C H R O M ATI N 287

t a nt (Fig The fibe rs of the net bec ome diss oci a ted
.

from one a nothe r i n such a wa y th a t l ong th re a ds a re

f ormed S ometimes one c onti nu ous fth rea d a ppe a rs ; a t
.

o the rs a n umbe r of sep a ra te th re a ds a re formed They .

g ra du a lly el ong a te a nd bec ome sle nde r a nd u niform i n

di a meter The na ph a se begins du ri ng which th e th re a d
.

sh ortens a nd thicke ns If the re is a si ngle th re a d it now

.

b re a ks i nt o a ce rt a in defi nite numbe r of sep a ra te seg

FI G 68 . .
—S el ecte d
S ta ges in C el l D ivisi on of C ells of th e R oo t
T i p o f th e S pid erw or t 1 T h e chr o m a ti n h a s or ga n
. . iz e d i tself into
a l on g slender t hrea d or sp i rem e a nd th e cy topl a sm begins to S h ow
i ndi ca t i ons o f th e orga n i z a t i on o f th e S pi ndl e 2 T h e spi rem e h a s
. .

b rok en i nto fra gments , th e ch rom o s om es , ea ch Of whi ch is spli t

l engthwise a nd a rra nged a t th e middl e of th e spindl e wh ere th e
two ex a ctly e quiv a l en t h a lv es a re b ei ng s ep a ra ted 3 Th e h a lv es . .

a re co mpl e te ly s e p a ra t e a n d m o vi ng t o w a r d th ei r resp e ctiv e p ol e s

o f th e S pin dl e 4 T h e ch rom os om es h a ve a rrived a t th e p ol es a nd
. .

a re a ggre ga te d i n a cl o s e m a ss b u t h a v e n o t y e t b een su rrou n de d

!

by a nu cl ea r m emb ra ne 5 T h e nu cl ea r m emb ra ne a nd nu cl ea r
. .

sa p h a v e f orm e d a n d th e ch ro m o s om es a re b e gin ni ng to u nd ergo

a p ro ce ss of v a cu ol a t i on whi ch will ev en
-
tu a lly re du ce ea ch on e to a

sta te m ore or l ess resembli ng a very mu ch b rok en rubb er sp onge .

p c p o l a r ca p of c y to pl a sm o u t o f whi ch th e spi n
. .
,
dl e fib ers a re orga n
i z ed ; nm nu cl ea r m emb ra n
. .
, e ; 3 spi rem e ; n n u cl eol us ; sp spi ndl e
,
’
, ,

fibers ; p ol e O f spindl e ; chr ch rom os om es ,

.

me nts c a lled chromosomes Th is numbe r is fixed a nd .

c onst a nt for th e cells of a ny p a rticul a r pl a nt or a nim a l .

2 88 —
CE L L DIVI S I O N

TH E F NU
OR M AT I O N O F T H E A CLE R S P I N DLE .
—At th e
time th e ch rom os omes begin to be devel op ed f rom th e
ch rom a ti nnet th e cyt opl a sm a rou nd the nu cleus p rese nts
a nun ch a nged a ppe a ra nce A s th ey p roceed with thei r
.

devel opme nt fibe rs begi n to a ppe a r a t two sides of the

,

nucleus They a re f ormed f rom th e cyt opl as m itself

.

me rely by a re a rra ngeme nt o f its st ruc tu re Ea ch fibe r .

h a s o ne end d i rected a t a p o i nt a sh ort dist a nce f rom the

nucleus tech nic a lly kno w na s the p ole o f the spi ndle a nd
, ,

th e o the r t ow a rd the nucleus itself (Fig S ome .

time du ri ng th e f orm a ti on of th e nucle a r fibe rs th e

nucle a r memb ra ne dis a ppe a rs th us le a vi ng th e ch rom o
,

s omes f ree i n the cell S ome of the spi ndle fibe rs the n
.

exte nd to th em a nd bec ome a tt a ched The spi ndle fibe rs .

c onve rge to two p oles on Opp osite S ides of th e nucleus ,

an d s ome fibe rs from e a ch p ole bec ome a tt a ched to e a ch

ch rom os ome The p ro cesses so fa r desc ribed bel ong to
.

th e p rep a ra t ory st a ge of divisi on t ech nic a lly k now n a s

,

p o p h a se
r -
.

T H E S P LI T T I N G OF T H E C H R O M OS O M Es
Afte r the .
-

fibe rs a re a tt a ch ed to th e ch rom o s omes th e l a tte r li ne up

an d f orm a fla t figu re midw a y betwee n th e p oles Ea c h .

th e nsplits lengthwise to f orm two ex a ctly simil a r h a lves

(Fig . The S plit occu rs i n a pl a ne a t righ t a ngles
to the spi n dle so th a t the fibe rs f ro m one p ole a re a tt a ch ed
to ch rom os ome h a lves on thei r own side
-
Th e resp ec .

tive h a lves next mig ra te t ow a rd th e p ole to which th ei r

S pi ndle fibe rs a re di rected Th is st a ge of nucle a r divisi o n
.

is c a lled me ta p ha se .

T H E A N AP H A S E (Fig c ove rs th e pe ri od d u ring

.

wh ich the ch rom o somes a re m ovi ng tow a rd the p oles of

the spi ndle I t is e a sy to Obse rve wh a t occu rs bu t diffi
.
,

c ul t to expl a i nh ow it is b ro ugh t a b o u t S ome O bse rve rs

.

a re o f th e o pi n i on th a t the fibe rs u nde rg o a c ont ra cti on

an d d ra w th e ch ro m os omes a l on g with th em O the rs .

thi nk th a t th e fibe rs a re no t s olid a t a l l but me rely rep re

se nt li nes o f f orce a l ong which th e fi ne p rot opl a smic
 T E L OP HA S E AND R E CO N S T R U C TI O N 2 89

g ra nules bec ome a rra nged like the i ronfili ngs i nthe field
o f a m a gn et Insu pp ort O f the l a tte r Opi ni oni s the f a ct
.
'

th a t the p ol a r e nd s o f th e fibers do no t seem to be a t

‘

ta ch ed to a n ythi ng s olid e nough to pe rmit th em to exe rt

a t ra ctive force on the ch rom os omes M ore ove r th e .
,

p ol a r e nds ofte n seem to diss olve a w a y eve n before the

ch rom os omes h a ve c ompleted thei r m oveme nt Whe n .

we c onside r th a t we do not yet u nde rst a nd a t a ll well

h ow a ny s ort of p ro t opl a smic m oveme nt is p roduc ed it ,

sh o uld not be su rp rising th a t this p a rticul a r s ort Sh ould

still be obscu re .

T H E T ELO P H A S E AN D T H E R E CO N S TR U CTI O N O F T H E
N U CLE U S (Figs . n
a d — Wh e n th e ch rom o
s omes a rrive a t thei r respective p oles they a t fi rst
d ra w cl ose t ogethe r a nd O fte n bec ome so c rowded
th a t it is difficult to disti nguish thei r individu a l ou t
li nes (telop ha se ) The next step ordi na rily c onsists
.

i n th e a ppe a ra nce of a new nucle a r memb ra ne cl o sely

su rrou ndi ng the c rowded ch rom os omes S h ortly it will .

be see n th a t th e memb ra ne is enl a rging a nd th a t the re

is now a sm a ll a m ou nt of nucle a r sa p I t is the cur .

re n t opi ni on th a t th e memb ra ne is merely a thi n l a ye r

of cyt opl a sm o f a m or e s olid ch a ra cte r th a n the rest In .

f a ct a ve ry S imil a r l a ye r is p rese nt on th e outside next ,

th e cell w a ll if th e re is one or f ormi ng the oute r c ove r

-
, ,

i ng o f one celled a nim a ls

-
I f one p u nctu res th e o ute r
.

l a ye r O f p ro t opl a sm with a fi ne needle the m ore fluid p a rt

a t on ce ru n s ou t th rough the h ole but a s Soona s it c omes
,

i n c ont a c t W ith th e su rrou ndi ng w a te r i t immedi a tely

f orms a new memb ra ne I t is supp o sed th a t the cyt o
.

pl a sm re a cts i nth is s a me wa y to the nucle a r sa p a s s oon

a s it is f ormed and thus gives rise to th e nucle a r
,

memb ra ne .

I N V I D UA LI TY O F T H E C H R O M OS O M Es
D I — As th e nucleus .

exp a nds th e rod like ch rom o somes a ls o begi n to exp a nd

,
-

an d to bec ome diste nded by v a cu oles p o ssibly o f nucle a r ,

sa p. This p rocess migh t be roughly like ned to the h oles

 290 E —
C LL DI VI S I O N

fo rm ed i nd ough by the ga s p roduced by ye a s t Th e fina l .

result is to red uce e a ch ch ro m os ome to a ra gged n etw ork .

These individu a l nets bec ome cl osely a ss o ci a ted in the

nucleus a nd p rese ntly it is no l onge r p ossible to tell wh e re
on e st ops a nd a no the r begi ns I n excepti on
. a l c a ses the

i ndividu a l nets ca nbe rec og nized a t a ll tim es a nd so we

knew th a t i nthese p a rticul a r org a nisms e a ch ch rom o s ome
m a int a i ns its own i ndi vidu a lity a t a ll tim es Al th o ugh
.

th is Ca nno t be di rectly obse rved in m ost pl a nts a nd a ni

m a ls the re is much indi rect evide nce to indic a te th a t it is
t rue f or th em a ls o .

T H E D I V I S IO N O F T H E CELL B ODY d oes n

-
o t occu r u n til
a f te r th e n ucleus h a s c ompleted its d ivisi on Ab out th e
.

middle o f the spi ndle the re is left a m a ss o f fibe rs which

‘

ex te nd e nti rely a c ross f ro m p ole to p ole a nd a re no t

a tt a ch ed to th e ch ro m os omes As th ey sh orte nth e e nds
.

a re d ra w n i n t ow a rd th e middle a n d th icke ni ngs a re

f ormed a t th e middle p oi nt E ve ntu a lly th ese t ouch a nd
.

S p re a d ou t to f o rm a s or t o f memb ra n o u s pl a te a c ro ss th e

middle o f th e cell A split a ppe a rs i nth is pl a te a nd th e

.

two h a lves sep a ra te slightly so th a t a cell w a l l ca n be

-

l a id d ow nbetwee na nd j o i ned a t its edges to th e ol d w a ll .

Wh e na ll this h a s bee na cc omplish ed it rem a i ns only th a t

the two new cells sh o uld a g a i ng row to full size a nd eith e r
bec ome diffe re nti a ted i nth ei r fi na l f orm or else repe a t the
divisi onp rocess .

S U M M AR Y. Th e imp ort a nt p oi nts to be a r i n mi nd

a re :

1Th a t th e nucleus c ont a i ns a defi nite fixed numbe r O f

.

ch rom o s omes for e a ch a nd eve ry ki nd o f a nim a l o r p l a nt

’

2 Th a t e a ch ch rom os ome m a i nt a i n
. s i ts own ide ntity
th ro ugh o ut a l l the ch a nges o f f orm .

3 Th a t e a ch i n
. c re a ses its subst a nce th rough growth ,

an d d ivides le n gthwise i nt o two ide ntic a l h a lves .

4 Th a t one h a lf o f e a ch g oes to e a ch n
. ew nucleus .

5 Th a t th e n
. e t result o f the wh ole p rocess is to gu a ra n
 S U MM A R Y 2 91

tee th a t e a ch cell Of th e pl a nt or a ni m a l b ody h a s ex a ctly

the s a me numbe r a nd ki nd of ch rom os omes .

NB. . I t will be sh ow nl a ter th a t a ctu a lly e a ch cell h a s

its ch rom os omes i np a i rs one membe r of e a ch p a i r being
,

origi n a lly de rived f rom e a ch p a re nt I t will a ls o be sh own

.

th a t th e membe rs of e a ch p a ir a re ne a rly th ough not

,

necess a rily ex a ctly a like

, .
 C HAPTER XX VI

R E P R O DU C TI O N I N A NI MAL S

R EP R OD U CT I O N species of livi ng thi ng pe rsists

. E a ch
f rom ce ntu ry to ce ntu ry wh il e th e life o f the i ndividu a ls
,

c omp osing i t is o f comp a ra ti vely b ri ef d u ra ti on A dog .

or ra bbi t f or ex a mple c omes i n

,
t o existe nce g rows to a
, ,

defi nite size lives its a ll o tted time a nd p rovides for the
, ,

c onti nu a ti on O f th e ra ce by p roduc ing off sp ri ng to t a ke

i ts p l a ce Th is p ro cess o f p roduci ng new i n dividu a ls a nd
'

.
,

thus i nsu ri ng th e pe rpetu a ti on of the species is k no wn ,

a s rep ro ducti on .

Inthe m a ny celled a n -
ima ls th e divisi onof l a b or a m ong
th e cells c omp osi ng th e b ody is a ss oci a ted with the de
v el O p me n t O f m a le a nd fem a le rep roductive or ge rm
cells which u nde r ce rt a i n c onditi ons to b e desc ribed
,

l a te r devel op i nt o new org a nism s Th is i s k now n a s

, . .

sexu a l rep roducti on In a h ost of l owe r a nim a ls th e

.

b ody divides to form two offsp ri ng ; or bud like ou t -

g rowths a re f ormed which sep a ra te f rom th e b ody a nd

bec ome new i ndividu a ls S ex cells i n such c a ses pl a y .
, ,

no p a rt wh a teve r a nd h e nce this type o f multiplic a ti on

,

is k now n a s a sexu a l rep roducti on In b oth types of .

p rop a g a ti onth e new i ndividu a l h a s its begi nni ng i none

or m o re cells d e rived f ro m th e p a re n t The livi ng ma .

teri a l is p a ssed f rom on e ge n e ra ti on to the next All

”
.

life f rom p re existi ng life i s to da y a rule with out a

- -

si ngle k now n excepti on .

S P O N TA N EO U S G E N ER AT I O N — Al th ough it h a s bee n
.

k now n for th o us a nd s o f ye a rs th a t a nim a ls a nd pl a nts

beget thei r ki nd it wa s a ls o held u ntil rece nt times th a t
,

th is rule is f reque ntly b roke ni nc a ses O f supp osed sp on

292
 B UDDING 293

ta n eou s ge n e ra ti onwh e re it wa s believed nonli vmg ma -

teri a l bec omes t ra n sformed i nt o livi ng subst a nce Fishes .

an d frogs we re th ough t to be devel oped f rom mud m a ny ,

i nsects f rom th e j uices of pl a nts a nd a h orde of org a nisms

,

wa s supp osed to origi na te i n dec omp osing m a sses o f

org a n ic m a tte r In the M iddle Ages ce rt a i n physici a ns
.

m a i nt a i ned th a t sc orpi ons we re ge ne ra ted i na pl a nt th e ,

sweet b a s il a nd the public wa s w a rned a g a i nst smelling

,

it lest sc or pi ons be ge ne ra ted i n the b ra in a nd i ns a nity

result . E ve ni nou r ownda y we h e a r o f h orseh a i rs th a t
.

a re s a id to ch a n ge i nt o sna kes a nd e a rthw orms a nd

,

m osquit o es th a t ra in from th e sky .

D oub t wa s fi rst c a s t on such a belief by th e I t a li a n

na tu ra list R edi wh o i n1 660 pe rf ormed th e simple exp eri
,

ment of tyi ng g a uze ove r the m ou th of a ja r c ont a i ni ng

dec a yi ng me a t a nd p rovi ng th a t fly ma gg ots w ere not
,

sp ont a ne ously ge ne ra ted Othe r i nvestig a t ors exp eri

.

mented with di ffe re nt f orms of life with the s a me result .

Th e bitte rest figh t wa s w a ged a b ou t th e b a cte ri a or ,

mi nute pl a nts which i n s ome c a ses a t le a st did a ppe a r

,

to a rise sp ont a n e ously In 1 86 1

. P a steu r th e g re a t
, ,

French scie ntist t ook th e field a nd a s a result of c a re

, ,

fully s a fegu a rded expe rime nts sh owed th a t eve n he re

sp ont a ne o us ge ne ra ti on d o es not o ccu r I ncide nt a lly it .

ma y be a dded th a t P a steu r s w ork l a id th e f ou nd a ti onfor

l

the s cie nce of b a cte ri ol ogy .

B U DD I N G
. I na sexu a l rep roducti onthe re a re two m a i n
—
types u nequ a l divisi ono f th e b ody or buddi ng (Fig
an d equ a l divisi onk no w na s fissi on(Fig At fi rst sigh t
.

it ma y a ppe a r th a t this is a s omewh a t a rtifici a l di sti nc

ti on but i nre a lity it is of fu nd a me nt a l imp ort a nce a s
, ,

wil l a ppe a r l a te r .

Hyd ra (Fig 7 ) a ff ords a n excelle nt ex a mple of bud

.

di ng in whi ch th e fi rst i ndic a ti on is a p ouch li ke ou t -

g rowth of the b ody w a ll of the side of the a nim a l As

-
.

this i nc re a ses in siz ete nt a cles a nd m outh a ppe a r on th e

free ext remity a nd a fte r fu rthe r g rowth the b ud con
,
294 R E P R O D U CTI O N IN A NI M AL S
st ricts its b a se a nd sep a ra ti ng f rom the p a re nt le a d s
, ,

a ni n de p e nde nt life th e re a fte r .

In m a n y sp onges sm a ll b irds devel op on th e i nside

o f th e b ody a n d with the de a th o f th e p a re nt o r eve n ,

before bec ome f reed a nd g row i nt o th e a dult c onditi on

, .

Am ong th e w orms especi a lly th e j o i nted or segme nted

,

w orms rel a ted to th e e a rthw orm th e re a re m a ny st riki ng ,

ex a mples of buddi ng In ce rt a i n species th e b ody

.

devel ops a h e a d ne a r th e m iddle a fte r wh ich the h i nde r

p orti on with th e new h e a d bec om es det a ch ed a s a new
i ndividu a l a nd swims a w a y I n th e c a se o f the p a l ol o
.

w orm o f th e Fij i a nd S a m oa nisl a nd s th is h i nde r secti on

neve r devel op s a h e a d yet it swims a b o ut i n th e f a ll
“

m onth s i nc o u ntless numbe rs a nd is c ollected a s a h ighly,

p rized a rticle of f ood .

FI G 6 9
. .
— B u ddi ng i n a M a rine W orm .

FIS S I O N . Fissi on or th e equ a l d ivisi onof a norg a nism

,

i nt o two d a ughte r i ndividu a ls is a widesp re a d m ode of ,

a sexu a l rep roducti on with the l owe r a n im a ls Am ong .

th e Unicellul a r org a nisms for ex a mple it is u nd oubtedly

, ,

th e m ost c omm on meth od O f m ultiplic a ti on In th e .

a m oeb a th a t ma y be t a ke n a s a typ e the n

,
ucleu s fi rs t ,

d ivides a nd th e n th e cell p ro t opl a sm g ra du a lly d ra ws

,

a p a rt u n til th e c onnecti ng st ra nd sep a ra tes a nd two O ff

S p ri ng thus a rise Inth e sea a nem one rel a ted to hyd ra
.
, ,

the b ody occ a si ona lly divides i nt o equ a l p a rts a nd a m ong ,

ce rt a in sm a ll species o f fl a tw orms fissi on a ppe a rs to

o ccu r whe n eve r the re is a bu nd a nt nou rishme nt .

I na l l th ese ca ses a n
'

d o th e rs th a t migh t be ci ted the

, , ,

h a lf sized d esce nd a nts g row to th e si z e o f th e origi na l

-

p a re nt a nd repe a t the p rocess if c ondi ti ons a re f a v ora ble

,
.
 R E S UL T S OF A S E X UAL R E PR O DU C TI O N 2 95

Before le a vi ng th is subj ect a n imp ort a nt disti ncti on

betwee nbudd ing a nd fissi onsh ould be no ted Anorg a n .

ism ma y p roduce a few or m a ny buds bu t it ultim a tely ,

dies Infissi on on the o the r h a nd the p a re nt v a nishes

.
, ,

i nthe c re a ti onof two desce nd a nts o u t Of the s a me m a te

ri a l a n d no de a th occu rs see Ch a p XX I I ) ,
.

R ES U LT S OF A S EX UA L R E PR OD U CTI O N Th e uppe r .

limits of sexu a l a ctivity a re re a ched i n th e c a se o f the

sh ipw orm or te red o a nd ce rt a i nspecies o f t a pew orms th a t
la y eggs a ye a r ; yet eve n this numbe r of i n
di v idu a l s is f a r S h ort o f wh a t is k now n to be a s exu a lly
p roduced in m uch less time by m a ny u nicellul a r org a n
isms InS tyl onychi a a cl ose rel a tive of P a ra m oecium th e
.
, ,

b ody divides once i n eve ry eight h ou rs In six d ays if .

,

f a v ora ble c onditi ons c o uld be m a i nt a i ned th e numbe r ,

of Off sp ri n g w ould be c ou ntless a nd thei r c ombined m a ss

w ould weigh a pp roxim a tely f ou r tee n p ou nds In th i rty .

d a ys the a gg reg a te i t h a s bee n estim a ted w ould be a

, ,

milli ontimes g re a te r th a nth e su n Alth ough such ex ces

.

sive numbers a re neve r re a lized th e re is ra pid multipli

,

c a ti on du ri ng fa v ora ble pe ri ods a nd s ome cells a re cer

'

t a i n to be c a rried i nt o situ a ti ons sufficie ntly fa v ora ble

to e n a ble them to pe rsist .

It will be See n th a t while this m ode of rep roducti on

'

te nds to p roduce a lm ost u nlimited numbe rs it obvi ously ,

p roduces i ndividu a ls th a t cl o sely resemble e a ch o the r

n
.

S exu a l rep roducti on onth e c o t ra ry f or re a s ons th a t will

, ,

be sh ow ni na subseque nt ch a pte r is mu ch m ore likely to

,

result i noffsp ri n g th a t a re u nlike a nd a re the ref ore m ore

f a v ora ble subj ects for the p rocess of ev oluti on .

R EGE N ER AT I O N . R ege ne ra ti on is the a bility o f a n

an im a l or pl a nt to repl a ce l o st p a rts In ce rt a i n specie s
.

a m on g th e l owe r a nim a ls th is ma y le a d to the p roducti on

of n ew i n dividu a ls , a nd thus be a form of a sexu a l rep ro
ducti on Th e sp onges m a ny rel a tives O f hyd ra w orms
'

.
, ,

o f v a ri o us kin ds the sta rfi sh es a nd seve ra l o the r a nim a ls

, ,

a re with o ut fi rm p ro tective e n vel op es a nd h e nce a re sub

296 R E P R O D U CTI O N I N A NI M AL S

ject to f reque nt mu til a ti on by the a cti on of w a ves for

, ,

ex a mple Ine a ch c a se th e d a m a ge is rep a i red a nd wh e re

.

th e b ody is t orn i n two th e missi ng p orti ons a re O fte n

repl a ced a n d two i ndividu a ls result A fl a tw orm ma y be .

b roke ni nt o m ore th a ntwo h u nd red pieces a nd e a ch will

f orm a n e nti re new i ndi vi du a l .

C ra bs i n
,
sects a nd spide rs wh e nseized by one of thei r
, ,

legs will o fte n sep a ra te the l imb f rom th e b ody wh ich

, ,

the nscuttles off to a pl a ce o f s a fety Liz a rds a nd s ome .

species of s a l a m a nde rs d rop th e t a il whe nro ughly tre a ted ,

an d wh ile th e a c tively w riggl ing a ppe n d a ge occupies the

a t te n ti on o f the e nemy th e rest of th e b ody ge ne ra lly
esc a pes M a ny sta rfi sh es f reque ntly u nde rg o a simil a r
.

p rocess of self mutil a ti on while a numbe r of species o f

-
,

the rel a ted sea cucumbe rs w ill th row ou t the e nti re d i

g es t i v e t ra ct wh e n m o lested I n m o st c a ses
. th e missi ng
p orti ons a re repl a ce d by the p rocess O f rege ne ra ti on
wh ich is thus seento be of p rime imp ort a nce to th e exist
,

e nce o f m a ny species O f a nim a l s .

R G N ERAT O N I N H GH R A N M A S
E E I I E I L — Am ong th e .

high e r a nim a ls th e p rocess o f rege ne ra ti onis l imited In .

th e f rog a nd rel a ted s a l a m a nd e r th e limbs th a t h a ve ,

bee nbitte nor cu t off will rege ne ra te but de a th results i f ,

the b ody is cu t i ntwo Ana mput a ted fi nge r i nthe c a se

n
.

o f ma n is eve r repl a ced th ough rege ne ra tive ch a nges

,

occu r th a t a re of h igh imp o rt a n ce Th e s ki ng rows ove r .

the d a m a ged a re a a nd be ne a th i t th e c onnective tissue

, ,

which ordi na rily bi nd s the b ody cells t ogethe r f orms a -

,

c omp a ct m a ss o f sc a r tissue This bec omes pe net ra ted by

.

rege n e ra ted bl ood vessels a nd by ne rves whe nthe i nj u ry

-

is sligh t B ro ke n b ones a re rep a i red but muscul a r a nd

.
,

ne rv ous tissues a re neve r rege ne ra ted to a ny l a rge deg ree ,

nor a re i nte rna l org a ns repl a ced by rege ne ra ti on .

A w orkm a ni na na u to m obile f a ct ory wh o h a s a lw a ys ,

m a de b olts c a nno t successfull y m a ke ti res And a pp a r

,
.

e ntly i nm uch the s a me wa y a ne rve cell th a t c onducts -

impulses c a nno t a ssume the du ties o f dig es ti on In

,
.
 S E X U AL R E P R O D U CTI O N 2 97

the r w ords the p owe r of rege ne ra ti on i n org a nisms is

n
o ,

m ost highly devel oped wh e re th e divisi o of l a b or is com

p a ra ti v el y sligh t a nd dec re a ses a s th e c ell s bec o me i n .

crea si n gyl S peci a lized .

— In eve ry a nim a l with th e

~

SEX UA L R E PR O U CT O N
D I .
,

excepti onof the simplest types the cells of the b ody b e ,

l ong to two ve ry disti nct g roups The re a re th ose con .

sti tu ti n g the b o dy p rope r an d o the rs kn o w n ,

a s r ep rodu c

tive or ge rm c ells Th e fi rst f orm by fa r th e g re a te r p a rt

-
.

o f the i n dividu a l a nd a re m ore or less speci a lized to p er

f orm a se ries o f a ctivities a ss oci a ted with food getti ng ,

a v o id a n ce of e nemies a nd th e c a re of th e y ou ng i nm a ny
,

c a ses T hese fu ncti ons h oweve r me rely se rve to ma m

.
, ,

t a i n th e life of th e individu a l a nd s oone r or l a te r a re ,

te rmina ted by de a th .

The ge rm cells on th e c ont ra ry t a k eno p a rt i nthese

-
, ,

b odily a ctivities but a fte r being no u rished for a time by

,

the b ody a re c a p a ble u nde r f a v ora ble c onditi ons of de

, ,

v el op men t i nt o new individu a ls th a t th us pe rpetu a te th e

ra ce . S ince these rep roductive eleme nts a re st ruc tu ra ll y
u nlike inthe m a le a nd fem a le a nd a re a s s o ci a ted with the ,

sex of the i ndividu a l thi s f orm of multiplic a ti onis know n

,

a s sexu a l rep ro ducti on In a subseque nt p a ra g ra ph it

.

will be sh ow n th a t even a m ong th e u nicellul a r a nim a ls

sexu a l rep rod ucti on occu rs a nd a c c ordi ngly is p rob a bly ,

ch a ra cte ristic of eve ry species O f a nim a l M ultiplic a ti on .

by th e sexu a l meth od depe nds onge rm cells or ga metes -

,

an d not onb ody cells while i nthe a sexu a l type th e n

-
,
ew

in dividu a l is p roduced by b ody cells a l one -

.

T H E S EX CELL S -
M a le a nim a ls p roduce ge rm cells
.
- -

k now na s sperms or spe rm cells wh ile the fem a le gives -

,

rise to eggs . Th e spe rms a re devel oped i n a c omp a ct

org a nn a med the testis ; th e eggs a re f ormed i na nequ a lly

well defined orga n the ov a ry E a ch egg or spe rm is a

-
,
.

single cell The Spe rms di ff e r i n sh a pe a s much a s the

.

species p rod uci ng them N eve rth eless a ll a g ree i nbei ng.

ve ry sm a ll a nd the m a j ority a re sle nde r th re a d lik e

,
-
.
2 98 R E P R O D U CTI O N IN A NI MAL S
b odies ; In a typic a l ex a mple (Fig 70 ) the nucleus i s .

a highly c omp a c t b ody i nth e he a d o f th e t a dp ole sh a ped -

cell,
while th e rem a i ning p ro to pl a s m is d ra w n o u t to
f orm a sle nde r th re a d th a t se rves a s a l oc om ot or org a n .

E a ch spe rm is thus a cell wh ose f orm fi ts it for a ctive

swimming m oveme nts e na bli ng it to c om e i n c ont a c t
,

with a negg i nthe p rocess of fe rtiliz a ti on .

E ggs with few excepti ons a re sph e ric a l cells with out
, , ,

th e p owe r o f m oveme nt a nd c ont a i n f ood m a te ri a l

, ,

usu a l ly i n th e f orm o f y olk Whe re the y o u ng a nim a l

.

e a rly shifts for itself a s i nthe c a se of the sea u rchi n or

, ,

FI G 7 0
. .
-
E gg a nd S perm C e lls S m a ll e r S pe rm o f s a m e ma g
.

n
i fi ca ti o na s e gg, i nwhi ch s o m e o f th e f o o d y o l k is sh o w n .

wh e re i t d ra ws its f o od supply from th e b ody o f th e

p a re nt a s i nth e ra bbit the a m o u nt o f y olk is sm a ll a nd
, ,

th e egg sc a rcely me a su res o ve r one o r two h u nd redth s o f

a nin ch i n d i a mete r N eve rtheless such ova a re m a ny
.

h u nd reds o f th ous a nds o f times l a rge r th a nth e spe rms o f

the s a me spe cies A t th e opp osite end o f th e se ries th e
.

a m ou n t o f y ol k is e norm o us In a J a p a nese sh a rk fo r
.
,

ex a mple th e egg is ove r eigh t i nches i nd i a mete r i n the

, ,

o st rich i t is o ve r th ree i nch es i n the d omestic f owl one ,

i nch a nd i n m a ny o the r fish es liz a rds a nd tu rtles th e

, , ,

egg cells a re only sligh tly sm a lle r .

A C CES S ORY S Ex OR GA N S i nthe highe r a nim a ls a re th ose

by me a ns of which spe rms a nd eggs a re b ro ugh t t ogethe r .
 A CCE S S O R Y SEX O R GAN S 299

Ve ry few eggs nd no spe rms ca n devel op int o new i n

a

di vidu a l s with ou t fusi on with th e g a mete of th e o the r ~

sex In s ome a qu a tic a nim a ls this is ve ry simple si nce

.
,

the m a ss of eggs which the fem a le l a ys i n the w a te r

a tt ra cts th e m a le p rob a bly by the stimul a ti on a ff orded
,

by s ome chemic a l subst a nce which th ey give off a nd ,

c a uses him insti nctively (see C h a p X V ) to dep osit the .

S pe rms i n c on t a c t with them In l a nd a nim a ls th is is

.

no t p ossible a nd i t bec omes necess a ry for the fem a le to

,

ret a i n th e eggs in a speci a l org a n whe re the m a le ma y

dep osit th e sperms i n orde r to pe rmit thei r fus ion with

on e an o the r .

FERTI LI Z ATI O N c onsists i n the fusi on of two cells a n ,

egg a nd a spe rm e a ch o f which is a highly org a nized u nit

,

with o ut th e p owe r of fu rthe r divisi on At the begi nni ng .

of th e p ro cess th e spe rm cell a tt a ches i tself to th e su rf a ce

-

O f h e egg (Fig 7 1 ) a n d a pp a re ntly is c rr ied inw d by

t . a a r

m ovements of th e egg p rot opl a sm D u ri ng i ts c onti nued .

j ourney i nt o the i nte ri or th e Spe rm nucleus g ra du a lly en

l a rges th rough the a bs orpti on of fluid u ntil fi na lly it
bec omes of the s a me size a s the egg nucleus with wh ich
it comes i nc ont a ct .

R ES U LTS O F F ER TI LI Z ATI O N The imp ort a nt results of

.

fe rtiliz a ti on a re two f old In th e fi rst pl a ce the u ni on

-
.

o f the two sex cells e a ch a l one i nc a p a ble of divisi on

-
, ,

supplies a stimulus th a t impels the fe rtilized egg to

u nde rg o a l a rge numbe r O f divisi ons resulti ng i n a new
i ndividu a l In th e sec ond pl a ce e a ch of these sex cells
.
-

c a rries a defi nite numbe r of ch rom os o mes ch a ra cte ristic of

th e species Let the numbe r be f o u r ; the nu cleus f ormed
.

by thei r u ni onthus c ont a i ns eigh t At eve ry cell divisi on .

these a re a ccu ra tely h a lved a nd dist ributed so th a t e a ch

cell of the b ody receives a nequ a l numbe r o f p a te rna l a nd
m a te rna l ch rom os omes Th e h igh imp ort a nce of this f a c t
.

w ill be c onside red i n c onnecti on with th e subj e ct of

he redity bu t a t this p oi nt it ma y b e p oi nted ou t th a t
,

ch rom a ti n is the be a re r o f he redit a ry t ra its a nd the off

300 R E P R O DU CTI O N IN A NI MAL S
sp ring i nh e rits eq u a lly f rom eithe r p a re nt The new .

individu a l is ce rt a i n to be sligh tly d iff ere nt f rom eithe r

p a re nt or thei r a ncest ors a nd th e re is a ch a nce th a t i t
ma y be bette r fitted to c onti nue th e ra ce (S ee Ch a p . .

XXX V ) .

I N DI V I DUA LI TY O F T H E C RO OS O
H M M E S — I t h a s bee n .

st a ted in the p recedi ng p a ra g ra ph th a t a ll of the b ody

cells a nd the rep rod uc tive eleme nts th rough o u t th e ,

g re a te r p a rt of thei r devel opme nt h a ve a defi ni te numbe r ,

o f ch rom os omes And this numbe r is est a blished a t the

.

FI G 7 1
. .
-
D ia gra m il l us tra ting
f ertili z a ti on of th e e gg a nd
th e
ea rly d e v el o pm e nt A e ntra nce o f th e S pe rm
.
, B a pp ro a ch o f .
,

spe rm to e g nu cl eus C u ni on of nu cl ei D spli tting o f ch rom o

.
, .
,

so m es (th o se o f S perm s olid bl a ck ) E se p a ra ti o n o f the chro mo

.
,

so m es p ri or to F th e di vis io n o f th e emb ry o into two cel ls

, .

time of fe rtiliz a ti onby th e u ni on of th e egg a nd spe rm

nucleus S ome of the i nsects for ex a mple h a ve eigh t
.
, ,

ch rom os omes ce rt a in sea u rch i ns S ixtee n s ome o f the

,
-
,

w orms twe nty f ou r wh ile the re a re f orty eigh t i n ma n

-
,
-
.

I f the re a re eigh t ch ro m os omes i n th e fe rtili zed egg this ,

ch a ra cte ris ti c numbe r ca nbe c ou nted a t e a ch o f th e th ou

s a nds of succeed ing divisi ons I ndeed i n s ome s p ecies .
,
 I N D I VID U AL IT Y O F C H R O MO S OM E S 301

i t is p ossible to f oll ow th e ch rom os omes f rom one cell to

an o the r
. In o the rs the re a re ce rt a i n size diffe re nces
a m on g th e ch rom os omes th a t a ppe a r u nch a nged f rom one
cell g ene ra ti on to the next a nd h ence suggest th a t he re
,

a ls o these b o dies a re self p erp e tu a ti n n

'

-
g u i ts Als o the re .

is a we a l th of evide nce i nc onnec ti onwi th b reedi ng exp eri

me nts which p ra ctic a lly a m ou nts to defi nite p roo f th a t
ch rom o somes ret a i nth ei r i ndividu a lity Ino the r w ords .
,

f ou r of th e eigh t a ssumed ch rom os omes i n eve ry b ody

cell a re th e di rect desce nd a nts of th o se i nt roduced by
th e spe rm while th e oth e rs a re f rom th e egg J ust a s
, .

in dividu a ls or cell s a re th e li ne a l O ffsp ri ng th ro ugh

th ous a nds of gene ra ti ons of o th e r i ndividu a ls or cells ,

so a re ch rom os omes th e desce nd a nts of p re existi ng -

ch rom os om es .

FERTI L I Z ATI O N OF U N I CE U LAR A N M A S

L L I L — Am ong
the u nicellul a r a nim a ls whe re O bvi o usly the re a re no spe
ci a l rep roductive cells th e r e is n
,
eve rth eless a fe rtiliz a ti on
p rocess th a t i n i ts esse nti a l fe a tu res resembles th a t o f
higher a nim a ls In P a ra m oecium for ex a mple two i n
.
, ,

di vi du a l s Come i n c on ta c t a n d p a rti a lly f us e to get h e r

f or a few h o u rs time a n d du ring the pe ri od u nde rg o a
’

c omplic a te d se ries of nucle a r c h anges With o ut c onside r

.

i ng the p ro cess i n det a il it ma y be s a id th a t ch rom a ti n

m a te ri a l is exch a nged a nd whe n the two cells sep a ra te
the nucleus of e a ch is a fusi onp roduc t a s i n th e c a se of
the nu cleus of th e fe rtilized egg E a ch i ndividu a l thus
.

ma y be s a id to h a ve fe rtilized the o the r Inseve ra l othe r .

spe cies the re a re sm a ll a ctively swimmi ng i ndividu a ls ,

usu a l ly c onside red to be m a les th a t pe rm a ne ntly fuse

,

with a nd thus fe rtilize l a rge r fem a le i ndividu a ls .

I t is ge ne ra l ly believed th a t th e i nt roducti on of new

ch rom a tin i nt o the cel l results i n a ch a nged i ndividu a l
th a t ma y be bette r fi tted to pe rpetu a te the ra ce th a nthe
p revi o us t ype Bu t th a t th e u ni onof the two u nicell ul a r
.

org a n isms results i na ni nc re a s ed c a p a ci ty for cell divisi on

is a deb a t a ble m a tte r Ins ome i ndividu a ls it a ppe a rs to
.
3 02 R E P R O DU CTI O N I N A NI MAL S
be the c a se ino the rs i t is quite th e c ontra ry Evide ntly
,
.

the two p rocesses a re no t so cl osely a s s oci a ted as i n

highe r typ es but the subj ec t requi res fu rthe r in
,

v es ti ga ti on .

E AR LY D EVELO P M E N T Th e u ni onO f two ge rm cells

.
-
,

a s h a s bee n p o i n ted ou t resul ts i n eve ry a n i ma l i n a

,

single celled fe rtilized egg Bey ond this p oi nt its devel

-
.

O p men ta l p a th depe n ds up onth e ki nd of org a nism it is ,

a lth o ugh th e re a re ce rt a i n fu nd a me nt a l resembl a nces

th rough o u t .

I nthe st a rfish wh i ch we sh a ll use a s a n ex a mple th e

, ,

fe rtili zed egg divides by th e ordi na ry p rocess of divisi on

i nt o two cells o f equ a l size (Fig
, These a g a i n .

divide i nt o f ou r th ese i nt o eigh t a nd so on u ntil

, ,

h u nd red s of cells o f ne a rly equ a l size a re p roduced D ur .

i ng this segme nt a ti on pe ri od a s it is te rmed whe n th e , ,

cells a re dividi ng with ou t f o rmi ng a ny defi ni t e org ans ,

a c a vity a ppe a rs i n th e midst o f th e cells a n d i nc re as i ng ,

i nsize a s the cells i nc re a se i nnumbe r bec omes th e h oll o w ,

o f a S phe ric a l o n e l a ye red g ro up -

The y o u ng st a rfish or .

emb ry o resembles a te nnis b a ll a nd a t th e end o f th ir ty ,

six h o u rs h a s a tta i ned th e bla s tu la s t a ge

’

T H E G A S TR U LA AN D B ODY L AY E R S — With th e i n crea s .

in g d ivisi o no f th e cells the bl a stul a bec o mes s omewh a t

,

fl a tte ned onone side a nd is fi na lly pushed i nw a rd i nm uch

,

the s a me m a nne r a s o ne p ushes i n the side o f a h oll ow

rubbe r b a ll .Th e cup like dep ressi ondeepe ns a nd fi na lly
-

bec omes a c omp a ra tively sle nde r sa c kno w na s the p rimi

tive gut or i n testi ne The emb ry o is no w in the ga s tru la
.

st a ge.

A t a b ou t this tim e ce rt a i ncells a ppe a ri ng like mi nu te ,

a m oeb a e mig ra te f ro m the tip o f th e p rimitive gut a n

,
d
w a nde r a b o u t i nthe fluid wi thi nthe c a vi ty betwee nth e
gu t a nd o ute r cells The emb ry o now c omp rises a noute r
.
,

inne r a nd middle cell l a ye r respe ctively na med the ect o

, ,

de rm e nd ode rm a nd mes ode rm T O a nticip a te it ma y be

, ,
.
,

sa id th a t th e e c tode rm gives rise to the S ki n se n se o rg a ns , ,

 GA S T R ULA AND B OD Y L A Y E R S 303

and ne rv ous system ; the e nd ode rm f orms th e li ni ng of the

digestive t ra ct a nd th e l a rge r p a rt of th e digestive gl a nds ,

such a s the live r a nd p a nc re a s ; while th e mes ode rm

devel ops i nt o a ll o f the o th e r org a ns of the b ody such a s ,

muscles bl ood b ody c a vity a nd ge rm cells

, , ,
-
.

FI G 7 2
. .
— Ea rly
D ev el o pm ent o f a S ta rfish . A , B , C , D , ea rly
segm enta ti on sta tes , E , ea rly bl a s tul a , F, l a te bl a stul a in section,
G, ea rly ga s trul a ; H, l a te ga s trul a sh owing bud di ng o f b o dy ca vi ti e s
from ti p of gu t (front V i ew ) , I , side V i ew of H , sh owing d evel opm ent
of m o t h
u .

L ATER D EVELO P M E N T Th e hyd ra a nd th e rel a ted sea

.

an em ones a nd j elly fi sh es a re little m ore th a n modi fi ed

-

g a st rul ae a nd a cc ordi ngly do no t devel op much bey ond

,

this p oi nt In the highe r g roup s on the c ont ra ry the

.
, ,
30 4 R EP R ODU CTI O N I N ANI M AL S
b ody l a ye rs u nde rg o m a ny ch a nges i na tt a ining the a dult
con di ti onf Th e st a rfish fo r ex a mple f orms two p ouch
, ,

l i ke o utg rowths on the tip o f th e i ntestine th a t fi na lly

seve r the i r c onnecti on, like bubbles bl ow n f rom a pipe ,

an d ultim a tely f orm th e b ody ca vity wh ich c on t a i ns th e ,

,

digestive t ra ct a nd the ge rm cell s

,
-
.

A t a b ou t the time the b ody c a vity 1 5 devel opi ng th e ,

m o uth c a vity a ppe a rs a s a dep ressi on i n th e ect ode rm

opp osite the f ree en d of th e gut with which it fi n a l ly

c omes in c ont a ct Th e w a lls a t the p o int of u ni on

.

bec ome pe rf ora ted a nd th e a lime nt a ry c a na l thus bec omes

a tube O pe n a t b o th e nds .

V ERTEBRAT E D EVE LO P M E N T The devel opme nt of th e

.

st a rfish a s th us desc ribed is i n a b roa d wa y esse nti a lly

, ,

th e s a me a s th a t occu rri ng i nm ost of th e l a rge r g ro ups of

an im a ls Bey ond th is st a ge the g rowth O f the i ndividu a l
.

is l a rgely a m a tte r O f d et a il with o ut p rof ou ndly m odify

in g th e fu n d a me n t a l pl a n O f th e b o dy Th e f ro g f o r .
,

ex a mple p a sses th rough a g a s t rul a st a ge devel ops a b ody

, ,

c a vity a nd the nceforth t ra vels a devel opme nt a l p a th th a t

,

i s ch a ra cte ris tic a lly ve rteb ra te The m ost st rik ing fe a tu re .

of this e a rly pe ri od of g rowth is th e devel opme n t o f th e

ce nt ra l ne rv ous system wh ich i n the l a t e g a s t rul a st a ge
,

m a kes i ts a ppe a ra nce a s two f olds cl ose t ogethe r th a t , ,

ex te nd th e le ngth O f th e a nim a l a l ong the b a ck (Fig .

Th ese i nc re as e i nh eigh t meet a nd fuse a l ong th e mid

,

line a nd th e tube th us formed sep a ra tes f rom the ski n

,

an d sinks be ne a th it .

Th e m outh c a vity th a t h a s f ormed a s in th e st a rfish , ,

devel ops i na ll cl a sses o f ve rteb ra t es f o u r or five gill slits -

th a t fi na lly o pe nto th e exte ri or a s i na fish Inl a te r life ,

.

th e gills dis a ppe a r i nth e a i r b re a thi ng ve rteb ra tes The

- .

digestive t ra c t le ngth e ns f o rms a st om a ch a nd d evel ops

, ,

p ouch es th a t fi na lly result inlive r p a nc re a s a nd excep t , , ,

i nthe fishes lu n gs O rg a ns o f speci a l se nse a re i nl a rge

,
.

me a su re m o dific a ti ons of th e e ct ode rm a nd a re o f too

c omplic a ted a ch a ra cte r to pe rmit o f a b rief desc ripti on .
 G R OW T H 30 5

Limbs a rise a S kelet onis f ormed th e bl ood vessels m a ke

, ,
-

thei r a ppe a ra nce a nd with g rowth ch a nges th a t give th e

,

b ody its fi na l sh ape th e ski n gives rise to th e sc a les ,

fe a the rs or h a ir th a t a re ch a ra cteristic of th e a d ult .

G RO WT H Inthe devel opme nt of a na n

. im a l f rom th e
egg to th e a dul t the b ody a bs orbs y ol k a nd o the r f ood
-

subst a nces f orms p rot opl a sm i n excess of th a t which is

,

dest royed a nd thus g rows In e a rly life especi a lly the

,
.
, ,

i nc re a se i nbulk is ve ry ra pid but it g ra du a lly sl ows d own

a s the a dult c on diti onis a pp roa ch ed D u ri ng this pe ri od .

FI G 7 3. . D ev el o pm ent of Fro g . A , ea rly d ev el opm en t of cen tra l

nerv ous syste m of fro g B , cl osi ng . of n erv o us sys tem f o lds . C, l a ter
s ta te wi th S kin cu t a w a y to S h o w co mpl e te s ep a ra t i o n o f nerv ous

system a nd a S imple sk el eton . Three gill sli ts a re S h ownnea r m ou t h

op en ing .

the a nim a l ma y be st a rved a nd rem a insm a ll bec oming a ,

dw a rf or i t ma y be a bu nd a ntly nou rish ed a nd grow to

,

l a rge size N eve rth eless g rowth h a s its limits excep t

.
, ,

pe rh a ps i n th e c a se of ce rt a in sp onges a nd a few Species

O f dev i lfi sh es A m ouse f or ex a mple ma y be supplied
.
, ,

W ith a n unlimited a m o u nt of f ood yet it neve r g rows to

th e S ize of a r a t th ough the b odily org a niz a ti on is
,

p ra ctic a lly the s a me i n b oth .

D I FFERE N TI ATI O N D evel opme nt p roceeds a s h a s .

,

been desc ribed from a st a ge W he re no org a ns exist

,

th rough a st a ge Wh e re they a re but roughly o utlined a nd

on ly g ra du a lly r e a ches th e p oi nt wh e re th ey a re i n a
,

c ompleted fu ncti ona l c onditi on I f the cells a re ex .

a mi n ed du ri n g this pe ri od they als o W ill be see n to p ro

306 R E P R O DU C TI O N I N A N I M AL S
g ress f rom a c onditi onwhe re a l l a ppe a r to be esse nti a lly
a like i n f orm a n d s tructu re They usu a lly m ove a b out
.

to a g re a te r or less deg ree i na ssumi n g thei r p rope r p osi

ti ons a nd it is only t ow a rd the cl ose o f devel opme t th a t
,
n
they a ssume the ch a ra cte risti c sh a pe a nd o rg a niz a ti ono f
the v a ri ous tissues D evel opme nt a nd d iffe re nti a ti on
.

thus p roceed a t a neq u a l p a ce .

M ETA M OR P H OS I S — The p rocess of devel opme nt h a s

.

b ee n pictu red a s a dv a nc i ng by g ra du a l st a ges f rom the

egg to the a dult The re a re m a ny a nim a ls h oweve r i n
.
, ,

which such is not the c a se The y o u ng butte rfly fo r ex

.
,

a mple , h a tch es f rom th e egg a s a c ra wli ng w orm like -

c re a tu re the c a te rpill a r or la rva th a t a fte r feedi ng on

, , ,

le a ves a nd i nc re a si ng i nS ize e ncl oses itself i na web or a t

,

le a st bec omes a qu iet p u p a ; whe n ce rt a i n c omplic a ted

ch a nges h a ve e nsued withi n th e b ody th e pup a l c a se ,

bu rsts a nd a wh olly di ffe re nt t ype o f i ndivid u a l eme rges

fi tted for a life on the wi ng a nd a diet o f nect a r Th is .

m a rked ch a nge of f o rm a nd fu nc ti onis k now na s a met a

m orph osis (Fig .Fly l a rv a e or m a gg o ts th e ,

wriggle rs o f m osquit oes th e y o u ng of s ta rfi sh es sea

, ,

u rchi ns c ra bs a nd w orms a re f reque ntly so u nlike the

,

a dults th a t it is on ly a fte r th e met a m orph osis th a t th ei r

r el a ti onship ca nbe dete rmi ned .

Am ong th e ve rteb ra tes th e f ro gs a nd t o a d s u nde rg o a

,

c omplic a ted m et a m o rph o sis whe reby the fi sh like w a te r -

i nh a biti ng t a dp ole bec omes t ra nsfo rmed i nt o th e a i r

b re a thi ng f ou r legged a nd t a illess l a nd a nim a l Inm a ny
,
-
.

eels th e ch a nge from the t ra nsp a re nt fl a tte ned l a rv a to

th e rou nd pigme nted a d ul t is l i kewise ve ry ma rked .

Fi na lly th e re a re m a ny l a rv ae especi a lly i nth e sea wh ose

, , ,

li fe hist ories a re u nk now n a nd wh ose s tructu re is so u n

l i ke th a t O f a ny k now n a d ult th a t thei r p ositi on i n th e
an im a l ki ngd om is p roblem a ti ca l .

C R E O F GGS
A E — Am ong m a ny o f th e l owe r a nim a ls ,

su ch as s ta rfi sh es sea u rchi ns a nd m a ny w orms the eggs

, , ,

a re l a id i nthe w a te r a n d a re f orthwith a b a nd oned by th e

 CA R E O F E GG S 30 7

p a rent On th e o the r h a nd th e g re a t m a j ority of egg

.
,

l a yi ng or ovip a ro us species i nsu re th e devel opi ng y o u ng

a ga i nst the a tt a cks of e nemies by p rovidi ng them with
s ome me a ns of p rotecti on Th e c ra bs for ex a mple a tt a ch
.
, ,

the eggs to a ppe nd a ges of the b ody a nd figh t off i nv a de rs ,

the spid ers O fte n c a rry th ei r ov a i n silke n c a ses m a ny ,

an im a ls const ruct b roo d p ouches a nd th e re a re th ous a nds

,

o f species th a t hide thei r eggs i n s a fe ret re a ts Am ong .

m a ny o f the b ees Wasps a nd ce rt a i n beetles not only

, ,
”
a re th e eggs hidde n but a ls o a supply O f f ood is pl a ced

withi n e a sy re a ch .

M a ny of the fishes such a s s a lm ona nd t ro ut bu ry the

, ,

The M eta m o rph o sis of th e M ona rch B tterfly

u .

eggs i n th e s a nd o the rs like th e sti ckleb a ck const ruct

,

el a b ora te nests for them th e sea h orses a nd rel ated pipe

,

fishes ret a i n th e eggs a nd y ou ng i n f olds o f ski n u nde r

ne a th the b ody Wh ile ce rt a i n o the r fishes a tt a ch th ei r
,

O ff sp ri ng to th e ski n or eve n c a rry them in the m o u th .

Th e eggs of frogs a nd t oa ds a re usu a lly l a id i n w a te r ,

th ough a few dry co u nt ry spe cies conce a l them i n f olds

o f the ski n on th e b a ck Th e eggs of bi rds a re usu a lly
.

pl a ced i nnes ts a nd the he a t requi red for devel opme nt is

p rovided by th e b roodi ng p a re nt In th e m a mm a ls or .
,
308 R E P R O DU CTI O N I N ANI MAL S
h a iry a nima ls the eggs,
a re r et a i ned withi n th e b ody of

the m o the r .

N OU R S M E N T OF Y O U N G
I H — V e ry little f ood y ol k is
st ored inthe eggs of m a ny species of a nim a ls a nd a lm o st ,

a t th e o utset th e y o u n g a re c ompelled to fora ge for

themselves Inothe r species wh e re the y olk is of g re a te r
.
,

a m ou n t a nd h a tch i ng a cc ordingly is del a yed th e st ruggle ,

f or existe nce begi ns a t a s omewh a t l a te r pe ri od In .

ei th e r eve nt h oweve r no p a re nt a l c a re is exe rcised a nd

, , ,

the newly h a tched off sp ri ng a re fully equipped for a n

a ctive in depe nde nt life This i n l a rge me a s ure con
.
, ,

tin u es to be the rul e i nn ume ro us species Whe re the y ou ng

a ctu a lly a re un de r th e c a re of th e p a re nts Am ong th e .

bi rds for ex a mple th e p a re nts no rm a lly supply s ome

, ,

nou rishme nt but i n the c a s e of th e ost riches ra il s

, , ,

pl ove rs ducks qu a ils a nd seve ra l rel a ted fa mi lies th e

, , ,

y ou ng ve ry e a rly bec om e i ndepe nde nt i n this rega rd .

I nthe m a mm a ls or h a i ry a nim a ls th e eggs a re ret a ined

i n th e b ody o f th e fem a le a nd with the excep ti on o f
,
-

th e cu ri ous d uckbill a nd th e Spi ny a nte a te r O f th e Au s

tra li a nregi on th e devel opi ng y o u ng a re p rovided with
speci a l memb ra nes a nd bl ood vessels fitted for the a b sorp -

ti ono f f ood subst a nces f rom th e p a re nt E ve na fte r bi rth .

th ey a re c omp a ra tively helpless a nd a re fu rth e r nou rished ,

by milk devel oped i nth e gl a nds of th e m othe r .

S E CO N D AR Y S EX UA L C H AR A CT ER S — Am ong th e l owe r
an ima ls a m a le usu a lly ca nbe disti nguish ed f rom a fem a le
on ly by dissecti onor mic ro sc opic a l ex a mi na ti on The sex .

o f th e high e r species o n th e c o n t ra ry is ge ne ra lly a ss oci

a ted wi th ce rt a i nst ruc tu ra l peculi a rities kn o w na s sec on d
a ry sexu a l ch a ra c te rs The m a les for ex a mple a re O f
.
, ,

l a rge r size th a nth e fem a les or th ey ma y be ch a ra cte rized

,

by h ighly devel oped tusks h orns excessive growth of , ,

h a i r b ril li a nt plum a ge spu rs c ombs sce nt gl a nd s or

, , , , ,

o th e r org a n s th a t a re d evel oped i nv a ri o us regi ons o f the

b ody . The fem a le o fte n p o ssesses highly devel oped
org a n s for th e p ro tecti ona nd no u rishme nt of th e y ou ng ,
 S E CO NDA R Y S E X UAL C HA R A C T E R S 309

but a side from these ch a ra cte rs h er m odific a ti onis usu a lly

less ext reme th a ni nthe m a le .

I n ce rt a i n c a ses these sec ond a ry sexu a l ch a ra c te rs a re

o f evide n t use H orns a re o f se rvice to th e m a le i n de
.

fe ndi ng the fem a le or i nb a ttli ng for a m a te Th e b rood .

p ou ches of m a ny fem a les such a s th e op ossum a nd k a ng a

,

roo se rv e to c a rry the y ou n

,
g Org a ns of sce nt ph osph or .
,

escent org a ns a nd Sou nd p roduci ng st ructu res a s i n th e

,

k a tydid a re believed to a tt ra ct membe rs o f th e opp osite

sex B u t the re a re ve ry m a ny o th e r ch a ra cte rs of this
.

cl a ss such a s the b rilli a nt plum a ge of m a le bi rds a nd

,

th e c ol ors of m a ny i nsects a s well a s spi nes k nobs , , ,

c ombs w a ttles etc wh ose sig nific a nce a t th e p rese nt

, ,
.
,

time is a n u ns olved p roblem .

B ODY AN D G ER M CELL S — AS a result o f th e g rowth

an d diff e re nti a ti ono f a sex u a lly devel op ed i ndividu a l two
cl a sses of cells a re p roduced Th e fi rst g roup c omp rises .

the rep roductive or ge rm cells which a re p ro tected a nd ,

nou rished by th e sec ond cl a ss th e b ody cells This l a st , .

na med a ggreg a te f ormi ng by fa r th e gre a te r bulk O f the

,

i ndividu a l pe rf orms m a ny duties i ncludi ng f ood g a th e r

, ,
;

in g, a v o id a nce o f e nemies a n d v a ri o us a ctivities

,
i n con
necti on With th e p rop a g a ti on a nd nou rishme nt of th e
y ou ng Th e ge rm cells on the c ont ra ry t a ke no p a rt i n
.
, ,

these pu rely i ndividu a l p rocesses a nd a re s olely c once rned ,

with the pe rpetu a ti onof th e ra ce .

Fu rthe rm ore i t is imp ort a nt to note b oth o f these

, ,

cell cl a sses a re equ a lly th e desce nd a nts of a n origina l

fe rtilized egg Th e h en by wh ich is me a nt the cells of
.
,

the b ody d oes not p rodu ce eggs ; it me rely la ys egg cells

,

th a t du ri ng the c ou rse of devel opme nt h a ve be come dif

ferenti a ted a n d set a side mu ch a s muscle or b one cells
a re diff e re n ti a ted a nd g rouped i nt o tissues E a ch egg .

ce l l whe nfe r tilized p ro du ces i n due time o the r eggs a s

, ,

well a s the p ro tecti ng b ody a nd wh ile the b ody dies ,

a fte r e a ch p e ri o d o f sexu a l m a tu rity s ome ge rm cells ,

pe rsist from ge ne ra ti onto gene ra ti on .

31 0 R E P R O DU C TI O N I N ANI M AL S
E AR LY H I S TOR Y O F G ER M C ELLS Alth ough th e rep ro
.

d u cti v e eleme nt s a re h eld wi thi n the b ody O f a na nim a l ,

much like pe a s i na p od this d o es no t me a n th a t du ri ng

,

th e m onths or ye a rs th a t they a re thus imp ris oned they

li e d o rm a nt . AS a m a tte r O f f a ct ge rm cells h a ve a ve ry
defi nite devel opme nt which begi ns a t a n e a rly a ge a nd
le a d s up to th ei r c ompleted f orm ; a nd thi s h ist o ry i ni ts
esse nti a l fe a tu res is the s a me for b oth th e spe rms a nd
the eggs .

In m a ny a n i m a ls the ge rm cells a re rec og n iz a ble i n

th e g a st rul a st a ge o r eve n e a rlie r th ough i n nume ro us ,

o the r species they a re n o t dis ti nguish a ble u ntil a much

l a te r pe ri od These origi na l or p rimitiv e sex cells a s
.
,

th ey a re te rmed a s a result o f repe a ted divisi o n (Fig 7 5 )

,
.

bec ome ve ry nume rous a nd cl osely p a cked t o ge the r

, , ,

ma y for a time ce a se to devel op fu rth e r S oone r or l a te r .

,

h oweve r th ey e nte r a g rowth pe ri od d u ri ng wh ich time

, ,

they inc re a se i n size owi ng to th e a bs orpti on of f ood

,

( y olk i n the c a se o f eggs ) a nd bec o me ,

a cc ordi ng to th e ,

sex Of the p are nt ei the r spe rm or egg m o the r cells They

,
.

now u nd e rg o two fi na l reduci ng or m a tu ra ti on divisi ons

th a t i n th ree respects a re O f sup reme imp ort a nce Th e .

m a tu ra ti ono f th e spe rm cells will fi rst be desc ribed .

R ED U CI N G D I V I S I O N I t will be re ca lled th a t i n th e
.

a ct O f fe rtiliz a ti o ntwo n uclei e a ch c a rryi ng f o u r ch rom o

,

s o mes i n the species ch ose na s a nillust ra ti on fuse a nd

, ,

the result a nt cell thus co nta i ns eigh t A t e a ch o f the .

succeedi ng d ivisi ons e a ch o f th ese ch rom o s omes splits

le ngthwise so th a t eve ry cell o f th e b ody includi ng th e ,

ge rm cells up to the m a tu ra ti on divisi o ns p ossesses this ,

ch a ra cte ristic numbe r A t th e begi nni ng o f the divisi on

.

o f a spe rm m o th e r cell eigh t ch rom o s o mes a ppe a r but

, ,

i ns te a d O f spli tti ng they u nite or conj ug a te inp a i rs one ,

m a te rna l wi th one p ate rna l (Fig I f the ch rom o .

s omes th a t c a me i nwi th th e spe rm a t th e time o f fe rtiliz a

ti onbe desig n a ted a b c d a n
,
d th o se o f th e egg be desi g
, , ,

na ted a s A B C D , the n the c onj ug a ti on O f the

, , ,
 R E DU CING DIVI S I O N 31 1

ch rom os omes is illust ra ted by Aa B b C c D d A spi ndle , , , .

forms a nd th e ch rom o s omes th a t h a ve p re vi ously u nited

i np a i rs sep a ra te with o u t splitti ng a nd t ra vel i nOpp osite
di recti ons Th is is k now na s th e reducti ondivisi on Si nce
.
,

two cells a re f ormed with on ly fo u r ch rom os omes a piece

in ste a d of eight E a ch of these cells divides by the usu a l
.

p rocess a nd the f ou r th a t result u nde rg o ch a nges a nd

Mtt l t i p l i é a t i on of e ar l y
g erm c e l l s .

Pa i r i ng 0
Chr omos ome

Reduc t i on di vi
s i on

ur e Egg

l / ‘ ert i nz a t i on of Egg

en
9 g Ea r l y Deve l og n t

FI G 7 5
. . D i a gra m S h owi ng d ev el opme nt Of th e ge rm cells .

bec ome m a tu re spe rmcells E a ch m a tu re spe rm thus

.

h a s one h a lf the u su a l numbe r of ch rom o s omes th a t ch a r

a cteri z e s th e cells o f th e b ody The egg cells u nde rg o a
.

simil a r reducti on divisi on so th a t th e u ni on O f a spe rm

,

with a n egg p roduces the specific numbe r th a t is eigh t , ,

.

We re there no reducti oni nth e ge rm cells a spe rm u ni ti ng

with a n egg w o uld p rodu ce a cell with sixtee n ch rom o
s omes a nd it is k now nfrom expe rime nt th a t th e devel op
,

me nt i nsuch a c a se w ould be a b norm a l .

R A N DO M A S S OR T M E N T O F C H R OM O S O M E S Inth e re .

ducti ondivisi on a s j ust desc ribed the ch rom os omes a re

, ,
31 2 R E P R O DU CT I ON I N A NI MAL S
sep a ra ted so th a t e a ch result a nt ge rm cell o f th e sec ond
o rde r c ont a i ns one membe r of e a ch p a i r Bu t wh ethe r
this me m
.

be r wa s de rived f rom th e spe rm or th e egg a t

th e time of fe rtiliz a ti onth a t is wh ethe r it is p a te rna l
, ,

FIG 7 6
. . D i a gra m illus tra ti ng the re du cti o no f the chr o m o so m es
.

A, n u cl e us o f spe rm m o th er cell , wi th 8 ch rom o som es B , pa i ring

.

o f ch ro m o s o m es C , di visi o no f s pe rm m o th e r cell , sh o wi ng 4 ch ro
.

moso mes p a ssing i nto ea ch d a ugh ter cell D , fo r imma tu re sp erms ,

. u
ea ch wi t h 4 ch ro m os o m e s , res l tin
u g f ro m two di vi si ons o f the s pe rm
m o th er cell .

or m a terna l is a m a tte r o f ch a n
,
ce I no the r w ord s th e
.
,

th ous a nds of spe rm m oth e r cells p roduce i n th e species ,

u nde r discussi o n sixtee n di ff e re nt types o f spe rm cells

,

depe ndi ng up on thei r ch rom os omes I f let ters a re used

.
 M ATU R ATI O N OF T HE E GG 31 3

as i nthe p recedi ng p a ra g ra ph to desig na te ch rom os omes

the n two cells ma y be formed Wi th p a te rna l (a b c d ) , , ,

an d m a te rna l (A B C D ) respectively or these ma y

, , , ,

exist i n c omb ina ti on (A b c d a nd a B C D , , , , , , ,

With eigh t ch rom os omes sixtee n c ombi na ti ons a re p os

sible ; with f orty eigh t a s i n ma n th e p o ssible numbe r
-

ru n s i nt o th e milli ons Th is ra nd om a ss ortme nt

.
,

a s it is te rmed r esults i n a
,
wide v a ri a ti on a s will
be le a rned i na subseque nt ch a p te r a nd is o f the highest
imp ort a nce i nc onnecti onwith the ev oluti onO f a species .

M ATU RATI O N OF T H E EGG As a result of the a ccu mu

.

l a ti on of food subst a nces du ri ng the growth pe ri od

m
,

e a ch egg oth er cell bec omes a rel a tively l a rge b ody ;

but while it is thus supe rfici a lly u nlike th e c orresp ond
i ng sm a ll spe rm cell its h ist ory du ri ng th e m a tu ra ti on
pe ri od is th e s a me It u nde rg o es a reducti on divisi on
.

an d the re is the s a me ra nd om a ss or tme nt o f th e ch rom o

s omes but th e f o u r cells f ormed a s a result of the two
,

m a tu ra ti ondivisi ons a re ve ry u nlike i nsize This me rely .

me a ns i n th e light of thei r subseque nt beh a vi or th a t

, ,

th e th ree sm a ll cells with out y olk (Fig 7 5 ) a nd i nc a p a ble

n
)
.

o f fe rtiliz a ti on a re rudime t a ry eggs W hile the r em a i n

,
i ng ,

l a rge cell a l one is c a p a ble O f devel opme nt F o u r fu nc .

ti on a l spe rms devel op f rom on e m o the r cell while one ,

fu ncti ona l a nd th ree vestigi a l eggs a re de rived f rom the

egg m oth e r cell .

D ETER M I N AT I O N OF SEX —U p to th e p rese nt time it

.

h a s beena ssumed th a t the numbe r of ch rom o s omes i nthe

b ody cells of th e m a le a nd fem a le is the s a me but a t ,

le a st one hu nd red species of a nim a ls a re k now n whe re

th e m a le h a s one or a t m ost a ve ry few less th a n th e
fem a le This oc ca si ons a m odifi ca ti onof the m a tu ra ti on
.

divisi ons Let it be a ssumed th a t the b ody cells of the fe

.

m a le h a ve eight ch rom os omes a nd the m a le seve n (Fig .

77 ) a s a ctu a lly h a ppe ns i n s ome species The hist ory .

of th e m a tu ra ti ondivisi on s of th e egg is th e sa me a s th a t
a lr e a dy desc ribed In th e m a le on the oth e r h a n
. d the , ,
31 4 R E P R O D U C TI O N I N ANI MAL S
c onj ug a ti ono f th e ch rom os omes results i nth ree p a ir s a nd
a n odd o r u n p a i red one (c a lled a sex ch rom os ome ) a s
-

th o ugh Aa Bb C O a nd d we re the c ombi na ti on Up on

, , .

th ei r sep a ra ti on th ree ch rom os omes p a ss to one p ole o f

th e sp indle a nd f ou r to the othe r Th e two cells th us
.

FI G . 77 . D i a gra m
illus tra ti ng th e d ete rmi na ti o n of se x A
__
.
,

nu cl eus s pe rm m o th e r cell wi th 7 ch rom o s o m es pa i ri ng o f 6 ch ro

of ,

mos o mes wi t h o ne o dd (s ex ) ch ro m o s o m e C divisi o no f sp erm m o th er

.
,

c ell S h o wi ng 4 ch ro m o so m es p a ssi ng i nto o ne cell a nd 3 into the

o th e r
. D s e co n
,
d a nd l a s t divisio nO f sperm m o th e r cell res ul ti ng in
two (a s y e t imm a t u re ) f e m a l e p ro d u ci ng sp e rms wi t h 4 ch rom o s o m es
-

ea c h a n d two m a l e p ro du ci ng wi th 3 ea ch
-
.

formed now divide a nd o f th e fo u r S pe rm cells th a t a re

p rod uced two c ont a i nthe O dd numbe r a nd two th e eve n
 D E T E R M I NATI O N OF S EX 31 5

one I f a spe rm with f ou r ch rom osomes u nites with a n

.

egg which i nv a ri a bly c a rries f ou r a fem a le results ; the

, ,

fe rtiliz a ti onof a negg by a spe rm with th ree ch rom os omes

gives rise to a m a le The re a re thus m a le a nd fem a le
.

p roduci ng spe rm s but only one typ e of egg

,
.

Ina numbe r of o the r species th e re a re di ffe re n ces b e

twee n th e sh a pes of th e ch rom os omes a nd eve ni nm a ny
an im a ls wh e re these a re n ot a pp a re nt the re is evide nce
de rived from b reedi ng expe riments th a t h e re likewise a
ch rom os ome mech a nism regul a tes sex .
 C HAPTER XX VII

R E P R ODU CTI O N I N P LAN T S

SEX UA L AN D A S EX UAL R E PR ODU CTI ON I nth e p reced

.

ing ch a p te r it h a s bee n Sh ow n th a t a nim a ls h a ve two

typ es O f rep roducti on sexu a l a nd a sexu a l Pl a nts a ls o .

exhibi t b oth types I t is p rope r a t th is p oi nt to re emp h a

.
-

size th e fa ct th a t while b oth meth ods secu re the p erp etu

a ti ono f the ra ce a n d ma y a ls o se rve to multiply it on e of ,

them — the a sexu a l — me rely rep roduces th e i ndividu a l

with out esse nti a l ch a nge bu t th a t th e o the r
,
the sexu a l
not only p roduces a new individu a l bu t a ls o e nd ows it
with ch a ra cte rs f rom b o th p a re nts In this wa y it ma y
n
.

a d usu a lly d o es h a p p e n th a t the n ew i ndivid u a l is not

ex a ctly like eithe r p a re nt .

TY P ES OF R EP R OD U CT I O N I N P LA N TS S i nce pl a nts ex
.

b ibit a f a r g re a te r v a riety i n th e de ta ils o f rep roducti on

th a n a nim a ls it f oll ows th a t no one ex a mple will suffice
to give a n a deq u a te p ictu re M ore ove r a lth o ugh o u r
.
,

ch ief i nte rest is with S eed Pl a nts it is quite imp ossible

,

to u n de rs ta nd th e sig nific a nce o f th ei r rep roductive p roc

esses with o u t s ome k nowledge O f th e ev oluti o a ry st a tes n
th ro ugh which they h a ve p a ssed a nd which a re still ex
h i bi ted by th e l o we r orde rs o f pl a nts For this re a son
.

this ch a p te r wi ll de a l i n o rde r of devel opme nt wi th the

f oll owi ng five types o f rep roduc ti on: (a ) By resti ng sp ores
an d swimmi ng S p o res a m ong th e l owest pl a nts — Al g ae
an d Fu n gi ; (b ) By simple types o f sex cells or g a metes

i n th e s a me l owe r pl a n ts ; (c ) By a l te rna ti on o f a
sexu a l a nd a na s exu a l ge ne ra ti on i nM o sses a nd Fe rns ;
d
( ) By seeds i n t h e h ighe r pl a nts ; a nd (e ) By v a ri o us
so rts o f p u rely veget a tive d ivisi on o f pl a nts such a s ,

bul bs cut t ings g ra f ts buds etc

, , , ,
.

31 6
 A S E X UAL R E P R O DU CTI O N 31 7

A S EX UA L R E P R OD U CT I O N IN E r TH E
a re AW —The e
m a ny pl a nts of much S imple r org a niz a ti on th a n th e
Fl owe ri ng Pl a nts the only ones with which m ost pe rs ons
,

a re a t a ll f a mili a r Th e l owest a nd simplest pl a nts b e

.

l ong to the Algae These pl a nts a re typic a lly a qu a tic ;

.

they h a ve simple b odies of no ve ry high deg ree of spe ,

ci a l i z a ti on a n
,
d simple meth ods of rep roducti on S ome .

o f th e l owest do n o t h a ve a ny sexu a l meth od a t a l l .

Two or th ree ex a mples will m a ke this cle a r .

Osci lla tort a (Fig 78) is a n a lg a of b l ue g reen c ol or

'

-
.

fou nd i nsh a ll ow temp ora ry p ools i nl a kes a nd st re a ms of ,

st a g na nt Wa te r a b out th e o utlets of sewe rs a nd oth e r

,

simil a r pl a ces whe re the w a te r c ont a i ns conside ra ble

a m ou n ts o f filth Its h a bits of life a re h oweve r
.
, ,

not the immedi a tely i nte resti ng subj ect but its meth od ,

of rep rod ucti on As th e illust ra ti ons S h ow a single

.
,

pl a nt c onsists of a simple row or fil a me nt of S h ort

cyli nd ric a l cells pl a ced with th ei r fla t su rf a ces i n
c ont a ct A c a reful i nspecti on will f a il to reve a l a ny
’

esse nti a l di ff e re nce a m ong th e cells except th a t th e ,

te rmi na l Ones h a ve th ei r free end rou nd ed ou t i n

ste a d of fla t Any cell or g ro up of cells ma y be det a ch ed
.

from the othe rs with out m a ki ng a ny di ff e re nce i n thei r

devel opme nt E a ch one g oes onm a nuf a ctu ring its f ood
8

an d g rowi ng i ndepe nde ntly o f th e rest A s e a ch cell .

g rows i t re a ches a p o i nt whe n divisi on o ccu rs m aking ,

two ou t o f i t This i nc re a ses the length of th e fil a me nt

. .

Any distu rb a nce such a s f requently occu rs i n the w a te r

,

i n wh ich it g rows ma y b re a k th e fil a me nt i nt o two or

,

m ore pieces Obvi o usly i n this pl a nt the re is no re a l

.

distincti on betwee n g rowth a nd rep roducti on Any cell .

divisi on ma y a cc omplish eithe r or b o th If th e p ond .

d ries up the pl a nts a re d ried a ls o F ortu na tely the p ro

'

top l a sm is a ble to e n du re such t re a tme nt with o ut killi ng

a l l o f the cells th ough m ost O f them do no t su rvive
,
.

Th ose whi ch do live need only th a t th e necess a ry he a t ,

light a nd m oistu re be a g a i n fu rnish ed them to begi n

,
31 8 R E P R O DU CT I O N IN P LAN T S

thei r life p rocesses j ust whe re they h a d left o ff P re .

s u ma bl y they n eve r bec ome so e nti rely dry a s to st op a ll

me ta b olism .

N os toc (Fig 7 9) is a n o the r blue g ree na lg a rel a ted to

.
-

O scill a t ori a I t c a rries oni ts w ork a nd g rowth i nm uch

.

the s a me m a nne r but sh ows two i nte resti ng a dv a nces in

FI G . 78 O s cill a to ri a
.
-
l A sh ort s ecti on o f a fil a m en
. . t sh ow
in g cel ls (1 , 2, 3 , 4 ) i n p ro cess o f divisio n c b , t he cen . tra l b o dy
. .

co rre sp o n di ng to the nu cleus of high er pl a nts ; ch, t he ch ro m a to ph ore

o r re gi o no f the cy to pl a s m i nwhi ch t he bluish - gre e ncol o ri n g m a tte r
1 8 l o ca ted 2 A si n
.
gl e cell o na l a rger s ca l e seenfro m th e end La bels
.
.

a s I nfig 1 . 3 A fil a m en
. t i nwhi ch o n
. e cell h as di e d c rea ti ng a w ea k
pl a ce a t whi ch t he fil a m ent will b rea k This is t he usu a l wa y i n
.

wh i ch t his pl a nt m u l tipli es the num be r o f fil a m ents 4 A si ngle . .

fi la m ent d ra w n a t 5 mi nu te i nte rv a ls to S h ow th e a m o u nt a nd type

-

of m ov em e n t 1 2 3 4 i nfig 4 in. di ca te su ccess ive p osi ti ons

, , , . .
 A S E X UAL R E PR O DU C TI O N 31 9

its meth ods of rep roducti on Inthe fi rst pl a ce not a l l th e

. ,

cells a re a like E x a mined a t a f a vora ble pe ri od i nits de

.

v el op men t th ree s orts o f cells will be f ou n d The l a rge r .

numbe r o f cells a re of th e ordi na ry veget a tive type wh ich

go r igh t a l ong pe r f o rmi ng thei r w ork o f f oo d m a nuf a ctu re ,

3 Gloeotr ichia

FI G 7 9 — O th er B lu e—
. . green Al gae 1 N osto c fil a m ents sh ow
. .

ing di vidi ng cells , h etero cys ts , h, whi ch s erv e t o b rea k th e fil a m ents

into sections a nd so mul tiply th eir numb er, a nd enl a rged S pore
cel ls , s tored wi th abu nda nt f o o d a nd surro und ed by a fi rm w a l l
for p ro tecti on du ring th e re s ti ng p eri o d 2 Gl o e o ca ps a , a S impl er
. .

f orm whi ch f orms sm a ll irregu l a r gro ups lls ins tea d of fil a ments
of ce ,

th e wh ol e en cl os e d i na m a ss o f g el a ti no u s w a ll m a teri a l 3 Gl o eo
. .

tro ch i a a s o m ewh a t m ore compli ca t e d type wi th v ery l a rge sp ore s

, ,

sp . f orm ed by th e u ni on of sev era l a dja cent cells T h e h etero cys t

, ,
.

forms next to th e sp ore a nd so s erves to deta ch th e S pore f rom th e

fil a m ent T h e sev era l figu res S h ow va ri ous s ta ges i nth e grow th of a
.

fil a m ent .

g rowth nd cell divisi on At i nte rv a ls i n th e fil a ment

,
a .

a re to be see n a secon d typ e of cells which a re l a rge r ,

h a ve thi ck w a lls a nd ve ry sc a nty cell c onte nts S ome of -

.
3 20 R E PR O DU C TI O N IN P LAN T S

them will a ppe a r to be enti rely empty These cells .

(he terocys ts ) h a ve th e fu ncti ono f b re a ki ng the fil a me nt

i nt o sh orte r secti ons As divisi ono f cells p rog resses a fil a
.

me nt bec omes l onge r a nd l onge r B ut h e re a nd the re

.

s ome cell tu rns into a hete rocyst This s ort o f cell is .

a tt a ched l oo sely to th e ordi n a ry cells o n eithe r side o f

i t with th e result th a t a S ligh t distu rb a nce such a s th a t ,

d ue to be ndi ng to a cc omm od a te g rowth ma y e a sily b re a k ,

th e fil ame nt Th e thi rd typ eo f cell is a ls o c onside ra bly

.

e nl a rged bu t i nste a d o f bei ng empty i t is de nsely fi lled

with p rot opl a s m a nd st ored f ood I ts w a lls a ls o bec ome
.

thick a nd so ch a nged ch emic a lly th a t they resist d ryi ng .

These a re c a lled res ti ng sp ores bec a use th ey do no t

divide u ntil a fte r th e o th e r cells i n th e fil a me nt h a ve
d ried or fro ze n Th is kills th e ordi na ry cells bu t d oes
.

not kill the resti ng sp ores On the re est a blishme nt of

.

g rowi ng c onditi ons the l a t te r resume g rowth a nd cell

d ivisi on a nd m a ke new fil a me nts I n this pl a nt th e re.

‘
i s see n to be a disti ncti on betwee n g rowth a nd rep ro
ducti on C ell divisi o n p rod uces g rowth but d oes not of
.

itself b ri ng a b ou t rep rod uc ti on Th e d iff e re nti a ti on of

.

s ome cells to form h ete ro cysts d o es h oweve r rep roduce

or mul tiply th e n umbe r o f pl a nts du ri ng th e growi ng sea
son Th e di ffe re n
. ti a ti onof th e resti ng sp ores t a kes c a re
o f rep rod ucti onf oll owi n g u nf a v ora ble we a th e r c onditi ons .

O bvi ously this a rra n geme nt h a s ce rt a i n disti nc t a d va n

t a ges to th e pl a nt for i t m a kes m ore ce rt a i nth a t th e fil a
me nts will be b roke n f reque ntly a nd th a t rep roducti on
wi ll occu r a fte r d rou h t .

U l o thrizr (Fig 80 ) is a S imple fil a me n

. t o us g ree na lg a
fou nd i nfresh w a te r st re a ms a nd l a kes E a ch fil a me nt is
.

a tt a ched a t on e en d to a stick o r st o ne a n d f ree a t th e

o the r en d The b a s a l a tt a chi ng cell is spe ci a lized for th e
.

p u rp ose a nd the tip cell is rou nded or s ometimes p oi nted .

Th e rem a i ni ng cells a re a t fi rst a ll simil a r Ea ch m a kes .

its ownf ood g rows a nd d ivides Whe nthe fil a me nt h a s

, ,
.

re a ched a suit a ble st a ge o f devel opme n t the re occu rs a

 A S E X UAL R E P R O D U C TI O N 3 21

diffe re nti a ti on a m ong th e cells A few me rely ce a se to.

grow but m ost of th em devel op a numbe r of sm a ll m o tile

,

rep roductive cells Th is is b ro ugh t a b out by a se ries o f

.

divisi ons of th e nucleus u ntil the re a re a b out 8 (or s ome

times 1 6 ) i n e a ch cell The cyt opl a sm the n divides up
.

i nt o a nequ a l numbe r of p orti ons so th a t e a ch nucleus is ,

su rrou nded by a sm a ll a m ou nt of i ndividu a lized cyt o

pl a sm N ext e a ch of these u nits u nde rg oes a p rocess O f
.

FI G 80
. . . vegeta t ive fil a m ent one of wh ose
U l o th rix A, p a rt o f a
cells is und ergoing di vi si on B , a
fil a m ent in whi ch th e ce lls a re
.

di vidi ng to f orm f o ur la r ge swimming S p ores C a S i ngl e swimm ing

.

sp ore dra wn to a la rger s ca l e D a swimming sp ore b eginni ng to

.
,

devel o p E a fur th er sta ge of dev el o pm ent F a nd G sm a ller

.
, .

swimmi ng sp ores devel oping i nto new fila m ent .

ro u ndi ng up i nt o a s ort of egg S h a ped b ody At th e -

.

sm a lle r end th e cy topl a sm p uts forth fine cili a — two i n

numbe r an d these new cells begi nto m ove a b out i nside
the Ol d cell w a ll P rese ntly th ey b re a k th rough th e w a ll
-
.

an d esc a pe i nt o th e w a te r They now begi n a series of

.

ra pid swimm in g m oveme nts a nd eve ntu a lly e a ch one con

3 22 R E P R O DU C TI O N IN P LAN T S

t a c ts with a su i ta ble obj ect to whi ch i t bec omes a tt a ch ed

by th e cili a ted end The cili a a re then wi th dra wna nd
'

thicke r blu nte r p rot rusi ons of the cell b ody a re ex te nd ed

o ve r the subs tra tum Th ey seek o u t a nd m a ke u se of its
.

i rregul a ri ties of f o rm to secu re a fi rm h old Th e p rotru .

si ons bec ome c ove red with a cellul ose w a ll a t th e s a me

t im e th a t the wh ole cell is f ormi ng its w a ll Whe n .

a tt a ch me n t h a s bee nc ompleted th e cell begi ns to el ong a te

an d p rese ntly d ivides The two new cells rem a i nj oi ned
.

an d m a ke a little two celled fil a me nt -

A c o nti nu a nce O f
.
'

cell divisi on el ong a tes th e fil a me nt u ntil i t p rese ntly

re a ch es a deg ree O f m a tu rity whe nit will i nits tu rnp ro

duce m o tile rep roductive cells O n a cc o u nt o f th ei r

.

numbe r a nd m oveme nt these a re c a lled swa rm sp o res o r

s ome times z oosp ores
’

E dogon i u m (Fig 81 ) is a s omewh a t m ore h igh ly d e

.

v el o p ed g ree na lg a th a n Ul oth rix but will se rve to illus

t ra t e th e s a me phe nome non o f rep roduc ti on by non

sexu a l sw a rm sp ores I t to o is a tt a ch ed a t the b a se a nd
.

f ree a t th e a pex I t g rows by cell divisi on Occ a si ona lly

. .

(o r f reque ntly if the c onditi ons a re j ust righ t ) s ome o f

its cells withd ra w thei r p rot opl a sm f rom c ont a ct with th e
e ncl osi ng w a ll a nd ro u nd up Ea ch cell th e n p rod uces
.

a si n gle sw a rm sp ore The Ol d cell w a ll b re a ks a nd the

.
-

sw a rm sp ore esc a pes a nd swims a b ou t vig orously by

me a ns of a c rowno f cili a a t th e sm a lle r end It fi na lly .

settles d ow ni nth e s a me wa y desc ribed for Ul oth rix a nd

g ro ws i nt o a new fil a me nt .

E S S E N T I A L S O F A S EX UA L R EP RO DU CT I O N Th ough .

the re a re a numbe r o f o the r d iffe re nt f orms o f non sexu a l -

rep ro du cti ona m o n g th e l owe r pl a nts it is p rob a ble th a t

,

these illust ra ti o n s will suffi ce to m a ke cle a r its esse nti a l

fe a tu res
. All a g ree i n th a t the rep roductive cells
wh ethe r th ey a re j ust ordi na ry b ody cells o r speci a l sw a rm
sp o es
r — h a ve a rise n th ro ugh ordi na ry cell divisi on a nd
h a ve ide nti ca lly th e s a me ki nd o f nuclei Ea ch a nd eve ry .

cell of th ese p l a nts whethe r ord i na ry veget a tive or rep ro

,
 A S E X UAL R E P R O D U C TI O N 3 23

du cti v e cells h a s th e s a me numbe r a nd ki nd o f ch rom o

,

s ome s I t na tu ra lly results f rom this th a t the su ccessive

.

ge nera ti ons a re ve ry S imil a r i ndeed no v a ri a ti ons ordi na r ,

ily occu r excepti ng such a s a re due to th e di ff e re nt envi

n
ro men ts u n de r which the successive ge ne ra ti ons ma y

FI G 81 . . (Edogonium . l ,
sh owi ng th e wa y i nwhi ch th e
2, 3 , 4 ,
con ten ts o f a cell tra n sf orm t h ems elv e s i nto a swimmi ng sp ore a nd
e s ca p e from th e ol d 0 6 1 1. 5 6 S h o w swimmi ng S p ores f orm e d f ro m
, ,

th e f ert il iz e d e gg sh o w ni nfig 83 . 7 8 9 S h ow th e germ i na t i onof

.
, , ,

swimmi ng sp ores to f orm new fila m ents .

g row The a dv a nt a ge of this type lies i n th e extra or

.

dina ry ra pidity O f multiplic a ti on e na bli ng th e pl a nt

to occupy a l l a v a il a ble sp a ce I t d oes not of itself give
.

r ise to a n v a ri a ti on
y .
3 24 R E P R O DU C TI O N I N PLANT S
S EXUA L R EP R OD U CT I O N by me a ns G a metes — eggsof

an d spe rms In m ost a lg ae a n

. d ne a rly a ll highe r pl a nts
the re is a sexu a l meth od of rep roducti on S ome pl a nts .

h a ve b o th me th ods A t this p oi nt it is necess a ry to de

.

fine m ore p recisely wh a t is me a nt by sex This ca nbest .

be d one by studyi ng a S imple form of it .

U lo th ri x h a s a ls o a sexu a l meth od O f rep roducti on(Fig .

I t is p a rticul a rly suited to i llus tra te th e ori gi nOf sea:

FI G 82
. .
—Ul th ix sexua l
o r , produ cti on A fil a m ent in whi ch
re .
,

th e cells a re fo rmi ng a b o u t 1 6 m o til e sex cells or ga m etes B C D E

-
.
, , ,

S h ow th e ga m e tes fusi ng i np a i rs F th e res ti ng sp ore d ev el o pe d fro m

.
,

th e fusi on o f th e two ga m etes .G th e germi n

, a ti on o f th e res t ing
S po re to p ro d u ce a n ew fil a m en t .

bec a use it is e a sy to obse rve th a t th e sw a rm Sp ores s ome

times t a ke p a rt i nsexu a l rep roducti on It will be rec a lled .

th a t i nUl oth rix e a ch cell p roduces eith e r 8 or 1 6 sw a rm

sp ores Th e more nume rous ones a re p rop orti ona tely
.

sm a lle r S ometimes th ey a ppe a r to be too sm a ll to p ro

.

duce a new pl a nt a l one a nd u na ided ; a cc ordi ngly th ey

fuse i np a i rs Th e u su a l sex cells a re p roduced i nth e s a me
.

m a nne r a s th e sw a rm sp ores bu t a re still m ore nume rous ,

1 6 to 64 i nnumbe r Th ese sm a ll m o tile cells a re c a lled

.

ga me tes or sea: c el ls a n d a lwa ys fu se i n p a i rs before

 S E X UAL R E P R O D U C TI O N 3 25
4

g rowi ng i nt o new fil a me nts F rom this i t a ppe a rs a s if

.

sex i n its simple st f orm h a s origi na ted by a ch a nge i n

the beh a vi or of ordi na ry sw a rm sp ores a nd th a t the
esse nce o f the p ro cess is th e fusi on o f two rep roductive
cells I t is a ni nte res ti ng f a c t th a t these g a metes a lw a ys
. .

S h ow a decided p refe re n ce for fusing with th o se f rom

s ome o ther fil a me nt ra the r th a n with th o se from the
m
s a e cell or fil a me nt Th e p recise ch emi cO physic a l re
.
-

l a ti o ns a t th e b ott o m of this beh a vi or a re no t yet k now n .

Th a t i t is a n a dv a nt a ge h oweve r is p retty cle a r f rom

, , ,

the f a ct th a t so m a ny pl a nts a nd a nim a ls h a ve ev olved

th is meth od of rep rod ucti on I t ce rt a inly gives a gre a te r
.

O pp ortu n ity for v a ri a ti on by b ri ngi ng t ogether th e ch a r

a cters of two p a re n ts i nste a d of one .

G dogon i a m (Fig 83 ) h a s a meth od of s exu a l rep rodu c

.

ti oni nwhich the egg (i e fem a le ) g a mete is ve ry much

. .

l a rge r th a n the spe rm (i e m a le ) g a mete A p a rticul a r

. . .

cell or cells i na fil a me nt swells up so a s to bec ome ne a rly

Sphe ric a l i n ste a d of th e el ong a ted cyli nd ric a l f orm fou nd
i nthe ordi na ry veget a tive cells I ts conte nts the nwith .

d ra w slightly f rom the cell w a ll a nd rou nd up to f orm a n

-

egg g a met e A sp ot on th e Wa ll s ofte ns a nd p rese ntly

.

forms a pe rf ora ti onth rough wh ich th e spe rm g a mete ca n

e nte r In s ome o th e r p a rt of the s a me fil a me nt or i n
.
,

s ome species i n a nothe r fil a ment s ome cell divides

’

, ,

ra pidly to f orm a n umbe r of ve ry sh ort cyli nd ric a l cells .

Ine a ch of these th e nucleus divides once Th e c yt opl a sm .

is the ndivided betweenthese two nuclei a nd e a ch ha l f b e

c omes org a nized to form a spe rm Wh e n these spe rms .

a re m a tu re they esc a pe th ro ugh th e b ro ke ncell w a ll a n d -

swim a b o u t by me a ns of a c row n of cili a a t th e sm a lle r

en d
. They a re a tt ra cted to the p e rfora ti on i n th e w a l l
o f the cell (o Ogon i u m) c ont a i ni ng the egg p rob a bl y b y ,

me a ns of s ome subst a nce di ffusing a w a y from it a nd se rv

i ng a s a di rective stimulus This is a c a se O f wh a t wa s
.

ca lled a t ropism i n C h a pte rs X I I a n d X I V whe re we ,

le a rned th a t a v a riety of exte rna l c onditi ons ma y a ct a s

‘

a ac R E P R O D U CTI O N IN PL AN T S

FI G 83
. . (Edo go n i um, se xu a l re p rod u cti on 1 a fi la m ent
. .

two e nl a rged cells co nta ini ng th e f em a l e se x cell or egg A t ta ch e d

-
.

to t h e sid e o f th e egg a re sm a ll m a l e pl a n ts whi ch i n this spe ci es

,

p ro du ce th e ma l e se x cel ls o r sperms 2 3 4 sh ow the d ev el o pment

- .
, , ,

o f th e e gg 5 a nd 6 S h o w t h e f e rtiliz a ti o no f th e e gg i na pl a n
. t (Bul
boc haa ta ) rel a te d to Oed o go n ium 7 sh ows the d ev el o pm ent of
.

spe rms ina nord ina ry fil a m ent whi ch i ns o m e speci es o f Oed o gonium
,

ma y a ls o co nta i ne gg cells 8 sh o ws sm a ll swimmi ng spo res whi ch

.

inthi s spec i es d e ve l o p the s o rt o f sma l l m a l e pla nts see ninfig 1 .

 THE DIFF E R E N TIATI O N OF S E X 3 27

di rective stimuli to c a use wh ole org a nisms or thei r org a ns

to m ove or be n d i nresp onse to stimuli The spe rm fi nd s .

the Op eni ng enters a nd fuses with th e egg As s oona s

‘

.
, ,

this fusi onh a s occu rred a ch a nge c omes a b out i nthe egg
r esulti n g i nth e form a ti onof a memb ra ne ove r its su rfa ce
p reventi ng the e nt ra nce of o the r spe rms This is a .

simple c a se of ch emic a l regul a ti on .

T H E D I FF ERE N TI AT I O N OF SEX In Ul o th rix we ca n

.

spe a k o f sex a lth ough the re is no disti ncti onbetwee nthe

sexes N ot only ma y th e two fusi ng g a metes c ome from
“

the s a me i ndividu a l bu t th ey a re b o th m otile a nd i ndis

tin u i sh a b l e i n eve r y r espect I n (Ed o go ni u m the e is
-

g . r

not only Sex but two sexes a t le a st i n respect to th e

,

g a metes th emselves One is a lw a ys sm a ll a nd highl y

.

m otile while th e o the r is l a rge sto red with f ood non

, , ,

m otile a nd rem a i ns In the cell wh ere it wa s f ormed In

, .

some ca ses the re is a disti n cti on of sex betwee n th e fil a

’

m en ts such th a t one fil a me nt forms S pe rms only a nd the

o th er eggs on ly In still o th e r sp eci es of (Edogoni u m
.
j

th e ma l e fil a me n
~

t is ve ry sm a ll c omp a red to th e fem a le .

Inthese Species the re a re a ls o two ki nds o f sw a rm sp ores

diff e ri ng insize The sm a ll ones p roduce m a le fil a me nts
.

an d the l a rge ones fem a les .

O the r illust ra ti ons of th e s a me s or t of di ffe re nti a ti on

o f sex ca nre a dily be f o u n d a m ong th e Alg ae a nd Fu ngi
’

M ore ove r it is e a sy to exte nd the ra nge of obse rv a ti on

,

an d fi nd illust ra ti ons i nte rmedi a te betwee n th ese so a s ,

to S h ow th a t sex h a s a rise n m a n y times a m ong pl a nts

th rough a ch a nge i nth e h a bits o f sw a rm sp ores a nd th a t
two sexes h a ve bee ndevel oped by a g ra du a l l o ss of m o ti on
an d i nc re a se i nsiz e a n d st ored f ood by one g a mete to de
v el op a n egg P ra ctic a lly a l l th e h ighe r pl a nts h a ve
.

sexu a l rep roducti on fu nd a me nt a lly like this but with

a dded c omplic a ti on s i n th e wa y the g a metes a re forme d
or i n the seque n ce O f devel opment du ri ng th e e nti re life
cycle .

A LTER N ATI ON OF GEN ERATI ON S i s f ou n d insome Al gae

3 28 R E P R O DU C TI O N I N PLAN T S
and Fu ngi a nd is a regul a r p a rt o f th e life cycle o f a ll
,

high e r pl a nts I t ca nbe m o s t re a dily u nde rst ood by con

.

si de ri ng the life h ist o ry o f a m o ss a nd the n c omp a ri ng

this wi th a fe rn .

M oss pl a nts (Fig 84 ) a re g ree n indepe nde nt l a nd

.

p l a nts c a p a ble of m a nu fa ctu ring thei r own f ood S ome .

FI G . 84 .
— M osses . m a l e ga m e to phy te wi th a clus ter of
1 .

an th eridi a i na ro s e tte l ea ves a t t he ti p 2 T he a nth eridi a wi th

of . .

t h e en cl o si ng l ea v es rem o ve d 3 A fem a l e pl a nt wi th a s po ro phy te

. .

a t ta c h e d to t h e righ t b ra n ch 4 A gro up o f a rch e go n

. . i a f ro m a ti p
lik e the one to whi ch the sp oro phy te O f fig 3 is a tta ch e d Wh en . .

t h e e gg i n t h e b a e o f th e a rch e g o n
s ium is rea dy fo r fe rtiliz a ti on a
S pe rm f ro m a na n th e ridi a ] b ra nc h swims i n the t hi n films o f w a te r

th a t a re p res en t o nth e m o ss pl a n ts a f te r ra i no r dew a n d goes d o wn

th e l o n g nec k of the a rch e go nium a nd u ni te s wi th the e gg From .

this f e rtil i z e d e gg t h e sp oro phy te a ris e s a n d rem a i ns a t ta ch ed by

i ts f o o t d erivi ng a ll i ts w a te r a nd s a l ts a nd p a rt o f i ts fo o d f ro m
,

th e l e a fy ga m e to phy te 5 T he upp er p a rt o f t he spo ro phy te ca ll ed

. .
,

t h e ca p s u l e o pe n a nd sh edding sp o res
,
.

time du ri ng thei r g rowth the re devel ops usu a lly a t th e ,

stem tips a g ro up o f sex orga ns The s o rt i nwhich spe rms

, .

a re f ormed a re ca lled a ntheridia E a ch one c o nsists o f a .

Sh ort st a lk a nd b ody The l a t te r h a s a no utside l a ye r of

.
 AN E XTR A G E N E R ATI O N 3 29

p ro te ctive cells a nd a m a ss o f inne r cells which f orm the

spe rms Ea ch spe rm is a sm a ll m otile cell swimmi ng by
.

mea ns of cili a The eggs a re f ormed i n org a ns c a l led

.

a rchegon i a sh a ped l ike l ong sle nde r fl a sks

,
Th e a ec om .

p a nyi ng illust ra ti ons will give a nide a of thei r sh a pe a nd

st ructu re E sse nti a lly they c onsist of th ree p a rts : st al k
.
,

ve nte r a nd neck Th e neck is l ong a nd sle nde r with a n

,
.

o utside l a ye r of p ro tective cells a n d a si ngle a xi a l row O f

cells k now na s th e neck c a na l cells The e nl a rged middle
-
.

p a rt c a lled th e ve nte r h a s one or m ore l a ye rs of p ro te o

tive cells onthe ou tside The row O f neck c a nal cells is .
-

c onti nued d ow n int o th e ve nte r by two cells know n re

sp ecti v el y a s th e eggx cell (a t the b o tt om ) a n d the ve nt ral
c a na l cell Wh e n the egg is ripe the ve nt ra l a nd ne ck
.

c a na l cells die a nd tu rn i nt o a s ort o f mucil a ge or j elly ,

which swells a nd f orces the top off th e neck This sub .

st a nce oozing ou t i nt o th e w a te r di rects th e sp e rm to the

openn eck Wh e n one fi nds its Wa y d ow n the neck a nd
.

fuses with th e egg a fe rtiliz a ti on memb ra ne is f ormed

“

a b o u t the egg a n d the mucil a ge h a rde ns a nd bl ocks the

p a ss a ge .

AN E X TR A G E N ER AT I O N — I t is p rope r a t this p oi nt
to c a ll a tte nti onto th e st riki ng peculi a rity o f these h ighe r
pl a nts Inth e pl a nts we h a ve bee ndiscussi ng he ret of ore
.

a fe rtilized egg ge n e ra lly devel ops a g a i n i nt o th e s ort of

pl a nt which p roduced it But h e re the re is a regu la r a lter
!

na ti onof two sor ts of p la nts i n the li fe cy cle The fe rti

’

l i z ed egg rem a i ns i n th e ve n t e r o f the a rch eg on ium a nd

begi ns to g row a nd divide E ventu a lly this y ou ng pl a nt .
,

whi ch we c a ll a nembry o rup tu res the w a ll of the e ncl os ,

i ng st ructu re a nd p ushes up i nt o th e a i r Whe n it is .

m a tu re it is f o u nd to consist o f th ree p a rts A t the .

b o tt om is a s ort o f f oo t embedded i n the end o f the

b ra nch (Fig wh ich b ore th e a rch eg onium Ab ove
. .

th a t is a sle nde r st a lk a ni nch or so l ong a nd be a ri ng a t its

top a ne n l a rged b ody know na s th e c a psule Th e f oo t is .

a norga nwh ich se rve s a s a me a n s of a tt a chment a nd a t

330 R E P R O D U C TI O N I N PLANT S
the sa me time a bs orbs supplies of w a te r a nd diss olved
f ood f rom th e origi na l pl a nt The c a psule a t th e top de
.

v el op s a multitude o f sm a ll wi n d b orne sp ores -

The .

st a l k se rves to lift the ca psule up wh e re th e sp ores ca nbe

a dv a n t a ge ously sc a tte red I t sh o uld be no ted th a t wh ile
.

this sm a ll a tt a ched pl a nt is depe nde nt onthe fi rst ge ne ra

ti on it bec omes g ree n a s s oon a s it is pushed o u t
,

o f th e e n cl osi ng st ructu re a nd is exp o sed to ligh t I t ca n .

,

the ref ore m a nu f a ctu re a p a rt a t le a s t o f i ts c a rb ohydr a te

,

f oods .

G A T P YT
M E O H E A N D S P ORO P H Y T (Fig
E — I n th e .

a b o ve a cc o u n t it is to be rec ognized th at a g ree n i nde

p e nde nt m oss pl a nt p roduces th e g ametes (eggs a nd
S pe rms ) a n d th a t the fe rtilized egg g rows i n t o a sm a ll
pl a nt which i nrespect to w a te r s a lts a nd p a rt o f its foo d
, ,

is p a ra si ti c on th e fi rst a nd which p rod uces a i r b orne -

sp ores which g row i nt o the g a met ophyte or sexu a l ge ne ra

ti on Th e fi rst st a ge is c a lled the ga me top hy te bec a use
.

it p rodu ces g a metes R ep roducti onis b rough t a b ou t by

.

the regul a r a lte rna ti ono f th ese two ge ne ra ti ons E ve ry .

a l te rna ti on O f ge n e ra ti ons i n th e highe r pl a nts i nv olves

esse nti a lly th is s a me cycle .

F ER N S S h ow th e s a me a lte rna ti o no f ge ne ra ti o ns a s do
m osses The re is h oweve r th is st riki ng di ff e re nce
.
, ,
the ,

fe rn pl a nt fa mili a r to eve ry one is th e sp orophyte

( Fig 8. 5 ) ge n e ra ti o n Th . e g a met o phyte (Fig 86 ) ge n .

c a ti o n is so sm a ll
r — a qu a rte r o f a n i n ch or less in
d i a mete r —th a t it is ordi na rily ove rl ooked excep t by
th ose wh o k now a b o ut i t a nd especi a l ly l ook for it An .

o th e r imp orta n t di ff e re nce is th a t whe re a s th e m oss

,

sp orophyte is sm a ll a nd a tt a ch ed (p a ra sitic a lly for a l l

it s w a te r a nd p a rt of its f ood supply ) to the gree n
g a me tophyte the fe rnsp o rophyte is p rop orti ona tely ve ry
,

l a rge g ree n a nd th e ref ore i ndepe nde nt

, ,
.

T H E S TR U CTU R E O F F ER N S For p rese nt p u rp o ses i t

.

is necess a ry to c onside r a few O f the det a ils o f st ructu re .

Inthe first p l a ce it is cle a r f ro m the m ost c a s u a l obse rv a

 T HE S T R U C TU R E OF FER N S 331

ti onth a t th e fe rnis a much l a rge r and m ore efficie nt pl a nt

th a n the m oss I t h a s l a rge le a ves a nd a n a mple roo t
.

system By cu tting sec ti ons of th ese org a ns a nd of th e

.

stems a c orresp ondi ng c omplexity a nd efficiency of org a n

i z a ti on is reve a led i n thei r i nne r st ructu re The sp ores .
,

wi th whi ch we a re chiefly c once rned a re c omm only p ro

‘

du ced on the b a cks of the le a ves Th e a cc omp a nyi ng

.

FI G 85
. .
l
T h e S p oro phy te Oi sexu a l pl a nt o f th e f ern 1 a wh ol e
. .

pla nt sh owing (a ) th e s te m (b) th e roots a nd (c) th e l ea v es 2 th e

, , . .

b a ck of a lea fl et sh owing cl us ters of sp ora ngi a a nd th e b ra nch ed

”
v ei ns thr o ugh whi ch w a ter a nd diss olved fo o d a re tra nsported .

3 a s ect i o nt h ro ugh a clus ter o f sp ora ngi a ona diff erent s ort o f fern
.

wi th a fla p lik e co ver or indusium 4 a S ingl e sp ora ngium sh owi ng

”
-
. .

th e s ta lk a n d th e r ing lik e an nulus a ro und th e ri m m a d e of

“
-

thi ck en ed cells wh o s e funct i on is to o p en th e sp o ra ngium a n

,
d dis
ch a rge th e rip e sp ores t h ro ugh t h e m o u t h lik e o pen
-
i ng a t th e l ef t .

5 a cro ss s ect i on o f th e s tem sh owi ng two d a rk b a n

. ds of s trong
s t iff eni ng tissu e a nd s evera l ro und or o va l b undl es of condu cting
“

illust ra ti ons S h ow v a ri ous det a ils of sp ore p roducti on At .

p rese nt we a re c once rned only with th e f a cts th a t th e

sp ores a re a ll a like a nd th a t they f a ll onm oist s oil , where
 3 32 R E P R O D U C TI O N I N PL ANT S
th ey give rise to the sm a ll but g ree n a nd i ndepe nde nt
g a met ophytes The det a il s of st ructu re o f th e l a tte r a re
.

a ls o sh ow ni nthe illust ra ti on s .

H ETERO S P ORY is a te rm used by b ot a nists to sig nify th a t

a pl a n t p roduces S p ores o f two di ff e re nt sizes (Fig .

Inm o st fe rns th e sp ores a re a ll a like I nS i ze a n d give rise

i ndi ff e re ntly to g a met ophytes of eithe r sex or s ometimes

‘
HR D u

FI G . 86 . T h e Ga m e to phy te
s exu a l genera ti on o f th e fern
or .

1 sp o res
. 2 a germi n a ti n
. g spo re wi th a sl e nd er ro o tl e t a nd a thi cker
.

bra nch wh i ch will d e v e l o p l a te r i nto a fla t s exu a l pl a nt su ch a s

.

3 a nd 4 n 4 a s exu a l pl a n
. t wi th a n
. th eridi a (a ) a nd ro o tl e ts o r rh i ,

z o id e (r) At (3 p ) th e S p o re is sh o wnf rom whi ch the wh o l e h as de

.

v e lo pe d 5 a p a rt o f a m a l e s e xu a l pl a n
. . t wi th two a n th erid ia sh o w
i ng t h e s tru c tu re a n d m e th o d o f dis ch a rgi n g the m a tu re sp erms .

6 m a tu re spe rms
. 7 a fem a l e ga m e to phy te wi th s e v e ra l f em a l e
. .

se x o rga n s (a rch ego ni a ) o n the u nd e r sid e a nd wi th a clus ter of

ro o tl e ts 8 a yo u n
. ger a nd old er a rch e go nium wi th a negg re a dy for
.

fertiliz a ti oni n the b a s e o f the o ld e r one 9 a fem a l e s e xu a l pla nt . .

(a ) wi th a y o u ng sp o ro phy te s till a tta ch ed to i t .

of b o th sexes The re a re a few fe rns h oweve r i nwh ich a

.
, ,

s ma ll sp ore is p rod uced i n one ki nd of sp ora ngium a nd

gives rise to a ma le ga me top hyte so sm a ll th a t it ca na l l be
 H E TE R OS P OR Y 333

ret a i ned with in th e origi na l sp ore w a ll In these fe rns .

th e oth e r ki nd of sp ora ngium p roduces a few — s ome

times only f ou r — l a rge sp ores Th ey a re eight to ten
.

FI G 87
. . S el a gi nel l a A , a b ra nch sh owi ng th e l ea v es a rra nge d
.

i n f o ur rows a nd th e b ra nch es tipp e d wi t h co nes or s trobili . B, a

sp ora n gi um b ea ri ng l ea f f ro m th e con
~
e wi t h a mi crosp ora ngium
fill ed wi th th e sm a ll so rt of sp ore or mi crosp ores C, a sp orophyl l
.

or sp ora ngi um b ea ring l ea f wi th a m ega sp ora ngium conta i ni ng f o ur

-

l a rge or m ega sp ores D a nd E S h o wi ng co mpa ra tiv e siz e O f mi cro

.
, ,

S p ores a n d m e ga sp o res T h e l a tter a re fro m 8 to 1 0 ti m e s t h e

.

di a meter of th e f orm er a nd cons equ ently f rom 5 00 to 1 000 tim es a s

l a rge i n contents .

C oul ter Pla n

, t S tructu res Appl eton
. .
3 34 R E PR O DU CTI O N I N PLAN T S
t imes th e di a mete r O f the sm a ll Sp ore a nd c onseque ntly
c ont a in f rom five hu nd red to a th o usa nd times 8 3

or 1 0 ) a s much subs ta n
3
ce Wh e n the y ge rmi na te they .

devel op i nto sm a l l fe ma le ga me top hytes wh ich a re

‘

a lm o st wh olly e n cl o sed wi th i n th e sp ore w a ll (Fig .

This diffe re nti a ti on of sp ores i n respect to size a nd the

rete n ti on O f th e g a met ophytes withi nthe sp ore w a ll led

FI G 88 . . m e ga spore conta ining a m a tu re ful l

Sel a ginell a . 1 . a

grownf em a l e ga m eto phy te wi th se vera l a rch ego ni a expo sed th rough

’

t h e rup ture d s po re co a t 2 a yo un
. g sp o ro phyte whi ch h a s d ev el o pe d
.

f ro m a ne gg f ertiliz ed i nsu ch a na rch e go nium a s sh owni nth e p re

ce di n g figu re N o te th a t this y o u ng pla nt rem a i ns a tta ch ed for a
.

co n sid e ra bl e tim e to th e fem a l e pl a nt a nd dr a ws i ts foo d fro m th e

supply o rigina lly stored i n th e l a rge spo re (8p ) the old spo re .
,

(r ) t h e fi rs t ro o t O f th e new sp o ro phy te (s t) a nd (l ) th e y o u ng s te m
,

an d l ea v e s 3 a nd 4 old e r s ta ge s O f t h e y o u n
. . g pl a nt (Re prod u ce d .

by pe rmi ss i o n fro m dra wi ngs by P ro fess o r C a mpbe ll ) .

to th e fu rthe r devel opme nt a l ong the s a me li nes of th e , ,

p olle n a nd seeds o f h ighe r pl a nts .

R E P RO D U CT I O N B Y S EED is a spe ci a lized devel opme n t of

the a l te rna ti on o f ge ne ra ti on I f o ne ex a mi nes a fl owe r .

( Fig 8
. 9 ) it will be f o u nd to be c o m p o se d o f a n o ute r
 R E PR O DU C TI O N BY S EE D 335

c ove ring of sep a ls a nd pet a ls which ma y be b rightly

c ol ored c onstituti ng a p rotecti on to th e e n
,

,
cl osed org a ns
du ring devel opme nt a nd a me a ns of a tt ra cti ng i nsec t
V isits a fte r the fl owe r ope ns a nd i nside st a me ns a nd
, , ,

pistils E x a mi na ti on o f the pistil sh ows th a t withi n

.

it a re th e rudime nt s of seeds c a lled ovules At th e top

, .

of the st a me n s a re e nl a rgeme nts c a lled a nthe rs I nside .

them a re sm a ll b odies know na s p olleng ra i ns Ina ppe a r .

an ce they resemble th e sp ores of the fe rna nd a re i nf a ct

a s we sh a ll p rese n tly see onl y a m odific a ti on of them
, .

FI G 89. . Pa rts of a l ga ns ) A a no penfl ow er

Fl ow er (C l a rki a e e .
,

sh owing (a ) a nth ers (p ) p eta ls (st) s tigm a ; B o pe n fl ow er a nd

, , ,

unopened bud p a rts l ettere d a s a b o ve i no penfl ow er B u d co vered

,
.

by s ep a ls (s ) a nd b elow th e O p enfl ow er th e o v a ry whi ch con

ta ins th e o v u l e s .

I t is well k now n to a lmost eve ry one th a t i nseed rep ro

'

ducti onit is necess a ry for th e p olle nto be pl a ce d onth e

tip of the pistil c a lled th e stigm a i norde r th a t fe rt i liz a
, ,

ti on ma y eve ntu a lly result a nd seed be p roduced Th e .

det a ils wh i ch u nde rlie this p rocess a re p ra ctic a lly u n

know n to the l a ym a n a nd nee d to be c a ref ully studi ed
3 36 R E P R O DU C TI O N IN P L AN T S

i norde r to u nde rst a nd rep roducti on i n fl owe ring pl a nts .

T H E O VU LE AN D F E M A LE G A M ETO P H Y TE (Fig .

Th e ovule a c tu a lly c orresp onds to the org a n(sp ora ngium )

on the b a ck O f a fe rn le a f i n which the sp ores a re p ro

du ced . Wh e nthe devel opme nt of th e ovule is c a refully

studied i t is fou nd th a t a t a ce rt a i nst a ge the re is devel
oped a sp ore m o th e r cell a n d th a t it u nde rg oes two succes
sive divisi ons ex a c tly c omp a ra ble to th ose desc ribed i n
C h a p te r XX V I for th e m a tu ra ti onof the a n im a l egg a nd ,

A
FI G . 90 .
— T h e Ovul e a nd Fem a l e Ga m e to phy te A , sh Ows a cross
.

i n th ro u gh th e b a s e o f th e pis til o f a 1i fl ow er Th ere a re

!

Se c t o .

t h ree ca vi ti es i n e a c h o f whi c h
,
a re two v ert i ca l rows o f o v u l es .

T h e figu re sh o ws six o vul es ea ch ,

co nta i ning a nea rly m a tu re fema l e
ga m e to phy te .

which will be p rese ntly desc ribed for pl a nt p olle n These .

t wo divisi on s result i nfo u r sp ores only one of which sur,

vives a nd m a tu res Th e othe r th ree rem a i nsm a ll a nd a re

.

eve ntu a lly c rush ed a nd dest royed Th e one fu nc ti ona l .

sp ore devel ops i nt o a sm a ll fema le ga me top hy te with o ut

eve r bei ng sh ed f rom th e o vule I t is the fa ct th a t the
.

S p o re is no t sh ed o n the g ro u n d a s it is i nthe fe rns a nd ,

th a t the g a met ophyte i s sma l l e nough to be re ta ined i n

'
 AN TH E R S AND M AL E GA ME T O P H Y T E 3 37

th e ovule th a t m a kes p ossible th e seed Wh e n th e en .

-

cl o sed a nd p a ra sitic fem a le g a met ophyte is full g row ni t

‘
-

c onsists of only seve n cells At one end is the egg cell

.

an d two a ss oci a ted cells ; a t the opp osi te en

'

d a re th ree
veget a tive cells wh ich ordi na rily a re of negligible v a lue ;
i nth e middle is a si ngle l a rge nucleus which wa s f ormed
by th e fusi on of two one of wh ich c a me f rom th e egg
,

g roup a t one end a nd the o the r of which c a me f rom the

B
B, a S i ngl e o vul e sh owing th e o u ter coveri ngs or inte gu m ents
and th e fem a l e ga m eto phy te i n th e middl e T h e l a tter sh ows i n
.

t his si ngl e s ecti ononly 4 of th e s ev encells a ctu a lly present

.

O pp osite g ro up of veget a tive cells Th is si ngle l a rge

.

d ouble nucleus is c a lled th e p rim a ry endosp erm nu cleu s .

It pl a ys a ve ry imp ort a nt p a r t i n the devel opme nt o f

ce rt a inseeds of ec onomic v a lue a nd must be fully u nde r
st oo d he re i f wh a t is to be s a id l a te r i nrespe ct to ce rt a i n
ph a ses o f he redity in c orn is to be u nde rst ood .

T H E A N T H ER S AN D M A LE G A M ETO P H Y T E (Fig .

Tu rni ng ou r a ttenti onnow to th e a nthe rs it will be f ou nd

3 38 R E P R O DU C TI O N I N PL AN T S
mp a ra bl e devel opme nt g o es on th e re A con
‘

th a t a co .

si dera bl e n umbe r of sp ore m othe r cells devel op i nth e c a vi

ties of e a ch a nthe r E a ch si ngle one u nde rg oes th e p roc
.

esses of reduc ti ona l d i v i s i ona nd p roduces f ou r sp ores j ust ,

a s e a ch spe rm m o the r cell i n th e a n im a l p roduces fou r

spe rms Th e sp ores which devel op i nthe a nthe r a ls o b e
.

g in t o g r o w a nd u su a lly c ont a i nt wo o r th ree cells a t the

time th ey a re shed a nd c a rried by wi nd or insect to th e tip
o f th e pistil He re th ey p rod uce el ong a ted p ro t rusi ons of
.

thei r w a lls c a lled p ollentu bes wh ich b ore th ei r wa y d ow n ,

th rough th e tissue to th e c a vity of th e pis til wh e re the “

FI G 91 . .
—T h e Poll en This figu
. sh ows a cro ss secti o nthrou gh
re

th e a nth er a t th e top O f a s ta m en T h ere a re fo ur ca vi ti es i nwhi c h

.

t h o us a nds of pollengra i ns a re f orm ed Ea ch po ll engra i nis a mi cro

.

spore wh en fi rs t f orm ed b u t u su a lly h a s germi na te d to f o rm a two

cell e d m a l e ga m eto phy te b e f o re i t is a ctu a lly sh ed a nd ca rri ed to
th e ti p o f th e pis til (S ee Fi g 92)
. .

o vules a re a nd e nte r a mi nut e ope ni ng i n the ovule

, ,

k now n a s th e mic ropyle a nd eve ntu a lly r e a ch th e egg

,

i nside Inth e me a ntime th e p olle ng ra i nh a s c ompleted

.

its gro wth i nt o a mi croscop i c ma le ga me top hy te I t is so .

redu ced inf a ct th a t it no rm a lly c onsists of only th ree cells .

One o f these is th e so c a lled tube nucleus wh ich a ppe a rs

-

to di rect th e a c ti ono f th e p oll e ntube a n d th e o th e r two

a re spe rm s .

F ERTI LI Z ATI O N EM BRY O AN D EN DO S P ER M (Fig

, ,
.

Wh e nth e two spe rm s a re disch a rged from th e p olle ntub e

 FE R TILI Z ATI O N , E M B R YO, E ND O S P ER M 339

int o th e fem a le g a met ophyte i nside the ovule of a Fl owe r

in g Pl a nt on e fuses
, with the egg t o fe rtilize it a nd th e ,

o the r u n ites with th e p rim a ry e nd osp e rm nucleus F rom .

the fe rt ilized egg devel ops the y o ung emb ry o sp orophyte

pl a nt . The e nd osp e rm nucleus now c onsists of th ree
fused nuclei a nd th e p rocess by which i t is f ormed is often
refe rred to a s t riple fusi on ” I t is s ometimes c a lled
.

doub l e ferti li z a ti on ona cc ou nt of th e f a c t th a t on

o
e of

the cells t a ki ng pa rt is a spe rm I t g rows a nd divides to .

FI G 92 . .
-
M a 1 e gametophyte or p ollen gra in a t ti me i t is sh ed
f rom a nt h er .
(t) tub e nu cl eus ; (8) sp erm m o th er cell wh i ch will divide
t o f orm th e two sperms .

p roduce a speci a l a nd p eculi a r tissu e c a lled endosp erm .

In s ome seeds th e re is ve ry little devel opme nt of

e nd ospe rm wh ile i no the rs like c orn the re is a l a rge de
, ,

v el op men t so th a t th e l a rge r p a rt of the m a tu re seed con

sists of end ospe rm .

T H E S EED is th e m a tu red ovule c ont a i ni n g a y ou ng

pl a nt a nd a st ore o f f ood (Fig 94 ) The emb ry o pl a nt h a s
.

beende rived by cell divisi on g rowth a nd d iff e re nti a ti on

, ,
3 40 R E P R O DU C TI O N I N PLAN T S
from th e fe rtili zed egg i nthe s a me m a nne r th a t th e a ni
ma l a rises f rom a fe rtilized egg S i nce the devel opme nt o f
.

th e a nim a l emb ry o h a s bee na dequ a tely desc ribed i nth e

l a st ch a pter it is u nn
ecess a ry to repe a t the st ory he re In .

th e devel opme nt of e nd ospe rm the re is cell divisi ona nd

FI G 93
. . A is a y ou ng o vul e wi th th e m ega spo re m o th er cell .

B sh ows th e m a tur e f e m a l e ga m e to phy te whi ch d ev el o ps f rom t he

m ega sp ore ; a t o ne end a re the egg 6 a nd two o th er cel ls ca ll e d
, ,

sy nergi ds s ; i nth e middl e a re two nu cl ei i nth e a ct o f fusi ng toge th er

,

to fo rm the e n d ospe rm nu cl eus ; a nd a t th e o th er end a re th ree cel ls

whi ch ordi na rily ta k e no fu rth er p a rt i nth e d ev el opment o f the s ee d .

Wh enth e p oll e ntub e rea ch es a fem a l e ga m e to phy te o ne o f i ts spe rms

u ni tes wi th the e gg a nd giv es ris e to t he y o u ng pl a nt o r emb ry o ,

t h e o th e r u n i tes wi th the e nd osperm nu cl eus a nd ma y giv e ris e to

a spe ci a l tissu e ca ll ed end o spe rm whi ch wh en p res ent conta ins
,

t h e f oo d supply fo r the emb ry o .

g rowth but little d i ffe re nti a ti on o f cells bey ond th a t in

v olved i n f ood s tora ge Wh e reve r a bu nd a nt e nd ospe rm
.

is devel oped its cells a re u sed to st ore a supply of st a rch

o r o the r f ood for th e y ou ng pl a nt Inpi nes a nd simil a r
.

pl a nts th e re is no e nd ospe rm a nd f ood is st ored i n the

l a rge fem a le g a met ophyte I nFl owe ri ng Pl a nts whe reve r
.
 D O R M AN C Y OF S EED S

ndosp erm i s impe rfectly devel oped e g the p ea f ood

'
‘

e ,
. .
, ,

st ora ge occu rs i n th e seed le a ves or c o tyled ons of th e

‘

emb ry o pl a nt Th e tissues o f th e ovule c onti ue g row

. th n
a l ong with the y oun g pla nt and bec ome di ffe renti a te d t o
f orm a h a rd c oat which se rve s to p ro tect the emb ry o
: ;

Pe a s o od . w1
t h s e e d co a t r e
mov e d a nd ‘o o t yl e d ons s a p s ;
r a t e d t o s h ow youn d s t em
an d r o ot .

Pi ne s ee d Co r n gr a i n

FI G 94 — T h e S tru cture of ' S eeds ;

. . T h e di a gra s sh ow t hree m
t yp es of s ee d (1 ) P ine s ee d i nwhi ch th e emb ryo i s l o ca te d i nthe
.
r

middl e of th e l a rge f em a l e g a m eto phy te a nd h a s m ost of th e foo d

” s tored in th e ga m e to phy te a nd s om e i n th e emb ry o ; (2) th e corn
gra i n whi ch h a s f oo d s tored l a r gely i n th e cells of th e endosp erm ;
an d (3 ) th e p e a s ee d , i n whi ch t h ere is no thi ng l ef t ei th er o f th e ,

fem a l e ga metophy te or of th e end osp erm , h a s th e f oo d s tored i n ‘

th e m u ch enla rge d s ee d l ea v es or co tyl edons N ote a ls o th at th e .

emb r y o of th e pi n e h a s s e vera l co tyl edon s of whi ch th e di a gra m

sh ows two , one onei th er side of th e s tem tip ; th a t th e cornemb ry o
‘

h a s b ut one v ery l a rge co tyl e d onor s ee d l ea f a nd t h a t th e s tei nti p

o ccupi es a n a pp a ren t ly l a tera l p osi ti on; a n d fin a lly t h a t th e pe a

ha s two l a rge l a tera l cotyl ed ons a nd a term ina l s te m ti p .

D OR M A N CY OF SEEDs
Th e seed .
-
f m most di ff e rs ro -

p a rts of th e pl a nt i n th a t it ca n bec ome a i r dry with ou t

i nj ury to th e p ro topl a sm of i ts cells a nd the n resu me !

”
growth whe n pl a ced in a suit a ble tempe ra tu re a nd pro .

v i ded with sufficie n t w a te r Th l S 1 s a ve ry imp ort a n

. t
ci rcumst a nce for it e na bles a pl a nt to su rvive l ong con
n
-

ti i i ed unf a v ora ble c on diti ons for growth or to b e ca rri ed '

l on n t n z d n e h s
‘

d ist ces with i j u y S ee Pl ts h t

'

o
’
!
u
’

g a r a a v . u

a g re a t a dv a nt a ge o ve r Fe rn s a nd the l ow e r
,
i
3 42 R E P R O DU C TI O N I N PLAN T S
p l a nts which h a ve less eff ective me a ns of rep rod uc ti on .

T H E CH RO M OS O M E H I S TO RY I N P LA N TS AN D A N I M A LS .

S ince th e pl a nt ch rom os omes beh a ve i na lm o st i denti

c a lly th e s a me m a nne r a s th ose of a nim a ls the re is no
necessity to rest a te th e f a cts a side from th e opp ortu nity
,

i t a ff ords to reite ra te th e esse nti a ls of the p rocess a nd

the reby fi x them m ore fir mly inmi nd I t sh ould h oweve r .
, ,

be remembe red in c onnecti onwith th is st a teme nt th a t

, ,

the reducti on divisi ons of pl a nts occu r a t a di ff e re nt

!

pe ri od in th e life h ist ory (Fig In a nim a ls redue

.

ti onoccu rs du ring th e p relimi na ry st a ges of the f orm a ti on

of g a metes i e , eggs a nd spe rms but i npl a nts it occu rs
. .
, ,

j ust p revi ou s to the f orm a ti on of sp ores The sp ores .

the ndevel op i nt o g a m et ophytes l a rge or sm a ll a cc ordi ng

,

to th e s ort of pl a n t u nde r c onside ra ti on Whe nm a t ure .

the g a met ophytes f orm th e eggs a nd spe rms but of c ou rse , ,

no reducti on i s necess a ry a t this p o int bec a use it h a s

a l re a dy t a ke npl a ce inthe f orm a ti ono f th e sp ores The .

m a tte r ma y be p u t a nothe r wa y thus : Inthe c a se of the

an im a l th e g a metes h a ve th e red uced numbe r of ch rom o
s omes Conseque ntly th e fe rtilized egg will h a ve th e
.

d o uble numbe r Si nce th e a nim a l b ody devel ops f rom th e

.

fe rtilized egg by ord ina ry cell divisi on eve ry cell of th e

an im a l b od y will a ls o h a ve th e d o uble numbe r I n the .

pl a nt th e g a metes a ls o h a ve th e red uced numbe r a nd th e

fe rtili z ed egg th e d ouble number The fe rti lized egg .

grows int o th e sp orophyte pl a nt by cell divisi on a nd

eve ry cell h a s th e d o uble numbe r R ed ucti on occu rs .

with the f orm a ti ono f sp ores a nd the refore th e g a met o

phyte h a s th e red uced numbe r F rom th is it will be see n
.

th a t th e re i s a n ext ra ge ne ra ti on the g a met ophyte

i nthe pl a nt life hist ory which h a s n o thi ng ex a c tly c orre
sp o n ding to it i n a nim a ls .

R EDU CT I O N AN D S EGR EGAT I O N or C H RO M OS O M ES IN

P LA N T S (Fig —This p rocess is m ost f reque ntly

.

studied i n th e a nthe r bec a use the re a re m a ny sp ore

m oth e r cells giving a nopp ortu nity to se e m a ny d iffe re nt
-
 R E D U C TI O N AND S E G R E GATI O N 3 43
'

a na i m

Body ca n mo th er

el s A im
Pa n al n
b ody 0 0 1 1 of
a nt mm fi oo

k g m
ot he r co

P l a nt

Moth er Ce ll

FI G . 95 . Th e Lif e- C y cl es nd An of P la nts
i m a ls C omp a red a .

T h e di a gra ms a re l a r gely s elf e xpl a na tory a nd sh ow th a t th e a n

- ima l
an d high er pl a nt b ot h origi n a te by th e u n i onof a sperm a nd a negg ,

t h ereby d o ubling th e chr om os o m e s n u mber so th a t ea ch cell h a s n

’

p a irs of ch rom os omes Wh enth e a ni m a l f orms i ts ga metes th e mem

.

bers of ea ch p a ir sep a ra te fro m one a noth er a nd go to opposi te po l es

of th e spin dle This resul ts in redu ci ng th e nu mb er by h a lf a nd
.

f urt h er pro vides th a t ea ch ga mete h a s b u t one ch rom osom e of ea ch

p a ir Fert iliz a tion a ga in uni tes a sperm a nd a n egg a nd d oubl es
.

th e n umb er a ga in Inth e pl a nt the history is th e s a m e i n so fa r

.

a s th e cy cl e u p to re du ct i on is con cern ed T he redu ct i on di visi on s .

gi ve rise to sp ores ins tea d of ga metes a nd spores dev el o p i nto mi cro,

s copi c ga m etophy te or sexu a l pla nts in the hi gh er pl a nts whi ch in

turnp ro du ce th e ga m etes On e sperm f us es wi th th e egg t o p ro d u ce
.

th e n ew pl a nt an d th e oth er i nm a ny pl a nts uni tes wi th th e end osperm

nu cl eus to pro du ce th e end osperm a p a rt of th e seed peculi a r to ,

certa i n fl ow erin g pla nts s u ch a s corn .

344 R E P R O DU C TI O N IN P LANT S

st a ges ine a ch mic rosc opic secti on I ne a ch sp ore m othe r

.

cell th e ch rom a ti n net res olves itself int o ch rom os ome

th re a ds Th ese the nc onj ug a te side by side Th e p a i red
. .

ch rom os omes nex t sh orte n a nd th icke n By th e time .

they a re m a tu re a S pi ndle h a s bee n f ormed which in ,

pl a nts h a s blu nt p oles a nd l a cks centrosomes such a s a re

a lw a ys p rese n t i n a nim a ls E a ch ch rom os ome p a i r h a s
.

FI G 96
. . C ell D ivisi o na nd R e du c tio n C o mp a re d .C e ll divisi o n
o cc u rs re gul a rly i n grow th , i ts ess e nti a l f ea tu re is the l o ngi tudi nal

spli tting of ea ch i ndividu a l chr o m o s o m e so th a t o ne- h a lf o f e a ch

goes to ea ch po l e o f th e spi ndl e a nd thus p ro vid es th a t th e two
new cells sh a ll be e x a ctly lik e e a ch o th er a nd e xa c tly li k e th e o ne
f rom whi ch th ey h a ve aris en .

In re du cti o n divisi o n the m emb ers of ea ch ch rom os om e p a ir ,

fi rs t j o insid e by sid e a nd th e na rra nge th ems elves o nthe spi ndl e i n

su ch a wa y th a t one goe s to ea ch po le th u s se grega ti ng th em f rom
,

on e an o th er Si n
. ce t h e p a i rs a re a rra nge d o nthe spi ndl e i na ch a nce
m a nner i t foll ows th a t this no t only red u ces the numbe r of ch ro m o
so mes th a t go to th e s po res b u t i t a ls o provid es the m ea ns by whi ch
new a sso rtm ents a rise so th a t th e S p o res p ro du ce d by a pl a nt will
co n ta i na ll th e po ssibl e a ss ortm e n ts o f ch ro m o s o m es
.
 V E G E TA T I VE R E P R O D U CTI O N 34 5

o ne ch rom os ome de rived origi na lly f rom e a ch p a re nt .

The one from th e origi na l p olle np a re nt is c a l led p a te rna l ,

an d th e on e f rom the origin a l seed p a re nt m a te rna l The .

membe rs o f e a ch p a i r a re ve ry S imil a r th o ugh th e diff e re nt

p a i rs di ffe r much from one a no the r Whe n th e p a i red .

ch rom o s omes a rra nge th emselves a t th e middle O f the

'

S pi n dle it is a m a tte r of ch a nce whethe r the m a te rna l or

th e p a te rna l one is tu rned t ow a rd a ny p a rticul a r p ole .

Next th e membe rs a re d ra w n a p a rt a l ong th e spi ndle

fib res This sep a ra ti on o f th e respective m a te rna l ch ro
.

mosomes f rom their p a te rna l fell ows is k now na s segrega

ti on I t is a ls o sp o ke n O f a s redu cti on B o th fe a tu res
. .

a re imp or t a n t R educti on i nnumbe r to one h a lf keeps

.
-

the ch rom o s ome numbe r c onst a nt gene ra ti on a fte r gen

,

era ti on a n d seg reg a ti ong ives th e Sp ores a new a ss ort

,

me nt a m a tte r O f g re a t imp ort a nce i nhe redity

,
.

V EGETAT I VE R EP R OD U CT I O N is ve ry c omm on i n pl a nts .

The re a re m a ny w a ys O f a cc omplishi ng it a cc ordi ng to ,

the st ructu re of p a rticul a r pl a nts A p ot a t o f or ex a mple ;

.
,

p roduces u nde rg ro und stems which bec ome e nl a rge d a nd

filled wi th s tored f ood chiefly st a rch Wh en th e pl a nt
, .

m a tu res a nd dies th ey a re eithe r h a rvested by ma nfor h is

own use o r rem a i n i n th e gro u n d i n the c a se of wild
,
’

pl a nts In eithe r eve nt they will begin to g row on th e

.

resump ti on o f f a v ora ble we a th e r a n d s o il c onditi ons .

N ew sh oo ts will be p ut ou t f ro m the le a f a xils p opul a rly ,

c a lled eyes a nd roots from the b a s a l p orti ons O f th e

,

sh oo ts thus givi ng rise to new pl a nts B ulbs put ou t

, .

side b ra nches which devel op new bulbs th a t a re used to

st a rt new pl a nts R ootst ocks O f I ris a re me rely p rost ra te
.

stems Th ey ma y be divided a nd used to st a rt new

.

pl a nts bec a use e a ch piece will p roduce new roots a nd

sh oots D a hli a roo ts ma y be used i nthe s a me wa y A
. .

twig of a will ow ma y be cut a nd stuck i nt o m oist s oil

whe re it will pu t ou t roo ts a nd g row i nt o a t ree A twig .

o f a pe a ch t ree ma y be cut a n d g ra fted i nt o a no the r t ree

whe re it will not only c onti nue g rowth but a ls o unite its
3 46 R E P R O D U CTI O N I N PLAN T S
tissues with th ose of the pl a nt to wh ich it h a s bee n
g ra fted A bud cut off a nd i nse rted ina slit th ro ugh th e
.

b a rk will g row in a si mil a r m a nne r .

Ina l l th ese a n d oth e r veget a tive meth ods O f rep roduc

ti on the re is one c omm on f a c t A piece o f th e pl a nt
.

c onsisti ng O f m ore or fewe r ordi na ry ce l ls is det a ch ed to

c onti nue its g row th elsewhe re Th ese cell s a rose th ro ugh
.

o rdi n a ry m ito tic cell d ivisi on a n d so i n respect to th eir

nuclei a re ex a ctly like a ll the othe r cells of the pl a nt N O
.

reducti onor seg reg a ti onof a n y kin d h a s occu rred Wh a t

.

eve r a ss ortme nt of ch rom os omes th e origi na l pl a nt h a s

i n i ts nuclei th a t ide ntic a l as s ortme nt will be f ou n
,
d in
the det a ch ed p orti on Th e new pl a nt is to a ll inte nts
.

an d pu rp oses me rely a p a rt of th e Old All th e na vel

n
.

ora n ge t rees O f Ca li f or i a now ru nni ng i nt o h u nd reds of

,

t h o us a nds h a ve bee n de rived i n this m a nne r f rom two

,

origi n a l t rees b rough t f ro m B ra zil .Th a t th ey a re not

a ll ex a ctly a like a n d equ a lly f ruitful i s due to a no the r
‘

c a use which will be expl a i ned i n a l a te r ch a p te r on

v a ri a ti on
.
SECTI ON 9

HERED I T Y
Gregor M end el , th e A u stri a nmonk wh o fi rst
,
correct ly
s ta te d the first two la ws of here dity
.
 C HAPTER XX VIII

M E ND E L S LAW S
’

VAR I ATI ON — Child re n neve r

. esemble eith e r p a re nt
r

ex a ctly i na l l p a rticul a rs a nd it is ve ry d oubtful wh ethe r

an y si n gle ch a ra cte r is a nex a ct duplic a te of th a t O f eithe r
p a rent S uch differences a mong i ndi vi du a ls rela ted by
.

descent a re c a lled va ri a ti on s by the bi ol ogist The .

stude nt sh ould note th a t th e di ffe re nces betwee na n oa k

an d a ma n a re not i ncluded i n th is defi niti on bec a use
these two org a nism s a re not ne a rly rel a ted by desce nt .

P UR E L I N E S AN D HY BRI D S — In p recedi ng ch a pte rs

.

th e stude nt h a s le a rned th a t the re a re two s or ts of r ep ro

ducti onf ou nd a m ong b oth pl a nts a nd a nim a ls ; viz sexu a l .
,

an d a sexu a l. In a l l a sexu a l meth ods th e re is only one

p a ren t a nd th e successive ge
, ne ra ti o n s a re a l l a like
(Ch a p XX V I I ) M ore ove r wh e n th e two p a re nts a re
. .
,

germi na lly a like or wh e nth e spe rm a nd egg a re regul a rly

p roduced by th e s a me i ndividu a l it is cle a r th a t sexu a l
,

meth ods will p roduce pu re li nes But when the two .

p a re nts bel ong to di ff e re nt species ra ces or v a rieties th e

, ,

O ffsp ri n g will receive a di ffe re nt c ont ributi on f rom e a ch

an d c onseque ntly will not be pu re b red a nd will no t b e
l ong to a pu re li ne The b reedi ng t ogeth e r of such dif
.

ferent p a re nts (i ndic a ted i n th is b ook by X ) is c a lled a

cross a n d th e Offs p ri ng is s a id to be h g b ri d
z .

M ODI F I CATI ON S — Ih a ny pu re li ne the re is a lw a ys

s ome v a ri a ti on a m ong th e v a ri o us i ndividu a ls For ex .

a mple two be a n
,
s f rom the s a me p od bel ong to the s a me
pu re li ne bec a use be a ns a re ordi na rily self p olli na ted -
.

Neve rth eless they w o uld sc a rcely eve r weigh p recisely

th e s a me n or wh e npl a n ted p ro duce pl a nts ex a ctly a like .

34 9
3 50 M E ND E L ’
S L AW S
Inm a ny c a s es the v a ri a ti onmigh t be k now nto be due to
s oi l or clim a te or s ome othe r e nvironme nt a l f a ct or In .

an y eve nt th e v a ri a ti o ns w o uld no t be t ra nsmitted to

their O ff sp ri ng This t ype O f v a ri a ti onis c a lled modifica
.

tio n .

M UTATI O N S E ve ni nth e pu rest ra ces or i npu re li n

. es
v a ri a ti ons s ometimes occu r wh ich a re t ra nsmitted to sue
ceedi n g ge n e ra ti ons Th ey a re k now n a s mu ta ti on
. s A .

no t u nc omm onillust ra ti ono f mu t a ti onis th e a ppe a ra nce

o f a white flowered pl a n -
t in a p u re l ine wi th c ol ored
fl owe rs I f self p ollina ted such wh ite pl a nts a lm ost i h
.
-

v a ri a bly b reed t rue ge ne ra ti on a fte r ge ne ra ti oni rresp ec

tive o f th e clim a te or c onditi ons of cultiv a ti on .

R CO B N
E M I A T I ON s — Wh e n p a re nts diff e r i n seve ra l
.

ch a ra cte rs th e hyb rid O ff sp ri ng usu a lly exhibit s ome of the

ch a ra cte rs O f e a ch p a re nt a nd a ls o l a ck s ome f rom e a ch
on e . V a ri a ti ons Of th is s ort due to rec ombina ti onof ch a r
a cters a re n eithe r m odific a ti ons nor mut a ti ons .

H ERED I TY is th e p rocess which is resp onsible for th e

p a rticul a r c ombi na ti on o f t ra nsmissible ch a ra c te rs p os
sessed by a ny org a nism It sh o uld be noted th a t th e .

defi niti on m a kes h e redity resp onsible for th e d iff e re nces

betwee n p a re nts a nd O ffsp ri ng a s well a s for thei r re
sembl a nces P rob a bly a nill ust ra ti onwill m a ke this cle a r
. .

Wh e na pu re b row neyed ma nm a rries a pu re blue eyed

- -

w om a n th e child re n a ll h a ve b ro w n eyes but if these

b row neyed off sp ri ng m a rry blue eyed m a tes a pp roxi
- -

m a tely h alf thei r ch ild re n will h a ve blue eyes a nd h a lf

b row neyes Th e p ro cess o f h e redity is equ a lly resp onsi
.

ble for b o th results .

M EN DE L S D I S CO VER Y O F T H E LAws O F H ER EDI TY

’
.

Al th ough th e f a c t of he redity h a s bee n k now n for two

th o u sa n d ye a rs o r m ore the l a ws h a ve only rece ntly bee n
disc o ve red The fi rst two O f them we owe to a nAust ri a n
.

m onk G reg or M e ndel wh o expe rime nted with g a rde n

, ,

pe a s a nd p ublished a n a cc ou nt o f his w ork i n 1 865 i n ,

which he m a d e the fi rst st a teme nt o f wh a t we know tod a y

 A S I M P L E ILL U S T R ATI O N 351

as : (1 )the La w of S eg reg a ti on a nd (2) the La w of R e a s ,

s ortment H is m ome nt ou s disc ove ry wa s a lm ost wh olly

.

neglected by his ownge ne ra ti ona nd only b rough t to th e

a tte nti onOf th e w orld i nge ne ra l i n 1 900 wh e nh is p a pe r
Wa s u ne a rthed by th ree E u rope a n b ot a nists (de V ries ,

C orre ns a nd T sch erma k ) wh o we re a ls o a ttemp ti ng to

,

disc ove r the l a ws O f h e redity .

THE LAW OF SEGR EGAT ION

A S I M P LE I LL U S TR AT I O N .
— Alth o ugh M e ndel w orked
with pe a s this l a w ca n be m ore re a dily u nde rst ood by
studying a diff e re nt expe rime nt (Fig Wh ena red .

FI G 97
. .
-
D i a gra m to
h ow th e resul t s of crossi ng a red four
S

O cl ock by a whi te on Th e upp er row sh ows th e re d a nd whi te

’
e .

pa rents Th e second row sh ows t h a t th e fi rs t genera ti on hyb rids

.

a re a ll a lik e a n d pi nk i nco l or T h e t hi r d ro w sh ows t h a t wh ent h es e

.

pink hyb rids a re s elf p ollina ted or b red wi th one a noth er a s econd
-

hyb rid genera t ion is pro du ced whi ch consis ts of one fou rth whi te -

fl ow ere d pl a n ts li k e o ne gra n dp a rent one h a l f pink hyb rids lik e th e

,
-

first hyb rid genera t i on a nd one f o urth red pl a nts lik e th e s econd
,
-

gra n dp a rent .

M orga n S tu rtev a nt M ull er B ridges The M echa ni sm of M endeli a n

, , , ,

Heredi ty .

fl we ed f ou
o r cl ock is c ssed with wh ite flowered one
r o
’
ro a -

the hyb rid off sp ri ng c a lled th e F ge ne ra ti on a re a ll pi nk

, 1 ,

flowered If th ese hyb rid pi nks a re self p olli na ted thei r

.
-

Offsp ri ng c a lled th e F ge ne ra ti on c onsists of th ree

, 2 ,
3 52 M E ND E L ’
S L AW S
cl a sses : one f o u rth a re red like one o f th e gra ndp a re nts ;
-
,

on e h a lf pi n
-
k li ke th e hyb rid F ge ne ra ti on; a nd one
, ,

f ou rth white like th e o th e r g ra ndp a re nt Th e reds a nd

,
.

whites b reed t rue i nsuccessive self p ollina ted ge ne ra ti ons -

b u t the pi nks a re a lw a ys hyb rids a nd a lw a ys p rod uce 1

red ; 2 p in k ; 1 wh i te wh e n self p olli na te d -
.

T H E P U R TY O F G A M ET S
I E — At th e time M e n . del
w orked with his pe a s no one k new th e st ructu re O f cells
s o h e wa s c ompelled to f orm ul a te a p u rely hyp o thetic a l

expl a na ti on i n orde r to u nde rst a nd why the g ra nd p a r -

e nt a l types a ppe a r i na defi nite ra ti o i n th e F a nd l a te r 2

ge ne ra ti ons a l ong with the hyb rids His expl a na ti ona p .

plied to th e f ou r O cl ocks w ould be a s f oll ows : (1 ) Th e red

’

p a re nt a nd th e white one e a ch t ra nsmi ts to th e pi nk h y

b rid th rough th ei r respective ge rm cells s ome s ort o f a de
te rmi ne r for fl owe r c ol or wh ose i ntera cti qn p roduces a
,

pi nk c ol or ; (2 ) Whe nth e pi nk hyb rid f orms its ge rm cells

o r g a metes th e two dete rmi n e rs a re sep a ra ted a nd a ny
p a rticul a r g a mete ma y receive eithe r the red dete r ” “

mi ne r or the wh ite o ne but not b oth (3 ) He nce some

“
.

g a metes a re pu re for red a nd a n equ a l numbe r a re pu re

fo r white ( )
4 Th e ch n ce c ombi n a ti o ns o f equ l numbe rs
”
a a
”
.

red an d white g a me tes will p roduce one f o u rth

“
of -

red x red (pu re red ) on

,
e h a lf red x white (hyb rid pi n
,
-
k) , ,

an d one f o u rth wh ite x wh ite (p u re wh ite ) c ombi na ti ons

-
.

T H E R E CO M B I N AT I O N R AT I O ca n be re a d ily ,

u nde rst ood by ex a mi ni ng th e a ccomp a ni ng di a gra m .

S P ER M

Wh ite Wh ite
FI G 98
. is a di a gra m sh owi ng p o ssibl e combi na ti ons o f red a nd
whi te sperms wi th red a nd whi te e g gs to p rodu ce the 1 : 2 1 ra t io .

The p ossible c ombi na ti ons a re f o u r C ombi na ti on 1 .

o ccu rs o ne time i nf o u r a nd p rod uces pu re red Combi na .

 THE BEAR ER S OF H E R E DI T Y ’

3 53

ti ons 2 a nd 3 b oth p roduce red x wh ite hyb rids or pi nk ,

an d c onstitute h a lf th e O ff sp ri ng Co mbi na ti on4 p roduces

.

pu re wh ite a nd occu rs one f ou rth O f the time This ma y

-
.

be f ormul a ted i n ge ne ra l te rms by s a yi ng th a t th e F 2

rec ombi n a ti ons will result i n 1 fi rst g ra nd p a rent a l type -

to 2 hyb rid F t yp es to 1 sec on

1 d g ra nd p a re nt a l type It -
.

is c omm only refe rred to a s the ra ti o .

T H E B E ARER S OF H ER ED T Y ARE C H R O OS O
I M M E S — In
th e p receding p a ra gra ph i t h a s been a ssumed th a t the
ge rm cells or g a metes receive s ome s ort O f dete rm i ne rs now
know na s fa c tors or genes which they p a ss onto th e O ff
sp ri ng I t is now desi ra ble to i nqui re wh ethe r the re is
.

an y mech a n ism suit a ble to pe rf o rm th is fu ncti on The .

r e a de r sh o uld n ow rec a ll th a t it h a s bee nS h ow ni nch a p

te rs XX V—XX V I I th a t (1 ) Th e g a metes a re th e s ole con

tri b u ti on of th e p a re nts to th e O ff sp ri ng (2) Th a t the ,

egg is O ftena th ous a nd times l a rge r th a nth e spe rm but ,

th a t thei r nuclei a re a pp roxim a tely equ a l (3 ) Th a t th e ,

ch rom o s omes a re th e on ly p a rts of th e nucleus wh ich

m a i nt a ina c onti nu o us ide ntity Th e l a rge size of eggs is
.

due to th ei r h a vi ng much m ore cyt opl a sm th a nth e spe rm

an d usu a lly to this bei n
, ,
g st ored with a bu nd a nt f ood for
the y ou ng org a nism The S pe rm l a cks this a lm ost wh olly
.
,

but is yet equ a lly p ote nt i nhe redity I t seems re a s ona ble.

to thi n k the ref ore th a t a t le a st the chief p a rt of th e

, ,

he redit a ry u nits c a lled genes a re c ont a i ned i nth e nucleus .

If th a t is so they must a ls o be c ont a i ned i nth e pe rm a ne nt

p a rt O f it the ch rom os omes Inf a ct th is i s k now nto be
,
.

t rue but its dem onst ra ti oninv olves a k nowledge of f a cts

,

with which th e stude nt is not yet f a mili a r .

T H E C H R O M OS O M E S CO N TA I N T H E G E N E S — Th e stu .

de nt is now a sked to rec a ll wh a t h e h a s a l re a dy le a rned

i n Ch a pte r XX V a b o ut the beh a vi or O f ch rom os omes .

The re it wa s sh ow nth a t i nordi na ry cell divisi one a ch a nd

eve ry ch rom os ome Splits le ngthwise i nt o two ex a ctly simi
l a r h a lves one O f which goes to e a ch O f th e new nuclei
'
!

.
,

This inv olves a S i mil a r splitting O f eve ry ge ne in th e

354 M E ND E L S LAW S
’

ch rom os ome a nd i nsu res th a t th e two new cell s h a ve

ex a ctly th e s a me numbe r a nd ki nds o f ch rom os omes I t .

a ls o f oll ows th a t eve ry cell i nth e b ody o f a pl a n t or a ni

ma l h a s ex a ctly th e s a me equipme nt O f ge nes h e redi
ta ry u nits ) . But si nce th ese ch rom os omes a re p rese nt i n
p a i rs o ne membe r o f e a ch p a i r h a vi ng bee nde rived f rom
,

e a ch o f th e org a nism s p a re nts th e re a re a ctu a lly two

’
,

ge nes p rese nt for eve ry sep a ra tely h e rit a ble ch a ra c te r ,

an d m ore ove r these p a i red ge nes a re i nopp osite membe rs

o f the s a me ch rom os ome p a i r .

S E GR E G AT I O N O F C H R O M O S O M E S A N D G E N E S — In th e
reducti o nd ivisi on s wh ich p recede the f orm a ti ono f sp ores
i n pl a nts a nd g a metes i n a nim a ls th e fi rst eve nt o f i m
p ort a nce for th e p rese nt dis cussi on c onsists in the mem
be rs O f e a ch ch rom o s ome pa i r pl a cing themselves side by
S id e in th e p ro cess c a lled s yn a p si s Afte r a se ries o f de
.

v el op men ts (which do n o t c on ce rn us a t the p rese nt

m ome nt ) e a ch ch rom os ome p a i r fina lly a rra nges its elf on
the nucle a r spi ndle Th e sec ond sig nific a nt step is th a t
.

th e two membe rs O f e a ch p a i r sep a ra te or segrega te f rom

on e an o the r a n d go to opp o site p oles This sep a ra ti ono f
n
.

th e m a te rna l ch rom o s ome f rom its p a te r a l m a te a l s o

sep a ra tes the ge nes wh ich they c ont a i n He nce if the .

m a te rna l Ch rom o s ome o f a hyb rid pink f ou r O cl ock which ’

c ont a i ns the ge ne for red fl owe r c ol or g oes to one p ole

the p a te rna l ch rom o s ome c o nt a i ni ng th e wh ite ge ne ” “

g o es to the opp o site p ole This beh a vi or O f th e seg re

.

g a ti ng ch rom os omes th us d oes two th i ngs F i rst i t i nsu res .

th a t th e sp o res or g a metes p rod uced will get o n ly o ne o f

th e p a i r o f ge n es (red or white i nf o u r O cl o ck for ex a mple )
’

an dn o t b o th o f th em . I nth e sec on d pl a ce it p roduces th e

two ki n d s O f g a metes i nequ a l numbe r As we h a ve a l .

re a dy see nthis is p recisely th e s ort O f me ch a n ism requ i red

to p roduce the ra ti o i nF 2.

Inthe f o u r o cl ocks
’
D O M I N A N CE A N D T H E R ATI O .

the hyb rid be twee nred a nd wh ite is pi nk a nd so is re a d ily

di st i nguished f ro m eithe r p a re nt ra ce In m a ny c ro sses .
 D OM IN ANCE AN D T HE R ATI O 355

this i s not th e c a se Whe n a t a ll p ea is c rossed with a

.

sh ort one th e hyb rid is a s t a ll a s the t a ll p a re nt a nd

ordi na rily i ndisti nguish a ble from it Insuch c a ses t a l l .

Pa re nt A Pa re nt B

nu c l e i o f th e
Th e
p a re nts s h ow i n g on e
c h ro mo s o me p a ir w it h
sa me g ne i ne a c h c h ro
e
mo s o me o f th e p a i r .

u l
N c ei of th e ga met es p ro
d u ce d by th es e p a re nts wit h
o nly n o e c h ro mo s o me
of th e p a i r .

N uc l e u s of t h e h b ri d y
w ith th e two c h ro mos m s of o e

t h e p i h vi n
a r a g d i ff e re nt genes
f o r f l owe r c l o r ( b ov )
o . a e

u l u
N c e s of o n e
N u l u
c e s of th e
oth e rki nd of g a me t e
ki nd o f g a me t e
p ro d uc ed by h y b ri d
u
p ro d ce d by t h e h yb ri d

FI G 99
. A dia gra m to sh ow th e se grega ti ona nd
. combina ti onof
ch rom os om es inhyb rid pin k f ou r O clocks ’
.
356 M E ND E L S L AW S ’

i s s a id to b e domi na nt to sh ort a nd sh ort rec essi ve

to t a ll Wh e n th e t a ll hyb rid s a re self p oll i na ted th ey
.
-

p roduce a n F ge ne ra ti on which c onsists o f 3 t a lls to 1

2

sh ort Obvi ously th is result is due to th e f a c t th a t th e

.

pu re t a lls a nd the hyb rid t a lls l ook a like I f e a ch t a l l .

pl a nt is self p olli na t ed a nd its seeds s owed sep a ra tely it

-
,

is f o u nd th a t two thi rds O f them a re hyb rid s

-

seeds p roduce b o th t a lls a nd Sh orts in a ra ti o ) an d

on e thi rd a re p u re t a ll s (p rod uci n
-
g only t a lls ) Th e .

ra ti o is th e ref o re o n ly a speci a l c a se O f th e ra ti o

an d re a lly c onsists o f o ne fo u rth pu re d omi n

-
an ts
t a ll ) : two f o u rth s hyb rid d omina nts : one f o u rth pu re
- -

r ecessives .

T H E LAW O F S EGR EGAT I O N — He redity i n a simple

.

c ro ss betwee n two i ndivid u a ls i nv o lving a si ngle p a ir of

ch a ra c te rs ma y be summ ed up i nth e f oll owi ng st a teme nt
or la w :
Th e p a i red genes wh i ch a re resp onsi ble for th e cha ra c
ter difieren ce segrega te fro m o ne an o the r i nthe redu cti on

di vi si ons f o n
h y bri ds
d t h e reab y g i v e ris e t o e q u a l nu m
bers of ga me tes p u re fo r on e o r the o th er gen e; a n d the se
g a m e tes t h en c o mbi ne b y c h a n ce a t fe r ti li z a ti o n to p ro

du ce the F or secon . d hy bri d gen e ra ti on wh i c h wi ll con ,

si s t of a pp roxi ma tely on e fou rth of th e i n

-
di vidu a ls p u re
an d li ke on e gra n dp a rent on e fou rt h p u re a n
,
-
d li ke the
o the r gra n dp a rent a nd one ha lf h y bri d whic h ha lf wi ll
,
-
,

a lwa ys be li ke th e firs t h y brid gen e ra ti o na n d ei the r li ke

on e gra n dp a rent or i ntermedi a te be tweenthe two gra n d
p ents
a r
—
.

T H E R ELAT I O N O F G E N ES T O C H AR A CT ER S Th ough .

the l a ngu a ge u sed i nspe a ki ng o f ge nes a nd ch a ra cte rs is

such th a t th e stude nt migh t re a dily g a i nthe i mp ressi on
th a t a p a r ticul a r ge ne is th e s ole c a use o f a c orresp ondi ng
ch a ra cte r this is fa r f rom th e t ruth A moment s con
f
.
,

side ra ti o nwill S h ow th a t a f o u r O cl ock fl o we r ca nndt h a ve

’

a red c o l o r u n less m a ny ge nes h a ve fi rst w rough t thei r

coOp era ti ve eff ect to devel op the p l a n t a nd fl owe rs All .
 R E LATI O N OF GE NE S TO C HA R A C T E R 3 57

th a t is re a lly me a nt i nth e c a se u nde r dis cussi onby the

”
exp ressi on ge ne for red is th a t red fl owe rs a re p ro
“

duced by a c ombi na ti on o f ge nes th a t diffe rs i n respect ,

to a S i n gle p a i r o f gene s f rom th a t which p roduces white

,
.

Thus t his si ngle ge ne is r esp onsible for th e di fference

be twee nred a nd White bu t it is not by a ny me a ns solely
r esp on sible for red or white fl owe rs .

AR E G E N E S T H E S O LE B EAR ER S O F H ER EDI TY ? This -

questi on will receive a dequ a te a nswe r i n l a te r ch a pte rs

bu t it seems desi ra ble to p oi nt ou t he re th a t th e p rese nt
st a te of k nowledge of he redity rec og nizes only two me a ns
O f t ra nsmitti ng ch a ra cte rs from p a re nt to off sp ring Th e .

g re a t m a j ority O f ch a ra c te rs which h a ve bee ni nvestig a ted

bel ong to the cl a ss which a re t ra nsmitted th rough ge nes
i n th e ch rom os omes A ve ry few ch a ra cte rs i n pl a nts
.
,

h a vi ng to do with th e g ree nc ol or a re now k now nto be

,

t ra nsmitted from th e o vule p a re nt di rectly to th e Off

sp ri ng by me a ns O f th e chl oropl a sts which a re c ont a ined
i n the cyt opl a sm o f the egg The re a re m a ny p opul a r
.

ide a s a b o ut the i nflue nce O f th e m o th e r on h er u nb orn

child which ; if t rue w o uld not f a ll i neithe r of these two
,

cl a sses M o st o f these beliefs a re Obvi o usly f a nciful a nd

.

h a ve no b a sis i n f a ct a t a ll All su ch ide a s a s th a t th e

.

m oth e r ca n c ont rol th e m ora l or i ntellectu a l ch a ra c ter

O f the child by di recti n g h er own th o ugh ts a l ong S imil a r
li nes a re i na ll p rob a bility me re fi gments o f the im a gi na
ti on.

AR E A CQ U I RED CH AR A CTER S I N H ER I TED ? Ana cqui red

ch a ra cte r is one whi ch th e org a nism h a s a cqui red du ri ng
!

its ownlifetime a s a result O f its owna ctivities or the a c

ti onon it o f exte rna l i nflue nces Th e ch ief di fli cul ty i n
.

an swe ri ng th e a b o ve qu esti onlies i ndistinguishi ng ge nu

in e a cqui reme n ts S O l ong a s one expe rime nts with ch a r
.

a cters a b o ut which the re is n o d o ubt th e a nswe r is a lw a ys

i nthe neg a tive For ex a mple the sm a ll feet of the h i gh
.

cl a ss Chi nese w om a na re ce rt a i nly a cqui red Her imme .

di a te a ncest ors a nd h er rem ote ones h a ve regul a rly b ou nd

3 58 M E ND EL S L AW S
’

up th eir feet to keep them sm a ll B u t if a ny one inth e.

se ries d o es no t h a ve h er feet b ou nd they a re no t sm a ll .

Cle a rly this s ort o f a na cqui red ch a ra c te r is n

_
ot i nh e rited
.

But if a t ra i ned t rotti ng h orse begets o the r f a st t rotte rs it

is O fte np opul a rly supp osed th a t th ey h a ve inh e rited th e
eff ects O f the p a re nt s t ra i ning I t is fa r m ore p rob a ble
’
.
,

h oweve r th a t they h a ve me rely i nh e rited a si mi la r a bi li ty

’

to be tra i ned . In a n y ev e n t th e bi ol ogist h a s not yet

.

notwithst a ndi ng l ong c onti nued a nd p a i nst a ki ng exp eri

-

me nt a ti on bee n a ble to p roduce a ny resul ts wh ich a re

,

no t re a dily explic a ble eith e r onthe b a sis O f th e ch rom o

s ome mech a nism Or ons ome org a nor subst a nce c ont a ined
i nthe cyt opl a sm o f th e egg C e rt a i nf a cts a re h oweve r
.
, ,

k now na b ou t th e st ructu re a nd met a b olism O f cells w hich

migh t O ff e r a n expl a na ti on i n th e eve nt th a t the i n
h e rit a nce O f a ge nui ne a cqui red ch a ra cte r S h ould eve r
be est a blished .
 C HAPTER XX I X

THE LAW OF I ND E P E ND E N T A S S OCIATI O N

T H E I N DE P E N DE N CE OF C H AR A CTER S —FOI the s a ke .

‘

O f simplicity i nth e discussi onof S eg reg a ti ono u r a tte n ti on

h a s bee nc onfi ned to a si ngle p a i r o f ch a ra cte rs a nd thei r
c orresp ondi ng genes Of c ou rse th e two p a re nts O fte n
.

di ffe r i nrespect to seve ra l ch a ra cte r p a i rs Wh a t is t rue .

,

h oweve r i nrespect to one p a Ir 1 8 a ls o t rue of a ny othe r

,

p a i r Inorde r to illust ra te the i ndepe nde nce o f ch a ra cte rs

.

inexp ressi ona nd i nhe redity a t le a st two ch a ra cte r p a i rs -

a re n ecess a ry I f a p u re t a ll wh ite flowered p ea is c rossed

.
-

by a pu re sh ort p u rple flowered one th e hyb rid Off sp ri ng

-

a re a ll a like a n ll n ll r —
d a t a ll a d a pu ple flowered In this .

it is to be no ted fi rst o f a l l th a t neithe r p a re nt h a s d omi

na ted ove r th e o th e r a s a wh ole Th e hyb rid h a s i nh e rited
.

its h eigh t from one p a rent a nd its fl ower c ol or f ro m th e -

o the r I f a t a ll p u rple 1 s c rossed by a sh ort white the

“

hyb rid is a g a i nt a ll pu rple Inthis c a se it resembles one

.

p a re nt exclusively From th e two exp e rime nts i t bec omes

.

cle a r th a t the exp ressi on of p a rticul a r ge nes depends on

th ei r own p owe rs a nd not on wh ich p a re nt th ey h a ve
bee nde rived f rom .

T H E C O M B I N ATI O N S OF I N DE P E N DE N T CH AR A CTER S I N
F , If the hyb rid t a ll pu rple pe a s discussed a b ove a re
self p olli na ted to p roduce a sec ond hyb rid ge ne ra ti on it
-

will be f ou nd to c onsist O f a pp roxim a tely 9 t a ll p u rp l e ;

3 t a ll white ; 3 S h or t pu rple ; 1 S h ort white I t is to be .

no ted th a t two Of th ese c ombi na ti ons a re th e s a me a s th e

g ra ndp a rents a nd two a re new c ombi na ti ons Th e result .

is the s a me whethe r the t a ll pu rple hyb rids h a ve bee n

,

p roduced by c rossi ng t a ll wh ites by sh ort pu rples or t a ll

pu rple by S h or t white Th us no t only do these p a i rs of
.

genes for t a ll S h ort a nd pu rple white exp ress th emselves

- -

3 59
360 LAW OF I ND E PE ND E N T A S S OCI A TI O N
i ndepe nde ntly o f one a nothe r but a lso they rec ombi ne i n
,

F2 i ndepe nde ntly O f one a no the r .

T H E M E C H A N I S M O F R EA S S ORT M E N T is th e s a me a s
th a t i nv olved i nth e seg reg a ti onof a si ngle p a i r o f ge nes .

I t me rely a dds to the seg reg a ti ono f one p a i r th e si mul ta

neou s seg reg a ti ono f one or m ore a dditi ona l p a i rs The .

di a g ra ms Sh ow h ow th is is a cc omplished .

Pa re nts

Re d u ctio nd ivis io na nd
s e g re g a t i o nof two p a i rs
o f c h ro mo s o me s .

Ga metes

Ih e H
’

yb ri d

FI G 1 00 is
. dia gra m of redu ctiondivisi ons
a of t he t a ll white a nd
sh ort pu rple p a rents etc , .

T H E C H R O M OS O M E S T A LL P U R P LE HY B R I D
OF T H E .

Th e p ea h a s seve n p a i rs O f ch rom os omes only two o f ,

which a re c once rned with th e p rese nt expe rime nt All .

fu rth e r me nti onof th e o th e r five p a i rs will the refore be

omitted Wh e n the reducti on divisi on occu rs e a ch p a i r
.
,

seg reg a tes (a s sh ow ni nth e di a gra m ) so th a t th e g a me tes

receive on ly one membe r o f e a ch p a i r Th e g a mete o f .

th e t a ll wh ite thu s receives on e ch rom o s o me c on t a i ni ng

th e T a ll ge ne T a nd a no the r c ont a i ni ng th e white “

ge ne p S imil a rly th e g a mete O f th e sh ort pu rple receives

”
.

on e ch ro m o s ome with th e sh ort ge ne t a no the r wi th ,

th e Pu rple ge ne P Th ese a re th e o nly ki nds of

g a metes th ese p a re nts ca nf o rm S i nce the re is only one .

ki nd o f ga mete formed by e a ch p a re nt o nly o ne c ombi na

ti o nca nresul t : viz th a t sh ow n i n the di a g ra m for th e
.
,

nucleus o f th e h yb ri df I no ne p a i r one ch rom o s ome h a s

the ge ne T a nd its m a te th e ge ne t I n th e o th e r p a i r .
,
 GA M E T E S OF TALL PU R P L E H YB R ID 361

o ne h a s th e ge ne P a nd th e o the r th e ge ne p This sort O f .

di ff e re nce i nthe ch rom os ome p a i rs is ch a ra cte ristic of th e

nuclei o f a ll hyb rids Alw a ys one or m ore ch rom os ome
.

p a i rs h a ve a t le a st one ge ne i none ch rom os ome di ff e re nt

from th e c orresp ondi ng one i nits m a te .

T H E G A M ETES OF T H E T A LL P U R P LE HY BR ID Th e re .

d ucti on divisi ons i nth e t a ll white a nd sh ort pu rple p a r

e nts p roduce only one ki nd of g a mete for e a ch bec a use th e
membe rs of e a ch ch rom os ome p a i r a re ex a ctly a like In .

th e T a ll Pu rple hyb rid m ore th a none kind of g a mete is

formed b eca u se i nth e seg reg a ti onof its two ch rom os ome
,_

p a i rs di ffe re nt ge nes go to th e opp osite p oles of the

spi ndle Fu rthe rm ore it h a s bee nnoted inp revi ous ch a p
.
,

te rs th a t it is a pp a rentl y wh olly a m a tte r O f ch a nce wh ich

wa y a ch rom o s o me p a i r lies on the nucle a r spi ndle a nd ,

the ref ore a ls o a m a tte r o f ch a nce to which p ole eithe r of

its membe rs g oes Th e di a gra m sh ows th e two w a ys i n
.

T i
_ W
“ W
f
o
.
‘

FI G 1 0 1
. is a di a gra m of two w a ys of a rra ngi ng chromosomes Of
th e ta ll pu rpl e hyb r id on th e sp i ndl e .

which these two ch rom os ome p a i rs ma y be a rra nged on

the S pi ndle of th e reducti on divisi on Of the hyb rid i n
refe re n ce to one a nothe r .

P ositi on I p roduces two ki nd s of g a metes TP (T a l l

Pu rple ) a nd tp (sh ort white ) P ositi on I I simil a rly p ro
.

duces th e g a metes Tp (T a ll white ) a nd tP (sh ort

Pu rple ) S i nce one p ositi on on th e spi ndle is j ust a s
.

likely a s th e oth e r it f oll ows th a t f ou r ki nds of g a metes

,

a re p roduced i n equ a l n umbe r .

T H E CO M B I N AT I O N OF T H E F OU R K I N D S OF GA M ETES .

S i nce the hyb rid ca n p rodu ce f ou r ki nds of spe rms a n d

f ou r ki nds of eggs it is a pp a re nt th a t i t ca n give rise to
,

a s ma n y ki nds O f desce nd a nts a s th e re a re p ossible com

bi n a ti on s o f g a metes Th e di a gra m (c a lled a P u nnett
.
 3 62 L AW OF I ND E P E ND E NT A S S O CIATI O N
squ a re a fte r th e ma nwh o suggested its use ) a ll ows one
to dete rmi n e re a dily wh a t c ombi na ti ons a re p ossible a nd
i nwh a t ra ti o e a ch wi ll o ccu r Al ong th e top O f th e di a .

g ra m a re w ritte n th e f o u r ki nd s O f eggs a nd a l ong the

left side th e f o u r kinds O f spe rms Inth e sixtee nsqu a res .

a re w ritte nthe v a ri o us c ombin a ti ons which f ou r ki nds O f

spe rms ca nm a ke with th e f o u r ki nds of eggs .

FI G 1 02
. is P u nnett S qu a re sh owing recomb ina ti ons O f the ga m etes
of th e ta ll pu rpl e hyb rid .

T H E 9 : 3 : 3 : 1 R AT I OI t wa s st a ted i na p revi o us p a ra
.

g ra ph th a t th e re w o uld be f ou r c ombi na ti ons o f ch a ra c te rs

i nth e sec ond hyb rid ge ne ra ti onwh e nth e t a ll pu rple hy
b ri ds a re sel f p olli n a ted a n
-
d th a t th ey w o uld o ccu r i nthe
ra ti o 9 : 3 : 3 zl These f o u r types ca nbe re ad ily ide ntified
.

i nth e d i a g ra m E ve ry one which h a s ei the r on

. e or two

T a ll ge nes T will be t a ll a nd likewise eve ry o ne with

“

on e or two Pu rple ge nes P will be pu rple Thus N os . .

,

1 2 3 4 5 7 9 1 0 1 3 a re a ll T a ll Pu rple a n
, , , , , , , , d a re n i ne ,

o u t o f th e to t a l o f sixtee n S imil a rly N os 6 8 1 4 a re

. .
, , ,

T a ll white a nd a re th ree o ut o f sixtee n N os 1 1 1 2 1 5 . .

, ,

a re sh ort Pu rple a n d th ree o u t O f sixtee n N o 1 6 is sh ort . .

wh i te Th is ra ti o is re a dily see n to be a ge ne ra l one

.

for a n y two i n depe nde nt d o mi na nt ch a ra cte rs .

G E N OT Y P ES AN D P H EN OTY P ES Th is d i a g ra m a ls o .

m a kes cle a r a no th e r imp o rt a nt f a c t Th e re is only one .

 L AW OF I ND E P E ND E N T A S S O R T M E N T 3 63

pl a nt ou t O f sixtee nwh ich is white a nd sh ort I t h a s b o th .

recessive ch a ra cte rs an d is pu re (ttp p ) f or b o th

,
Th e .

th ree S h ort pu rples h oweve r a re not a ll a like N O 1 1 is

, , . .

pu re for b o th ch a ra cte rs (ttPP ) bu t b o th 1 2 a nd 1 5 a re

pu re only for sh ort (tt ) a nd hyb rid for pu rple (P p ) Th e .

s a me di ffe re nce is to be no ted a m ong th e t a ll whites a nd

t a ll pu rples ; only one o f e a ch g roup is re a lly pu re the ,

o th e rs bei n g hyb rid f or one or b o th ch a ra cte rs F rom .

this it a ppe a rs th a t th e re a re m ore comb i na ti ons o f ge nes

’

th a na re rec ogniz a ble from the a ppe a ra nce of the pl a nts .

Inorde r to disti nguish th e combi na ti ons or ge nes f rom ,

the c ombi na ti ons o f ch a ra cte rs bi ol ogists give th em

sep a ra te na mes Any p a rticul a r combi na ti onof genes is
.

S p oke nO f a s a gen o typ e wh e re a an m b i n t i n f h

“

o a r
s yp ,
o a o c

a cters is c a lled a p h en o typ e th a t is t o sa y visible type

, ,
.

The re a re sh ow ni n th e Pu nnett squ a re 4 phe no types or

visibly disti nguish a ble types T a ll Pu rple T a ll white , ,

sh or t Pu rple a nd S h ort wh ite ) but the re a re ni ne ge no

, ,

types Or c ombina ti ons of ge nes a s f oll ows : (1 ) TTP P ,

.

(2 ) TT P p ( N o s 2 a n d (3
.
) T t P P ( N OS 3 a n d .

( ) Tt
4 (N O S 4 7 1 0 .
,
a l l O f wh ich l ook a like
, ,

an d a re t a ll a nd pu rple ; (5 ) TTp p (N O (6 ) T tp p
.

NOS 8 .
,
a l l t a ll a n d wh ite ; (7 ) ttPP (N O (8) .

tt ( N os 1.2 1 5 ) a
,
ll S h o r t p u rple ; a n d ( ) pp (
9 t t N o 1 6 ) .

sh ort white .

T H E LAW OF I N DE P E N DE N T A S S OR T M E N T OF GE N ES
AN D C H AR A CTER S Wh en two p a rents whi ch di ffer i n
.

resp ect to two di ff eren t cha ra cter p a i rs a re crossed th e -

hy bri d wi ll exh i bi t bo th domi na nt cha ra cters i rresp ecti ve

of whi ch p a ren t con tri bu tes th em .

When the hy bri d forms i ts ga me tes the i ndep endent

g en e s ,wh i c h a r e i n s ep a r a te c h ro m o s o m e p a i rs s eg g
re a te ,

an d form fou r ki n ds of ga me tes .

At ferti li z a ti onthe fou r ki nds of ga me tes u ni te to form

ni ne di fierent genotyp es or combi na ti ons of genes a nd
n
,

wh enbo th cha ra cter p a i rs s how domi na nce fou r di ffere t

-

p h en o typ e s o r co m bi n a t i o n s o f ch a r a c te rs .
 C HAPT ER XXX

LIN K AG E PH E N O M E NA

TH E T H I R D LAw Limi ta ti onof I ndepe nde nt Ass ort

.

me nt Inth e l a st ch a p te r i t wa s sh ow nth a t m a ny ch a r
.

a cte r p a i rs a ss o r t th emselves i n
-
depe nde ntly of one a n
o th e r . Bu t S i nce this i ndepe nde nce is d ue to th ei r
r espective ge n es being ca rried i n di ffe re nt ch rom os ome
p a i rs it is cle a r th a t the re ca n be no m ore i ndepe nde nt
p a i rs O f ch a ra cte rs th a n the re a re p a i rs of ch ro m o s omes .

For ex a mple the re a re seve n p a i rs O f ch rom o s omes i n

,

p ea s. M e ndel f o u nd j ust seve n p a i rs O f i ndepe nde nt

ch a ra cte rs nor h a s th e exte nsive w ork since his t im e
,

disc ove red a ny m ore This f a ct is a ls o t rue of a ll o th e r

.

pl a nts a nd a nim a ls so fa r studied I t ma y be st a ted a s a

.

th i rd l a w o f h e redity The nu mber of p a i rs of cha ra c

ters whic h ca na t the sa me ti me a ssort i n dep endently i n
a n y orga ni s m i s e q u a l to th e n u m b e r of c h ro mos o me p a i rs

i n i ts ce l ls .

T H E F O URT H LAw C ha ra cters wh ose genes a re lo

.

ca ted i n t h e s a me ch ro mos o me a re li n ked i n heredi ty .

B o th the thi rd a nd f o u rth l a ws a re Obvi o us c o nseque nces

o f th e beh a vi o r o f the ch ro m o s omes S i nce these ch ro m o
.

s omes a re defi ni te i nnumbe r a nd m a i nt a i nthei r i ndi vid

u a l i ty th e t o t a l n
,
umbe r O f ch a ra c te rs must b e a ssorted
'

i n a n equ a l numbe r o f li n ked g ro ups Th a t this is a c

.

t u a lly t rue is bei n g a bu nd a ntly p roved by i nvestig a ti o ns

with b o th a nim a ls a nd pl a nts G roups o f li nked ch a ra c
.

te rs a re a l re a dy k no w n i n seve ra l pl a nts i ncludi ng the

,

sweetpe a i n wh ich the phe nome non wa s origi na lly dis

co ve red Li nk a ge i s a ls o know n i n seve ra l a nim a ls
.
,

including ma n .
 THE VIN E GA R FL Y 3 65

T H E V I N EGAR FLY I nvestig a ti ons i n he redity h a ve

.

p rob a bly m a de g re a te r p rog ress i nthe l a st d oze nye a rs or

so th a ni na l l p revi o us time Th is h a s i n l a rge me a su re
.
-

bee n m a de p o ssible th rough th e use of th e vi neg a r fly ,

D ros ophil a This insect (see Fig 1 0 5 ) is sm a ll a nd b reeds

. .

re a dily i nc a ptivity It mut a tes (see p 3 50 ) f reque ntly

. .

an d its life is S h or t Ina little ove r tenye a rs m

. ore ge ne ra
ti ons of th is fly h a ve bee nre a red a nd c a refully Obse rved
u nde r the mic rosc op e th a n th e re h a ve bee n ge ne ra ti ons
of men si n ce the d a w n of rec orded h ist ory .

The nucleus (Fig 1 03 ) O f the cell of t h e vi nega r fly is

.

a lm o st ide a lly suited for studies i nh e redity Inthe fi rst .

FI G 1 03
. . T h e N u cl ei of th e C ells o f V in e ga r Fli e s T he fi g re
. u
to t h e l ef t sh ows t h a t th e fem a l e h a s 4 p a i rs O f ch ro m oso m es i nea ch
ce ll nu cleus a nd th a t th e m emb ers of ea ch p a ir a re visibly al ik e T h e .

righ t- h a n d figu re sh ows t h a t th e m a l e h a s t hr ee p a i rs li k e t h ose of

th e f em a l e b u t t h a t t h e m emb ers of th e l ow er p a i r a re vis ibly dif
ferent fro m on e an o t h er S O f a r a s inve s t iga ti ons h a v e go ne up to
.

th e p resen t th e on e wi t h t h e h o o k (t h e Y -
c h ro m o s o m e s o -
ca l l e d )

i s no genes or h eredi ta ry fa c tors In certa in a bnorm a l fli es

ca rr e .

i t is enti rely missing a nd t h a t a pp a rent ly wi th o u t m a ki ng a ny

e ssen ti a l di ff erence i n h eredi t y .

M organ Stu rtev a nt M u ller Bridges M echa ni sm of M endeli a n

, , , ,

Her edi ty .

pl a ce it h a s only f ou r p a i rs of ch rom os omes a nd he nce

ca nh a ve on ly f ou r gro up s O f li nked ch a ra cte rs In the .

sec ond pl a ce its ch rom o s ome p a i rs ca nbe disti nguished

,

from one a noth e r by th ei r f orm a nd size This h a s bee n .

3 66 LIN K AG E P H E N O ME NA

o ne of the me a ns by wh ich it is p ossible to tell i nwh ich

ch rom os ome p a i r th e ge nes for th e d iff e re nt li nk a ge
group s of ch a ra cte rs a re l oc a ted Th e fi rst p a i r a re .

st ra igh t rods th e sec ond a nd thi rd ones a re be nt rods

, ,

an d th e f o u rth p a i r a re sm a ll rou n d b odies .

T H E SEX C H RO M OS O M ES O F D R OS O P H I L A Inthe fe .

m a le th e two membe rs Of th e fi rst p a i r O f ch rom o s omes

a re i n distinguish a ble a nd a re c a lled X ch rom os omes In .

th e m a l e one of these ch rom os omes is sligh tly di ff e re nt

,

f rom its m a te One O f them is like th e p a i r inth e fem a le

.

an d is i nf a c t a nX ch ro m o s ome Th e o the r one is c a lled

.

th e Y ch rom os ome a nd is no t k now nto c a rry a ny ge nes

,
.

T H E D ET ER M I N AT I O N O F S Ex Whe nth e vi neg a r fly

.

f orms its eggs th e two X s seg reg a te f rom one a nothe r

,
’

a l on g with the membe rs O f th e o the r th ree p a i rs so th a t

eve ry egg receives one _X Inth e m a le the X seg reg a tes
.

from th e Y So th a t one h a lf th e spe rm s receive a nX a nd

th e o th e r h a lf receive a Y Th e re a re thus two ki nds of
spe rm
.

s a nd bu t one O f eg g s I f a n X c ont a i ni ng Spe rm

.
-

fe rtilizes a n egg a fem a l e results XX ) but if a ,

Y c on
-
t a i ni ng spe rm fe rtilizes i t a m a le (X Y ) resul ts .

This ge ne ra l meth od O f sex dete rmi na ti o na p pe a rs to be

-

c omm on a m ong i nsects bi rds a nd m a mm a ls i ncl udi ng

'

, ,

ma n .

L I N K AGE I N V I N EGAR FLY CR OS S ES M a ny h u nd reds

.

o f expe rime n ts h a ve bee nc a rried ou t i nstudies O f li nk a ge

with this i nsec t so th a t th ere is now a ve ry l a rge b ody
,

o f d a t a wh e re a s o u r k n
, owledge o f this phe n ome n onis fa r

less c omplete inrespect to a ny o the r a nim a l or pl a nt .

Fo r this re a s o n th e f oll owi ng p a ra gra ph s will d e a l wi th

vi neg a r fly c rosses with th e u nde rst a nd ing th a t th e p ri n
ci p l es d evel oped a re inge n e ra l a pplic a ble to othe r a nim a ls
an d pl a nts .

T H E B A C K CR O S S is a meth od of b reedi n g i nwh ich th e

fi rst ge ne ra ti onhyb rids a re m a ted to pu re recessives i n
ste a d o f to one a no th e r I t is much used i n a na lyzi ng
.

th e ge ne ti c c omp osi ti ono f hyb rids bec a use it gives simp le r

 B A CK CR O S S 3 67

r esults th a nm a ti ng hyb rids with one a noth e r i nth a t th e ,

recessive p a re nt being p u re p roduces only on

,
e ki n,
d Of
g a mete M ore ove r since these g a metes h a ve only reces
.
,

sive genes they will not obscu re a ny d omi na nt ge nes of

th e hyb rid with which they u nite Th e b a ck c ross i s .

th us p a rticul a rly well suited for studying li nk a ge ph e

nomena Th e next p a ra g ra ph will S h ow h ow it w orks
.

i n c rosses i nv olvi ng i ndepe n de nt ch a ra cte rs .

B A C K CR OS S I N V OLV I N G I N DEP E N DE N T CH AR A CTERS .

Whe nth e hyb rid t a ll p u rple pe a s discussed i nth e p reced

in g ch a p te r we re m a ted to o ne a no the r or self p o lli na ted -

the sec ond hyb rid ge ne ra ti onc onsisted of f ou r phe no types

i nth e ra ti o O f 9: 3 : 3 1 I f this s a me hyb rid is b a ck c rossed
.
-

to th e d o uble rec essive S h ort white the s a me f o u r phe n

,
o ,

types will be p rod uced but i nthe ra ti o Th e t a ble

S h ows why this is SO .

Fou r k i nds of e ggs whi ch a re p rod u ce d i nequ a l nu mb ers

by th e ta ll pu rp l e hyb rid
,
.

FI G 1 0 4
. . P unne tt S qu a re (sh owi ng res u l ts of b a ck -
cro ssi ng a ta ll
pu rple hyb rid to a pu re sh ort w hi te) .

L I N K ED CH AR A CTER S I N T H E F IR S T HY BR ID GEN ERA

T ION beh a ve j ust like i ndepe ndent ch a ra cte rs I n th a t it ,

m a kes no di ffe re nce from which p a re nt th e d omina nt

ch a ra cte rs c ome For ex a mple (1 ) if a l ong wi nged g ra y
.
,
-

b odied fly is m a ted to a vestigi a l wi nged bl a ck b odied one - -

the hyb rid Offsp ri ng will a ll be g ra y b odied a nd l ong -

wi nged bec a use these two ch a ra cte rs a re d omi na nt (Fig .

(2) If th e s a me two ch a ra cte r p a irs a re i nt roduced -

i nto the c ross by m a ting a g ra y b odied vestigi a l wi nged - -

fly with a bl a ck b odied l ong wi nged one th e result will

- -

still be g ra y b odied l ong wi nged hyb rids j ust a s would

- -
,
36 8 LI NKAG E PH E N OME NA
be the c a se if the exp e rime nt c once rned indepe nde nt ch a r
a cters (Fig Al th ough G ra y L ong x bl a ck vestigi a l
.

an d G ra y vestigi a l x bl a ck L on g give ide ntic a l fi rst gen

FI G 1 05. Il lustra ti ng th e Be h a vi o r O f Li nke d C h a ra c te rs inthe

.

vinega r fly A bl a ck-b o die d fly wi th ve s tigi a l wi ngs is ma te d to a

.

gra y b odi e d fly wi th l ong wi ngs T he fi rs t hyb rid ge ne ra ti o nis lo ng

-
.

winge d a nd gra y b o die d Whe nth is is b re d to a bl a ck ve s tigi a l fly

-
.

b k c osse d to th e d o ubl e rece ssive t ype ) th e gra n dp a renta l

'

(.i e . a c - r

type s a ppe a r in th e se co n d hyb rid genera ti o n In the ch rom o some

.

dia gra ms the ge ne b for bl a ck b ody is conta i ne d i nthe sa m e ch rom o

so m e wi th th e ge n e v fo r ve s tigi a l wi n gs so th a t the two rem a in
li nk ed to ge th er th rough ou t a ll the ce ll divisi ons o f grow th a nd
th rough t he re du ct i o n divisi on s wh ich p rodu ce the ga m ete s , a nd
t h ere fo re re a p pe a r to ge th er a g a i n i n t he se co nd hyb rid genera tio n .

T he s a m e is a lso , o f co u rse , t ru e fo r the d o mi n ant ge ne s B fo r gra y

bO d y a nd V fo r l ong wi ngs .

Morga n, The P hysi ca l Ba si s of Heredi ty Lipp inco tt

~

. .
 LIN K E D C HA R A C T E R S 3 69

cra ti on hyb rids th ese hyb rids do not (a s hyb rids with
,

i ndepe nde nt ch a ra cte rs do ) give the s a me phe notypes i n

th e sec ond hyb rid ge ne ra ti oneithe r wh enb a ck c rossed or -

wh en m a ted with one a no the r, a s is set f orth in det a il

inth e nex t p a ra g ra ph .

B A C K CROS S I N G L I N K ED C H ARA CTER S — I f th e l ong

-

FI G 1 06
. . T h e C ro ss B etw een Ve s tigi a l Gra y a n d Long B l a ck .

T h e symb o ls use d i nt his di a gr a m a re th e s a m e a s t h o se i n t h e l a st

on e . Howe ver i t S h ould b e no te d t h a t i nt his cro ss ea ch p arent con
tribu te s o n e d o mi n an t gen e , wh ere a s i n th e l a s t , o ne p a ren t con
tribu te d b o t h d o min an ts Inthis ca se th e fi rs t hyb rid ge n
. era t i onw i ll

sh ow b oth d omina nts a nd so , t h ere ore, resemble one p a rent in

f
resp ect t o b o dy co l or a nd th e o t h er i nre sp ect t o wi ng l engt h . As th e
di a gra m S h ows t his doe s not i nv o lve a ny distu rb a nce of th e link a ge
since th e gra ndp a renta l co mbina tions of genes a nd t he ir corre sp ond
ing ch a ra cters rea pp e a r a ga i ni n th e se cond hyb rid gen era t i on.

M org a n, P hys B a si s
. .
 70 LIN KAG E PH E N OME N A
w inged g ra y b odied hyb rid m a le flies f rom the fi rst type
-

O f c ro ss whe re the t wo d omi n

, an t ch a ra c te rs G ra y
an d L ong ) ew e f rom the s a me p a re nt a re m a ted to ,

d ouble recessive fem a les bl a ck vestigi a l ) the re w i ll

be j ust two types of Off sp ri ng ; viz G ra y L ong a nd bl a ck .
,

vestigi a l like thei r g ra ndp a re nts I f f ree a ss ortme nt h ad .

o ccu rred the re w o uld h a ve bee n i n a dditi o n rec ombin a

ti oncl a sses : viz G ra y vestigi a l a nd bl a ck L ong a nd a ll

.
,

f ou r cl a sses i n equ a l numbe rs a s wa s expl a i ned in a ,

p recedi ng p a ra g ra ph for the p ea c ross .

I f the sec ond (2) type of g ra y l ong hyb rid m a les a re

c rossed to bl a ck vestigi a l fem a les they p rod uce a ls o o nly
two cl a sses O f off sp ri n g : viz G ra y vestigi a l a nd bl a ck .
,

L ong like thei r g ra ndp a re nt s a nd no rec om bi na ti on

cl a sses .

Inb o th expe rim e nt s the re is a t o t a l f a ilu re to p rod u ce

th e rec ombi na ti on cl a sses to be expected a cc ord ing to
the l a w O f f ree a ss ortme nt I n o the r w ords th e ch a ra c.
,

te rs th a t we nt i nt o the c ross t ogeth e r h a ve st a yed to

gethe r or S h ow c omplete l ink a ge Th e a cc omp a nyi ng .

di a gra m a nd th ose i nth e nex t p a ra g ra ph S h ow why these

r esul ts a re O bt a i ned .

S p erms of the two s or s t

o f gra y long hybri ds .

C a se Gra y Lo ng sperm X Gra y Lo ng F2

(1 ) i b la c k ves tigia l sp e r mx bl a ck v es tigi a l F2

C a se Gra y ves ti gi a l sp erm X G ra y v estigi a l F2

(2) i b la ck L ong sp erm X bl a ck Lo ng F2

T H E S EGREGAT I O N L I N KED G EN ES S i nce it h a s

OF .

a l re a dy bee n sh ow n th a t ge n es which a re i n diff e re nt

ch rom os ome — p a i rs a ss ort indepe nde ntly it is evide nt th a t
ch a ra c te rs which do no t a ss ort freely must be i nth e s a me
ch rom o s ome p a i r The a cc omp a nyi ng di a g ra ms (Fig
-
. .

l 0 7a a n d b ) sh ow th e l oca ti ons o f th e ge nes f or vestigi a l

wi ng a nd bl a ck b ody c ol or i nthe sec ond ch rom o s ome p a i r -
 T HE S T R E NGTH OF L I N K AG E 371

a nd th e wa y they a re dist ributed by segreg a ti on i n the

two c ro sses u n de r discu ssi on .

T H E S TR E N GT H O F LI N K AGE In the l a st p a ra g ra ph p

m a le hyb rids of the vi neg a r fly we re b a ck c rossed to -

fe m a le reces si ves bec a use the li nk a ge O f ge nes i n the

ch rom o s omes O f m a le vi nega nfli es is c omplete i e the re ,
. .
,

is neve r a ny sep ara ti onof th e ge nes i nthe s a me ch rom o

” '

s ome a nd c onsequently the ch a ra cte rs th a t origi na lly

,

S p er m m ot h e r ce l l o f Eg g m o t h e r c e l l of
Pa re nt I Pa re n t ll
Gra y l
b a ck
Lo ng v e s t ig ia l
GL GL ] gl
‘

g _

Th e Gra yLo ng Hyb ri d

( ma l e)
GL gl

( bl a ck ve s tig ia l )
5 !

FI G 1 07. .
(a ) S h owi ng th e se gre ga tion of li nk e d genes in gra y
long hyb rid of t yp e GL gl .

we nt i nt o th e c ro ss c a me ou t t ogeth e r a g a i n with out f a il

“

in the sec ond hyb rid ge n e ra ti on This peculi a rity Of .

link a ge i nm a le V i neg a r flies viz th a t l i nked ch a ra cte rs

,
.
,

neve r sep a ra te is not t rue for th e fem a le hyb rids or for

,

m a les of a l l species O f a nim a ls By m a ti ng a fema le .

(Fig 1 08) hyb rid G ra y L ong vi neg a r fly O f type (1 )

.

the Off sp ring of G ra y L ong a nd bl a ck vestigi a l ) with a

3 72 LIN K AG E P H E N O M E NA

d ouble recessive a bl a ck vestigi a l ) m a le a ll fou r , ,

phe notypes occu r i n the sec ond hyb rid ge ne ra ti on but ,

they do not occu r i n the ra ti o to be expected i n

the c a se o f f ree a ss ortme nt I f a fem a le hyb rid o f type .

ffsp i ng f g ra y vestigi a l by bl a ck l on g ) is
( )
2 O r o

m a ted to the d ouble recessive bl a ck vestigi a l ) m a le ,

Th e g ra y lo ng h yb ri d

(m a le )
GI g L

m
S pe r s of th e
h yb fi d

Gra y vesti g ia l
GI

FI G . 1 07 .
—(b) S ho wi ng th e se grega ti onof l ink ed genes in gra y
l o ng hyb rid of t ype Gl gL .

th e sa me f ou r ph e notyp es occu r bu t th e ra ti o s a re a g a i n
di ff e re nt Inth e fi rst c a se th e re a re a b o u t five times a s
.

m a ny G ra y L ong a nd bl ac k vestigi a l flies g ra nd

p a re nt a l c ombi na ti ons ) a s the re a re G ra y vestigi a l a nd
bl a ck L ong or rec ombi n a ti oncl a sses Inth e sec o n d c a se .

the re a re a ls o a b o u t five times a s m a n y g ra ndp a re nt a l

t ypes this t im e G ra y vestigi a l a nd bl a ck Lo ng ) a s
 T HE S T R E NGTH OF LINKAG E 373

FI G . 1 08
. C rossing- O ver i n a Cro ss B etweeiI B l a c Ves tigi a l
k
an d Gra y Long . This exp eriment is li ke th e l a s t two ex cep t th a t
i nt his ca se a fe m a l e hyb rid is b re d to a d ouble recessive m a l e . Th e
second line sh ows th a t f o ur kinds of eggs a re p ro du ced a s if free
a ss ortm en t h a d occ rre du Th e diff erence li e s inth e f a ct t h a t t h ere
.

a re m a n y m ore eggs wi th th e origi na l combina ti on Of genes th a n

wi th th e recombina ti ons a s is sh ownby th e f a ct th a t i nth e second
hyb rid genera t ionth ere a re a b ou t fi v e t imes a s m a ny gra ndp a renta l
t yp e s a s t h ere a re re co mbi n a t i on t yp e s
. C ro ssing-o v er o f gen e s in

t h e vi n e g a r fly h a s occu rr e d up to t h e p resen t on l y i n t h e f em a l e
an d n o t i n th e m a l e . m
Hence f e a l e hyb rids a re se d wh en i t i s
u
desire d to dem onstra te cro ssing-o ver a nd m a les wh en link a ge onl y
is to b e sh own .

M orga n, P hys B a si s. .
 74 LIN KA G E P H E N O ME NA

r ec ombi na ti on types O the r ch a ra cte rs Sh ow th e s a me

.

s ort o f phe nome na but the st re ngth o f li nk a ge v a ries with

th e p a rticul a r ch a ra cte rs .

CR OS S I N G OV ER is th e tech nic a l na me give n to the

-

b re a k a ge O f li nk a ge betwee n ch a ra c te rs wh ose ge nes a re

c a rried i nthe s a me ch rom os ome p a i r I nthe p recedi ng -
.

p a ra g ra ph s the ch a ra cte rs bl a ck b ody a nd vestigi a l

,

wi ngs we re sh ow nto be li nked a nd d u ri ng th e f orm a ti on

,

o f g a metes usu a l ly to rem a i nt ogethe r but to sep a ra te i n ,

a ce rt a i np rop orti onO f th e g a metes S ince li nk a ge is due

.

to th e ge n es bei ng i nth e s a me ch rom os ome it is evide nt

th a t the re must be s ome me a ns by which a ge ne ma y be
exch a nged f ro m one ch ro m os ome to i ts m a te .

TH E M E CH A N I S M O F CR O S S I N G O VE R -
Ina ve ry e a rly
.

st a ge O f th e rediI cti on divisi ons i n th e m othe r cells o f

g a metes in a nim a ls (a nd sp ores i n pl a nts ) the ch rom o
s omes a re d ra w n ou t i nt o sle nde r th re a ds A t this time .

th ey bec om e as s oci a ted sid e by side th ro ugh out thei r

e nti re le ngth i nth e p ro cess c a lled sy na psis Th ey bec ome .

so cl o sely twisted a b o u t on e an o th e r th a t i t is difficult to

disti nguish one f rom th e o th e r D u ri ng th eir l a te r .

devel opme nt th ey bec ome m uch sh orter a nd thicke r I t .

is supp osed a nd th e re is much i ndi rect evide nce to p rove

, ,

th a t wh ile they a re twisted a p a rt o f one ma y c ome to

repl a ce a c o rresp on di ng p a rt o f its m a te The di a g ra m .

Fig sh ws h w c ssi n g ve ff ects th e p o si ti on

( . 1 1 0 ) o o ro o -
r a

o f the bl a ck a n d vestigi a l ge nes i na p a i r of ch rom os omes

Of a hyb rid vi n eg a r fly .
T H E F I FT H LAW O F H ERED I TY ma ybe st a ted a s f oll ows
Li n ked gen es m a y c r os s-
o v e r fr o m o n e c h ro m o s o m e o f a

p a i r to i ts m a te b y e x c h a n g e o f c o rr e sp o n d i n g s e g m e n t s

of the p a i red c h ro moso mes .

TH E S I X T H LAw st a tes th a t t h e ge n e s i n a c h ro m o so m e

a re a rra n g e d i n l i n ea r o r d e r This
. l a w r ests o n the f a ct
th a t th e re is a defi nite pe rce nt a ge o f c rossi ng o ve r be twee n -

an p
y two a r O f i s ge n a
es n d th a t i t is g r e a te r th e fu r th e r

a p a rt they a re . This is o f c ou rse O bvi o us f ro m th e

, ,
 SE X LIN KAG E 375

di a g ra m for th e f a rthe r a p a rt th e ge nes a re the m ore

,

ch a n ces th e re a re th a t a b re a k will occu r betwee nthem .

SEX L I N K A GE S i nce the sex o f a ni ndividu a l i nm a ny

.

an im a l species is dete rmi ned by its ch rom o s omes it w ould

na tu ral ly be expected th a t s ome ch a ra cte rs w ould be

FI G 1 09
. Cr o ssi ng O ver i n Cross B etw een Gra y
.
-
Ve s tigi a l a nd
B l a ck Lo ng Th e symb ols a n
. d re sul ts a re t h e s a m e as i n th e l a s t
di a gra m e xcep t th a t th e genes a re li nke d diff erently . T h e se cond
hyb rid genera t i onsh ows a ga ina b ou t 5 tim es a s m a ny gra ndp a renta l
typ e s (h e this t im e gra y v es t igi a l a n
. d bl a ck l ong) a s recombina
t i o n typ e s .

M orga n Phys B a si s
,
. .
376 LIN K AG E P H E N O ME N A

linked i nth e sex ch rom o so me This is i n fa ct t rue a nd .

we ma y now p roceed to ex a mi ne this m a tte r m ore cl o sely .

S ince th e vi neg a r fly h a s 2 X s i nthe fem a le a nd one X ’

an d on e Y i nthe m a le it is evide n t th a t th e fem a le ca n

,

be p u re h a ve th e s a me ge nes i nb oth ch rom os omes )

FI G . 110 — D i a gra m to sh o w cro ssing o ver O f genes I sh ows

.
-
.

th e a rra n gement of genes inth e 2nd p ai r of chr omos omes of the hy

b rid b efore cross ing o ver h a s occurred I I S h ows the recombi na t ions
- .

a f te r cross in g over h a s occu rred S egrega tionof the chromos omes inI
- .

wi ll give ga metes lik e th e p a rents ; of th e ones inI I wi ll giv e reco m

b in a ti o ns . In th is p a rti cul a r insta nce cross ing O ver o ccu rs ina bo u t -

17 o f t h e g a m e tes B = gra y b o dy ; b b la ck b ody ; V = lon

.
g w ings ; z

v = v es ti gi a l win gs .

f or ge nes c a rried i n th e X ch rom os omes b u t th a t the ,

m a le c a nnot bec a use h e h a s o nly o ne X a nd the Y h a s

,

no t bee n sh ow n to c a rry a ny ge nes An illust ra ti on o r .

two will m a ke the m a tte r cle a r .

R ED EY ED F E M A LE X W H I T E—
-
E Y ED M A LE FLY (Fig .

Th e ge ne for eye c ol or i nthis c ross is l o c a ted i n

-

th e X ch rom os ome R ed is d omi n ant to white Th e

. .

re d eyed fem a le p roduces eggs a l l o f wh ich h a ve o n

-
e X

ch rom os om e c onta i ni ng th e ge ne for red eye H a lf th e -

.

spe rms h a ve a nX wi th th e ge ne for wh ite eye a nd h a lf

”
-

h a ve th e Y ch rom o s ome Whe n wh ite X spe rms fer .

-

ti l i z e th e red X eggs red eyed fem a les result ; bec a u se

-
,
-

th e red is d omi n an t t o the wh ite Whe nY spe rms fer

‘
-

”
.

ti l i z e red X eggs red eyed m a les result bec a use the

”
- -
, ,

on ly X p rese nt h a s a red ge ne “
.
 —
R E D E Y E D FE M AL E 377

Wh a t:

SPERM

FI G . 111 .
-
Sex-Link a ge . ss invol ving th e gene s red a nd
A cro
w hi te eye whi ch a re l oca te d i nth e sex ch rom osome s Th e X ch ro
, .
-

mosomes O f t h e fem a l e p a rent a re co l ore d bl a ck t o indi ca te th e

p resence of th e gene for red eye a nd th e X ch rom osom e o f th e m a le
- -

lef t whi te to indi ca te th e white gene T h e Y ch rom os ome of th e

.
-

m a le is a lso left bl a nk b eca u se i t d oe s not conta ine i th er gene T h e .

di a gra m S h ows th a t the fi rst genera t ion m a le s a nd fem a l es a re b oth

red e ye d b e ca use ea ch rece iv e s a gen
-
e f or re d ey e In th e second
- .

genera t i on a ll fem a l es a re red e yed since th ey h a ve ei th er one or

-

two ch ro m o s o m e s wi t h th e red gen e wh erea s on ly h a lf th e m a l es

receive th e ch ro m o s o m e s wi t h th e red g en e T h e o th er h a lf rece ive
.

a ch ro m o s om e wi t h th e whi te ge n e an d a re whi te e y e d -
.

-M orga n S tu rtev a nt M uller B ridges M ech a ni s m of M endeli a n

,

I—
, , ,
H cr e di t I nl t
37 8 LI N KAG E PH E N O ME NA
R ed eyed hyb id fem a les X
-
eyed m a les If the h y
r red- .

brid red eyed fem a les o f th e l a st p a rag ra ph a re b red to

-

th ei r red eyed b ro the rs (or a ny o the r red eyed m a les ) a ll

- -

the d a ughte rs a re red eyed bec a use they receive a d o mi -

na nt red X f rom th e f a th e r a nd eithe r a red o r a

“ “

“ ”
white X f rom th e hyb rid m othe r H a lf th e s ons a re
”
.

red eyed bec a use they receive a red

“
-
X f rom thei r
m o the r a l ong with th e i na ctive Y f rom the fa th e r Th e .

o the r h a lf o f the s ons a re white eyed bec a use they receive -

a wh ite X f rom th ei r m oth e r a nd a ni na ctive Y f rom

th e f a the r Th e a cc omp a nying figu re (1 1 1 ) sh ould m a ke
.

this p o i nt cle a r .

R ED EY ED M A LE X W H I TE E Y ED F E MA LE
-
R efe re nce -
.

to Fig 1 1 2 sh ows th a t th e hyb rid d a ugh te rs h e re receive

”
.

a
“
white X f romth e m oth er a nd a red f rom th e
f a th e r a nd a re the ref ore red eyed Sons receive a Y
,
-
.

f rom f a th er a nd a white X from m oth e r a nd a re the re

f ore white eyed I f th ese wh ite eyed m a les a re b red to
-

'
.
-

thei r h yb ri d red eyed sis te rs h a lf the d a ugh te rs a re red

-

eyed hyb rid s a nd h a lf a re p u re white eyes Likewise -

.
,

h a lf the s ons receive a red X f rom thei r hyb rid m othe r

an d a Y f rom th ei r f a th e r a nd a re red eyed a nd h a lf re -
,

cei v e a wh ite X f ro m th e m o the r a nd a Y f rom th e

“

f a th e r a nd a re white eyed -
.

S U M M AR Y OF PR I N CI P LE S OF H ER EDI TY .

1 Th e fi rst l a w or p ri nciple st a tes th a t th e ge nes for

.

p a re nt a l ch a ra cte r p a i rs seg reg a te i nth e hyb rid

-
.

2 Th e sec on
. d p ri nciple g oes a step fu rthe r a nd st a tes
th a t e a ch p a i r of ch a ra cte rs ma y seg reg a te i ndepe nde ntly
o f othe r p a i rs .

3 Th e thi rd p ri n
. ciple is th a t i ndepe nde nt seg reg a ti on
is l imited to p a i rs wh ose ge nes a re i n sep a ra te ch rom o
s ome p a i rs
-
.

4 Th e f o u rth p ri n
. ciple is th a t ch a ra cte rs wh ose g enes
a re i nth e s a me p a i r O f ch rom os omes will be li n ked i e ,
. .
,

te nd m ore or less st rongly to rem a i nt ogethe r i nh e redity .

 P R IN CI P L E S OF H E R E DIT Y 379

WHITE

FI G . 1 12 Sex-Link a ge
. .White-e ye d fem a le by red-e yed m a le .

Th e symb ols use d a re th e s a m e a s i n th e l a s t figure . T h e re s ul ts

a re so m ewh a t diff eren t a l th o ugh t h e p ri ncipl es i nv olved a re identi ca l.

The fi rst gen era t i on fe m a le s a re red-e ye d b e ca use t h e y rece ive a

red gen e f ro m t h e i r f a t h er. T h e m a l es a re whi te-e ye d b eca u se t h e y

re ce ive a whi te gen e f ro m th e m o t h er a n d on l y a n ina ct iv e Y - ch ro

mosome from th e f a t h er Th e res u l ts i nthe se cond genera t i on a re

.

e xpl a i ne d i n th e s a m e wa y .

M orga n, Stu rteva nt, Mu ller, Bridges, M echa n i sm of M en deli a n

Heredi ty .
3 80 LI N KAG E P H E N OM E N A

5 Th e fifth p inciple is th a t the c ssi n

. r ro -
g ove r or ex
ch a nge O f c orresp ond ing p a rts o f h om ol og ous ch rom o
somes c a uses th e b re a ks i nli n k a ge of ch a ra cte rs
.

6 Th e six th p rinciple a sse rts th a t by c omp a ri ng th e

.

pe rce nt a g es of c rossi ng ove r i t is p ossible to sh ow th a t

-

th e ge nes a re a rra nged i nth e ch rom os omes inlinea r orde r .

7 S i nce sex i n h ighe r a nim a ls is d ete rmi ned by th e

.

ch rom os omes ch a ra cte rs wh o se ge nes a re l oc a ted in th e

,

sex ch rom os omes S h ow sex li nk a ge .

 C HAPTER XX X I

THE C HA R A CT E R E XPR E S S I ON OF GE N E S

T H E D I FFER E N TI A L GE N ES FA CTOR S Ina p reced

OR .

in g ch a pte r i t w a s S h o w n th a t n o ch a r a cte r is eve r th e

result O f th e a ctivity of on e si n gle ge ne or f a ct or a l one .

( P a ge N eve rth eless it is t rue th a t wh ethe r a p a r

ti cu l a r ch a ra cte r of a pl a nt or a nim a l a pp ea rs or n
. o t o fte n

depe nds on th e a ctivity O f one or a rel a ti v el y few genes

'

which ma y th erefore be c a lled diff erenti a l genes or fa c

, ,

t ors All th e ex a mples which h ave bee n used thus fa r

.

i n this b o ok h a ve depe nded fo r thei r fina l exp ressi on on

s ome one f a ct or The re a l S itu a ti onma y p ossibly be m a de
.

cle a re r by the rel a ti ons existi ng i n th e vineg a r fly i nre

spect to eye c ol or Ab o u t twe nty fi v e di ff e re nt modi fi ca
-
.
-

ti ons O f eye c ol or i n th is fly h a ve a l re a dy bee n studied

-
.

M a ny genes o f c ou rse i nte ra c t to p ro duce th e na tu ra l

, ,

wild type with red eye Th e o the r c ol or s a re due to a

.

cha n ge i n s ome si n gle ge ne for e a ch bu t not a lw a ys th e

s a me ge ne or eve none i nthe s a me ch rom os ome .

T H E R ELAT I VE P OS I TI O N O F G E N E S — I t h a s a l re a dy
been S h ow n th a t th e two di ff e re nti a l genes resp onsible
for a p a i r of c ont ras ti ng ch a ra c te rs a re c a rried i n th e
s a me ch rom os ome p a i r th e one i n one ch rom os ome a nd
-
,

the othe r i nits m a te a nd so S h ow seg reg a ti on M ore ove r

,
.
,

the genes a re a rra nged i nline a r orde r so th a t c orresp ond

in g ge n es li e a t ex a ctly th e Sa me r el a tive p o siti on s i n thei r

respective ch ro m o s omes Th is h oweve r f a ils to expl a i n

.
, ,

fully th e he redit a ry beh a vi or o f m a ny ch a ra cte rs a s for , ,

ex a mple eye c ol or i n vi neg a r flies A red eyed fly (Fig

,
-
.
-
.

1 1 3 ) c rossed to eith e r a W h ite eyed or a che rry eyed fly - -

S h o ws th e ordi n a ry seg reg a ti oni nth e sec on d hyb rid gen

3 81
3 82 C HA R A C T E R E X P R E S S I ON OF GENES
era ti onint o nd white or red a nd che rry a s the c a se
red a
,

ma y be A white eyed fly c rossed to a ch e rry eyed fly

.
- -

simil a rly gives white a nd ch e rry inthe sec ond ge ne ra ti on .

S ince a ny two of these ge nes (red ch e rry white a nd a ls o , ,

e osi n) for eye c ol or ma y p a i r with one a nothe r they must

-

w
hi te f ema l e Ch e rr y ml a e Ch e r r y f emal e Be d ma l e

I
Re d f e ma l e

I I

r
Ch e r r
x
y ml ’
a
I
o

Chbr r y ma l ’ Red ma l e
g

F1 0 . 1 13M ul tipl e All e l o m o rphs C h err y is a m o difi ca ti on of

. .

th e sa m e re d ge n e whi ch wh e n m o difi e d i n a n o t h er wa y p ro du ce s

whi te eyes I t is a lso knownto u nde rgo s till a no th er mu ta tio nwhi ch

.

give s ri se to eos ineye s T he dia gra m sh o ws the be h a vi o r o f the sex

.

ch ro m o so m es i nwhi ch th e se ge n e s a re co nta i ne d i n th e two cro s se s

whi te fe ma le by ch erry m a le a nd ch erry fem a l e by re d m a l e Th ey .

sh o ul d be comp a re d wi t h the cro ss be tw ee na whi te fe m a l e a nd a re d

m a le in the l a s t ch a p te r From a nex ami na tio n O f th ese di a gr a ms
.

i t will be O bse rve d th a t a n y two o f th e se ge n e s will p a ir wi t h o n e

an o t h e r be ca u se th e y a l l occu py th e s a m e re l a t iv e p o si ti o n i n th e i r

re spe ct iv e ch ro m o so m e s S u ch a sys te m O f se v era l ge n

. e s a bl e to

p a i r with one a nother cons ti tu tes a sys te m of m ul tip le a ll el om orphs .

 M UTATI ON S O F G E N E S 3 83

b e l oc a ted i nth e sa me rel a tive p ositi on, but o f c o u rse

'

a ll

an omosome ca n h a ve but on
'

y p a rti cu l a r ch r e o f them .

Th u s we h a ve i nsuch a c a se a system inste a d of a p a i r o f

genes a ll due to diffe rent ch a nges O f the s a me ge ne
, .

M UTATI O N S O F G E N ES If th e wild red— eyed vi neg a r .

fly is a ctu al ly the origi na l type it is cle a r th a t a ll o the r

eye c ol ors h a ve bee n de rived f rom it by a ch a nge or
-

mu ta ti onOf s ome one or m ore of th e ge nes c oncerned i n

the p roducti on o f red In the m ut a ti ons which h a ve .

give nrise to wh ite ch e rry a nd e osi n we h a ve see n th a t

, ,

the mut a ti on occu rred a t the s a me l ocus or p o siti on

i n th e ch ro m os ome O the r eye c ol ors e g ve rmili on
.
-
,
. .
, ,

h a ve a rise n by mut a t i on of o the r ge nes i n th e s a me

ch rom os ome p a i r Whe n white eyed m a les a nd v er
-
.
-

—
mili oneyed fem a les a re c rossed th e F m a les a re 1

ve rmili on like th e m o the r a nd the F fem a les a re red

, , 1 ,

bec a use they receive the norm a l red gene c orresp ond i ng
to white from th e m othe r a n d the norm a l ge ne c orre
sp on di ng to ve rmili on fro m th e f a the r An i nspecti on .
.

o f the a cc omp a n yi ng figu re (1 1 4 ) will S h ow th a t t hey a re

hyb rid for b oth ge nes white a nd ve rmi li on and the ref ore , ,

h a ve red eyes bec a use the norm a l ge nes a re b o th d omi

,

na nt M u ta ti ons of genes a lw a ys p recede a nd a re i nf a ct

.

the ca u se of mu ta ti ons i n cha ra cters Of pl a nts a nd

an im a ls .

T H E GA M ET I C R ATI OS O F HY B R I D S — The re a de r .

a l re a dy kn ows th a t th e hyb rid betwee n p a re n ts di ffe ri ng

i na si ngle p a i r o f ge nes a nd ch a ra cte rs p roduces j ust two
ki nds of g a metes a nd fu rth e r th a t the hyb rid betwee n
, ,

p a re nts di fi eri ng i nrespec t to two p a i rs of ge nes p roduces

fou r kinds of g a metes i n equ a l numbe r p rovided the .
,

ge nes i n questi on a re i n di ffe re nt p a i rs o f ch rom o s omes

an d hence not li nked I t is e a sy to go a step fu rth e r.

an d ge ne ra lize this I f p a re nts 1 a nd 2 di ffe r i nrespect

.

to th e i n depe nde nt ge nes a a nd a a nd b a nd b th e hyb rid ’ ’

,

h a s the ge no typ e o u bb a nd ca np roduce equ a l numbe rs

’ ’
,

O f g a metes a b a b a b a b N ow supp ose a thi rd p a i r

’ ’ ’ ’
.
, ,
,

,
3 84 C HA R A CT E R E X P R E S S I ON OF GENES
of ge nes 0 a nd c to be inv o lved Th e a b g a met es ca n
’
.

the nbe eithe r a bc or a bc ; i e the re a re twice a s m a ny ’

. .
,

kind s o f a b g a metes a s before Th e s a me th ing is t ru e .

fo r th e o th e r ki nds Thus e a ch a dditi ona l p a i r o f ch a ra c

.

te rs a nd ge nes for which a pl a nt or a nim a l is hyb ri d '

d oubles th e numbe r o f diff e re nt ki nds O f g a metes it ca n

Ve r mi l i o n f ema l e Pa r e nts Wh i t e ma l e

d
Re d - e y d f e ma l Ve r mi l i n ma ]
/
e Th e F hy br i d s o
1 '

\1 h
‘
e F
2
h b r i
‘
d s y
/

l
Ve rmi l i o n/f ema l e Re d ! sh a l e ma l g
k

FI G 1 14
. .
—
Re d-Ey e d Hyb rids p rodu ced by cro ssi ng whi te-e yed
an d v ermi l i o np a re nts T he F1 m a l e is v ermil i o nb eca se he re ce ive s
. u
his singl e X - ch ro m o so me f ro m hi s m o ther T he F1 fem a le is re d-e y e d .

be ca use sh e receives a nX -chrom o so me wi th the no rm a l re d f a cto r V

f ro m the fa th er a nd a no th er wi th the norm a l red fa cto r W f ro m
the m ot h e r T he re sul ts o f i n
. terb ree ding the F1 m a les a nd fe ma l es
is sh o wn i n th e low er ro w o f the di a g ra m .

p rod uce Or i n ge ne ra l te rms a n o rg a nism hyb rid for

.
, ,

np a i rs o f i ndepe nde nt ge nes a nd ch a ra cte rs ca np roduce

2 ki n
“
ds of g a metes .
 T HE N U MB E R OF G E N O T Y PE S 3 85

TH E N U M BE R G E N OT Y PE S I N T H E S E CO N D G E N ERA
OF

TI O N is c onst a n t for a ny p a rticul a r type o f m a ting a nd

depends of c o u rse on the n
, u mber o f ki n
, ds O f ga metes
wh ich th e fi rst ge ne ra ti on hyb rid ca n p ro duce .

He re only b a ck c rosses a nd i nte rb reed ing ca n be con

-

si dered .Th e fi rst is so simple a s b a rely to need me nti on ,

f or if n ki nds O f pu re g a metes a re m a ted to on e ki n d of

S pe rm (or egg ) (a n d a pu re recessive ca n f orm only on e

ki nd ) they w ill of c ou rse give rise to ndi ffe re nt c ombi na

ti ons O f ge nes or ge notyp es a nd a l l will o ccu r i n equ a l
numbe rs In i nte rb reeding the m a tte r i s not quite so
.

S imple bu t is e a sily u n de rst ood in the f oll owi ng wa y :

‘

P a re nts a X a p roduc e th e hyb rid a a a nd it p roduces

’ ’

g a metes a a nd a which c ombine i nth e sec ond ge ne ra ti on

’

geno typ e c ombina ti ons l a a ; 2a a ; l a a i e 3 ge notypes ’ ’ ’

,
. .
,
.

I f a sec ond ge ne a nd ch a ra cte r di ffe rence b b is a dded -

’

ge no type a a ma y bec ome a a b b a a bb or a a b b i e is ,

’
,
’ ’
,
. .
,

multipli ed by 3 Th e s a me thi ng is t rue for th e a a a nd

‘
’
.

a o ge n o typ es he n ce a ddi ng a noth e r ge ne a nd ch a ra c te r

’ ’
,

d iffe re nce i n the hyb rid multiplies the number of F

genotypes by 3 S t a ted ge ne ra lly a hyb rid i nv olvi ng n
.
,

p a i rs of i ndepe nde nt ch a ra cte rs a nd ge nes f orms 3 ge no 11

types i n th e s ec ond hyb rid ge ne ra ti on by i nte rb reedi ng .

T H E P H E N OTY P E S OF T H E S E CO N D GE N ER AT I O N Two .

i ndepe nde nt p a i rs o f ge nes a ss ort i n the reducti on divi

si ons of the hyb rid S O a s to give f o u r ki nds o f g a metes
an d these u nite by ch a nce to p roduce f o u r ph eno types i n
th e sec ond hyb rid ge ne ra ti on I f th e ge nes a re li n
,

. ked
two phe n o types ma y be missi n g a lt ogethe r or whe n ,

c rossi ng ove r occu rs a ppe a r i n gre a tly reduced numbe rs

-
,

i n c omp a ris on with ra ti o s expected i n i ndepe nde nt

a ss ortme n t I t is now p o ssible to c onside r a sec ond r e a s on
.

for v a ri a ti onf rom th e usu a l ra ti o s o f i ndepe nde nt a ss ort

ment The ra ti os discussed i n p revi ous p a ra g ra phs a re
n
.

th ose wh ich o ccu r wh e n a ch a ra cte r di ff e re nce depe ds

on a si n le di ffe re nti a l ge ne bu t the re a re m a ny ch a ra c
g ,

ter di ffe re n ces which depe nd onm ore th a none diff e re nti a l
 C HA R A C T E R E XP R E S S ION OF GEN E S
ge ne. I t is to be emph a sized th a t the ch rom o s ome
mech a nism dist ributes th ese ge nes i nthe usu a l wa y a nd
gives rise to exa ctly th e s a me numbe r a nd ki nds of ge no
typ es a s i n o the r ch a ra c te rs bu t th a t th e numbe r a nd
,

kind of ph e no types v a ry with th e wa y i n wh ich the

p a rticul a r ge nes exp ress themselves Th e f oll owi ng p a ra
.

g ra phs ill ust ra te c omm on t ypes O f inte ra cti on of ge nes

CO M P LE M E N TAR Y G E N E S a re such th a t th e p rese nce O f
b o th is necess a ry to p ro duce th e ch a ra cte r The re a re .

two s orts of wh ite c orne a ch o f wh ich h a s on e o f the co m

p l emen ta ry ge nes necess a ry to p roduce red c o rn Whe n .

they a re c rossed th e hyb rid is red a nd the F c onsist sof 2

9 red to 7 wh ite This resul t ca nbe re a dily u nde rst ood

.

by c onside ri ng th e c ombi na ti ons o f ge nes i n the sec ond

ge ne ra ti on If we supp ose th a t th e one white p a re nt
.

c a rrie s the ge nes A a nd b a nd th e o the r c a rries the ge nes

a an d B b o th d omi na nts bei ng necess a ry to p roduce red
,

c ol or we ca na cc o u nt for a ll th e f a cts Th e h yb rid the n

,
.

h a s th e ge n o typ e Aa B b (si n ce one p a re nt is supp o sed to

be AAbb a nd th e o the r a a B B ) Th e hyb rid p rod uces
.

fo u r ki nds o f g a metes AB Ab a B a b i nequ a l numbe rs

, , ,
.

Th ese rec ombine a cc ordi ng to La w 2 to give 9 AB ; 3 a B ;

3 Ab ; 1 a b (see Pu n nett squ a re C h a p XXIX ) Th e
,
. .

9 AB a re of c ou rse red ; th e 3 Ab a re wh ite like on e g ra nd

p a re nt ; th e 3 a B a re wh ite like th e o the r g ra ndp a re nt ;
an d th e 1 a b i s a ls o wh ite Th e seve na re white bec a use
.

th ey l a ck eith e r one o r b oth of th e ge nes nec essa ry to

m a ke red .

S UPP LE M EN TAR Y G E N ES This na me is give nto ge nes

.

wh ich ch a nge o r m odify the ch a ra cte rs p roduced by o the r

ge nes bu t do no t p roduce any visible e ff ect of themselves .

For ex a mple i f p u re red c orn is p olli na ted by a ce rt a i n

,

ki nd o f wh ite com a pu rple hyb rid is p roduced The .

wh ite c a rries a m odifyi ng or suppleme nt a ry ge ne which

ch anges red to pu rple bu t p rod uces no c ol or i n th e
,

a bse n ce of eithe r of the ge nes for red I f this pu rple .

hyb rid is self p olli na ted i t gives pu rple white a nd red

-
, ,
 I NH I B I T O R Y GE N E S 3 87

phe no typ es i n th e sec ond hyb rid ge ne ra ti on In this .

p a rticul a r c ross the red c ornh a s the ge no type AA B B cc

( A a n d B bei ng the ge n es which p roduces red ) a n d th e
white one ma y h a ve the ge no type AA bb C O (i n wh ich
C is the ge ne wh ich ch a nges red to p u rple ) The F .
1

hyb rid is AA B b Cc a nd p roduce s g a metes AB C Abc , ,

Ab C Abc which c ombine to p roduce F O f 9 AB C ; 3 AB c ;

, 2

3 Ab C ; 1 Abc Th e 9 AB C a re p u rple ; the 3 AB c a re red ;

.

the 3 Ab C a nd th e 1 Abc a re wh ite bec a use they l a ck one

O f the ge n es for red .

I N H I B I TOR Y G E N ES a re ge nes which p reve nt th e u su a l

a cti on of o th e r ge n es I f a ce rt a i n ki nd of wh ite c orn
.

is c rossed with red c orn th e hyb rid is wh ite notwi th ,

st a nding th a t i t receives b o th red ge nes from the red

p a re nt Th e re a re seve ra l w a ys i n which this may h a p
.

p e n O f which on e O f th e simple r will be expl a i ned Th e ..

red p a re n t h a s th e geno type AAB Bi i a nd th e wh ite

p a rent AAb I wh ich gives the hyb rid the c omp ositi on
o f AA B b I i . I t p roduces g a metes AB I AB i AbI Abi , , ,

an d they rec ombi ne to p ro duce a sec ond hyb rid gene ra

ti on of 9 AB ] ; 3 AB i ; 3 AM ; 1 Abi Al l th ose with I .

a re white bec a use this ge n e p reve nts the f orm a ti onof red ,

an d Abi is wh ite bec a use it l a cks B Th e re a re the re

.

f ore 1 3 white to 3 red i n the sec ond hyb rid ge ne ra ti on

o f this c ross Othe r c rosses ca n b e m a de with diffe re nt
‘

ki nds of wh ites but a ll c ont a ining I which will of c ou rse , ,

give still di ff e re nt phe notype ra ti os i n F 2.

C U M U LATI VE GE N E S a re th o se which p roduce di ff e re nt

i ntensities or a m ou nts o f the s a me ch a ra cte r a cc ord
i ng to h ow m a ny o f t hem a re p resent A ve ry simple
.

c a se of this wa s desc ribed i n th e f o u r o cl ock whe re th e ’

re d p a re n
”
t h a s two red ge nes R R a nd th e white h a s
“
‘

two c orresp on ding ge nes rr Th e hyb rid h a s only one

.

red ge n e (B r ) a nd I S only a b out h a lf a s red p ink )

a s th e red p a re n t A ce rt a i n red g ra i ned wh e a t c rossed
.
-

with a wh ite gives a h a lf red hyb rid Wh e n th e-

.
‘

hyb rid is self p olli na ted it p roduces a sec ond ge ne ra tion

-
3 88 C H AR A C T E R E XP R E S S I O N OF GE NES
Of pl a nts in wh ich the re a re S i x S h a des o f red besid es
wh ite Wi th o u t g o ing int o det a il i t ma y be s a id th a t
.

this result depe nds on th e f a c t th a t the red g ra ndp a re nt

h a s th ree p a i rs o f ge nes f o r red a nd a l l a re p ra c tica lly
eq uiv a le nt They rec ombi ne i nth e sec ond ge ne ra ti onso
.

th a t s ome pl a nts h a ve 6 red ge nes s ome 5 s ome 4 s ome, , ,

3 s ome 2 s ome 1 a n
, , d s ome n
,
one th e reby givi ng rise to
,

a s ma n y c ol or typ es The wa y this w orks o u t i s sh ow n

.

i nth e a cc omp a n ying P u nnett squ a re (Fig .

RED REd Re D Re d rED rEd re D

FI G 1 1 5
. . Th e F, of a Cert a i nWh ea t C ross wh ere the red p a re n t
(RREED D) diff ers f ro m th e whi te o ne (rreedd) i n respe ct to th ree
se p a ra te a nd i nd e pe nd ent p a i rs o f genes , a ll o f whi ch a re ca p a bl e
o f p rod u ci n g so m e red co l o r T he F, is a b o u t h a lf as re d a s the red
.

p a re nt Wh enthis h a lf red hyb rid (RrEeDd) is se lf- p o lli na te d i t p ro

.
-

du ces the geno type co mbi na ti o ns sh o wn i n the Pu nne tt sq u a re .

N o te th a t si nce e a ch o f th e ge n es R , E , a nd D a re e quiv a l e n t and

th e i r e ff e cts cu mul a t ive , th ere wil l be 1 pl a nt wi th 6 d ose s o f red ,

6 pl a n ts wi th 5 d o se s , 1 5 wi th 4 d o se s , 20 w i th 3 d oses , 1 5 wi th 2
d ose s, 6 wi th 1 d o se , a nd 1 wi th no red, i e whi te . . .
 S UM M AR Y 3 89

S U M M AR Y . 1 The numbe r a nd ki nds o f g a metes

.

p roduced by a hyb rid depe nds onth e numbe r o f di ffe re n

ti a l ge nes i nv olved i n the c ross a nd on whethe r a ny of
th em a re li nked i nthe s a me ch rom os ome p a irs -
.

2 For a n
.
y p a r ticul a r type o f m a ti n g ( e b a ck c ross
.i .
-

or i n te rb reeding ) the ki nds a nd ra ti os o f c ombi na ti ons

of ge n es ge no types ) wh ich occu r i n the sec ond
hyb rid ge ne ra ti on depends on th e ra ti o a nd ki nds of
g a metes f ormed by the fi rst ge ne ra ti on hyb rid .

3 The ra ti o of phe n
. o typ es i nthe sec on d hyb rid gene ra
ti on depends on the wa y i n wh ich th e i ndividu a l ge nes
exp ress themselves a s well a s onth e ra ti o of g a metes p ro
du ced by th e fi rst ge ne ra ti on hyb rid .

4 Th e c omp a ra tively simple ra ti os to b e expected

'

a cc ordi n g to th e l a ws of M e ndel 1 st a nd 2md La ws )

a re ve ry o fte ng re a tly m o dified by (1 ) li n k a ge a nd c ross
i ng ove r a n
-
,
d (2) th e wa y i n which the ge nes exp ress
themselves .
 C HAPTER XXX II
PLANT B R E E DI NG

T H E HI S TOR Y P LA N T B R EEDI N G — The re is no

or .

rec ord o f whe n men fi rst beg a n to a ttempt to imp ro ve

pl a nts Wh e reve r th e expl ore r g oes a m ong s a v a ge a nd

.

p r imitive pe oples he fi nds th em usi ng pl a nts o f s ome sort

a s f ood . H e o fte nfi nd s s ome pl a nts cultiv a ted a nd they
a re likely to be bette r th a n the w ild on
‘
es I t seems .

p rob a ble th a t c onsci ous ch o ice o f bette r pl a nts for seed

i nm a ny c a ses c a me s oona fte r ma nre a lized th e effi cie ncy
o f cultiv a ti on C e rt a i n it is th a t m o st imp ort a nt c rop
.
,

pl a nts h a d not only bee nimp roved by th e time a t wh ich

th e re i s th e e a rliest hist oric a l t ra ce of them but a ls o th a t
,

th is imp roveme nt h a d h a ppe ned so l ong bef ore th a t it ,

is a m a tte r o f difficulty eve nto t ra ce thei r origi na l s ou rce .

For ex a mple it is only a few ye a rs a go th a t wh e a t wa s

,

f o u nd g rowi ng i n th e na tu ra l wild st a te i n P a lesti ne ,

a lth o ugh i t h a d bee ni n cultiv a ti on bef o re the d a w n o f

h ist ory .

D u ri ng th e l a st 200 ye a rs th e a ttempts to b reed bette r

pl a nts h a ve bee n nume rou s a nd s ometime s st riki ngly
successful I t is h oweve r only withi n th e l a st twe nty
.
, ,

fi v e ye a rs th a t sufficie nt k nowledge o f th e l a ws o f he red

i ty h a s bee n a v a i l a ble to p u t pl a nt b reedi ng on a
-

scie ntific b a sis th a t e na bles it to p roceed c onfide ntly a nd

ra pidly t ow a rd its desi red e n ds E ve ry civilized g ove rn
.

me nt now empl oys nume ro us pl a nt b reed e rs S c ores .

a re w o rki n g i n expe rime nt st a ti ons sch ools a nd com

, ,

merci a l seed est a blishme nts .

T H E ECO N O M I C I M P ORTA N CE or P LA N T B REED I N G In .

th e U nited S t a tes th e N a ti ona l G ove rnme nt th ro ugh the ,

D ep a rtme nt o f Agricul tu re a nd the v a ri ous st a tes th rough

,

3 90
 P R OB LE MS OF PLAN T B R E E DING 3 91

t heir expe rime nt st a ti ons a nd c olleges of a g ricultu re ex '

pe nd l a rge sums of m oney on pl a nt b reedi ng These .

f a cts a re well k nowna nd s ometimes form a b a sis for u n

f ou nded c riticisms bec a use c omp a ra tively few pe ople
re a lize the v a lue i n d o ll a r s a n d ce nts o f this w ork As .

a m a tte r of f a ct the expe n se is a me re tri fi e c omp a red

to the retu rn s A few ye a rs a go M r Luthe r Bu rb a nk
. .

pl a ced on th e m a rket a new p o t a to I t wa s estim a ted .

th a t wheni t h a d c ome int o gene ra l use it h a d i ncrea sed

‘

the va lu e of th e a n nu a l p o t a t o c rop by a n a m ou nt
g rea te r th a nthe e nti re c ost of ru nni ng the U S D ep a rt . .

me nt of Ag ricultu re for one ye a r J ust before th e G re a t .

Wa r th e M a rquis whe a t p roduce d by D r Ch a rles E , . .

S a u nde rs D omi ni onC e re a list h a d c ome i nt o ge ne ra l cul

, ,

ti v a ti on i n the a r e a s o f C a n a da a n d the D a k o t a regi on

o f th e U n ited S ta tes which a re suited to i t I t is a ve ry .

i nte resting f a ct th a t th e i nc re a se i nyield due to its i nt ro

ducti oni n these a re a s is a lm ost ex a ctly th e s a me a s the
a m ou n t o f wh e a t which the two c ou nt ries c ould sp a re for
exp ort to ou r a llies Wh a t th e l a ck of this a dditi ona l
.

whe a t w ould h a ve me a nt the re a der ca n re a dily j udge

,

for himself Its me re m onet a ry v a lue to th e p roduce r

.

wa s m ore th a n a h u nd red milli on d oll a rs Whe n Pro .

f essor R obe rts O f K a ns a s is ol a ted his K a n red wh e a t h e

a dded to th e a n nu a l i nc ome of eve ry K a ns a s owne r o f a

1 60 a c re wh e a t field e n
-
o ugh m on ey to buy a new a ut o
m obile a nd th a t with o ut th e l ea st i nc re a se O f l a b or
. .

T H E P R OBLE M S O F P LA N T B R EED I N G a re a s v a ri o us a s
the desires O f men I nc re a se O f yield is one of th e m o st
.

freque nt I t is o ften desi ra ble to exte nd th e ra nge ove r

.

which a c rop ca nbe g row n a nd so it bec omes necess a ry

to p roduce a n ew v a riety th a t will th rive u n de r the new
c onditi ons o f g re a te r c old or highe r tempe ra tu re d rie r or ,

wette r s o ils m ore or less su nshi ne etc In m a ny c rops

, ,
.

a pples p ot a t oes ) u n
,
if ormity O f size a nd a ppe a ra nce
is highly desi ra ble In ora nges a nd g ra pes seedless
.
,

va rieties a re v a lu a ble a s well a s th ose high i nsug a r con

3 92 P LAN T B R E E DING

te nt In c orn th e pe rce nt a ge o f sug a r st a rch p ro tein

.
, , ,

an d Oil a re a ll imp ort a nt c onside ra ti ons for p a rticul a r

uses a nd the b reede r s trives to p rod uce a ra ce wi th the
m ost suit a ble c omp ositi onfor e a ch Th e f oll owi ng p a ra .

g ra phs will fu rnish c onc rete ex a mples of h ow the b reede r

h a s s olved s ome o f these p roblems :
T H E M ET H ODS OF P LA N T B R EED I N G n a tu ra ll y v a ry

a cc o rd ing to the p roblem but ma y be g ro uped u nde r two

,

ge ne ra l h e a ds selec ti on a nd hy bri di z a ti on Th e stude nt

, .

will u nde rst a nd f rom his p revi ous studies th a t s ome kinds
o f pl a n ts a re na tu ra lly self p olli na ted a nd so exist i n -

pu re li nes re a dy for th e b reede r to ide ntify a nd select ,

while o the rs a re na tu ra lly c ross p olli na ted so th a t p op ul a -

ti ons o f them c onsist of m a ny a nd u su a lly a ll p ossible

ge no types Some pl a nts a re self ste rile a nd so c a nnot be
.
-

self p ollina ted the reby i nc re a si ng the l a b or of b reedi ng

-
,

them Fu rthe rm ore it freque ntly h a ppe ns th a t i t is

.
,

necess a ry to c ross two species or v a rieties i n orde r to

b ri ng t ogethe r th e d esi red c ombi na ti on o f ch a ra cte rs ,

self p oll ina te or i nte rb reed the hyb rids a nd ch oose f rom
-
,

a m on g th e rec ombi na ti ons i n F a nd l a te r ge ne ra ti ons 2

the ones th a t a re desi red .

P UR E L I N E S ELE CT I O N Wh e a t is na tu ra lly self p ol li

.
-

na ted bu t is o cca si ona lly c rossed i n th e field A field of .

m ost c omme rci a l v a rie ties is likely to c onsist of a me re

me ch a ni ca l mixtu re o f nume ro us diff e re nt p u re li nes One .

meth od o f imp rovi ng it is th e refore to m a ke a ve ry c a re

, ,

ful a nd c omplete study o f th e g rowi ng g ra i ni norde r to

ch oo se th e best pl a n ts These a re th e nh a rvested sep a
.

ra tel y Th e seed s of e a ch pl a nt will ordi na rily be pu re

.

an d c onstitute a pu re li ne Th ese pu re seeds a re s owed

.

the nex t ye a r i nth e testi ng pl ot o ne ro w to e a ch col lec ,

ti onO f seed for c omp a ris ona nd selecti ono f th e best pu re

,

li nes Th e next ye a r s a mples f rom th e best pu re li nes

.

a re s owed i nl a rge r pl o ts for m ore a ccu ra te c omp a ris on .

It requ i res a b o u t five ye a rs o f testi ng i n this wa y to

m a ke a fina l selecti on I t wa s i nthis wa y th a t P ro fess or
.
 I M P R OVING INDIAN CO R N 393

R obe rts is ol a ted th e K a nred wh e a t which h a s p roved to ,

be so supe ri or to the whe a t s p revi ously g row n i n the

Pl a ins regi on
—
.

I M PR OVI N G I N D A N COR N
I N O c rop is m ore imp or
.

t a nt i n the U nited S t a tes th a n m a ize or I ndi a n c orn or ,

h a s received m ore a tte nti ona t th e h a nds O f th e b reede r ,

b oth a m a teu r a nd p rofessi ona l I t is believed th a t this .

pl a nt p rob a bly origina ted i nM exic o Ina ny eve nt the re .

,

we re a l re a dy m a ny v a rieties in cultiv a ti on a m ong th e

Ame ric a nI ndi a ns wh e nw hi te menfi rst m a de its a cqu a i nt
an ce These v a rietie s h a ve si nce the n bee n i nc re a sed
.

an d imp roved u ntil now the re a re excelle nt v a rieties

a d a p t ed to a lm o st eve ry pu rp ose a n d to a lm o st eve ry s ort
of clim a te or s oil withi nthe ra n ge O f th e pl a nt Cornis .

na tu ra lly wi nd p olli na ted a nd th e ref ore usu a lly c rossed

. , ,
.

Acc ordi ngly m ost com pl a nts a re hyb rids Th ey ca nbe

, .

reduced to a pu re c on diti onby a rtifici a l self p olli na ti on -

,

but th is Imp a i rs th e vig or o f th e pl a nt so g re a tly a s to

m a ke th e meth od imp ra c tic a ble excep t for speci a l p ur
p os es Inp ra ctice the b reede r to begi nh is w ork eithe r
.
, , ,

selects di rectly a numbe r O f g oo d pl a nt s f rom th e v a riety

h e wish es to imp rove or else h e fi rst c rosses two ,

p resum a bly suit a ble v a rieties a nd selects d esi ra ble

pl a nts from th ei r offsp ri ng Wh icheve r wa y th e fi rst .

selecti onis m a de th e ch ose npl a nts a re is ol a ted i na sep a

ra te b reedi n g pl ot so a s to be p ro tected from p olle n of
,

o th er v a ri e ti es an d a ll owed to p olli na te one a no the r

’

.
,

Th e best pl a nts a re a g a i nch ose na nd th e p rocess repe a ted .

Eve ntu a lly a rel a tively u nif orm ra ce is secu red but it
, ,

is no t of c o u rse p u re Wh a t h a s a ctu a lly h appe ned by

, , .

this meth o d of selecti onis to m a ke th e ra ce rel a tively pu re

for ce rt a i n ch a ra cte rs (wh a teve r ma y be desi red ) a nd ,

hyb rid for m a ny o the rs i n orde r to m a i nt a in th e

necess a ry vig or O f the pl a nt .

B REEDI N G C OTT O N The m o st v a lu a ble p a rts of th e

.
-

c ott on pl a nt a re l ong silky h a i rs a tt a ched to th e seeds

”
i nside th e c ott on b oll or seed p od These h a i rs v a ry
“
.
3 94 PLAN T B R E E D ING
innumbe r, le ngth nd st re ngth I t i s desi ra ble for m ost
, a .

p u rp oses a ut om obile ti re c as ings ) to h a ve them a s

l ong st rong a nd fle xible a s p ossible Cott on i n na tu re
, , .

re a dily c rosses so th a t pu re seed ca nbe p ro duced o n

,
ly by
gu a rdi ng a g a inst c ross p oll i na ti on Th e meth od O f a o
-
.

c omplish i ng this is b o th u nique a nd e ff ective Ince rt a i n .

is ol a ted v a lleys o f the a rid s outhweste rn U nited S t a tes

c o tt onca nbe g row nby i rrig a ti ona nd ca ne a sily be kep t
pu re p rovided only one v a riety is g row n i n e a ch v a lley
G ro we rs i na v a lley a g ree ona ce rt a inst ra i na n
.

d th e np re
ve nt by l aw th e pl a nti ng of a ny o th e r ki nd S electi on .

for imp roved seed i s the nm a de i nis ol a ted b reedi n g pl a ces

u nt il a suit a ble v a riety h a s bee nsecu red a nd the nonly ,

p u re seed is s ow ni nth e v a lley I t rem a i ns pu re me rely

.

bec a use it h a s no ch a nce to c ross .

S EEDLE S S ORA N GES illust ra te a s ort o f pl a nt b reedi n g

wh ich ca nonly be p ra cticed with p e re nni a l pl a nts whi ch
ca nbe multiplied i na sexu a l w a ys by g ra fts cu tt in gs etc , , .

All th e W a sh i ngt on na vel ora nges i n Ca lif orni a a re de

sce nded f rom two t rees origi na lly b ro ugh t f rom B ra zil .

J us t a s th ese fi rst seedless ora nges origi na ted f rom se ed ed

v a rieties by mut a ti o n so they i n thei r tu rn h a ve p ro
,

d uced m a ny new st ra i ns f ro m bud sp orts By a bud .

sp ort is me a nt th a t a mut a ti on occu rs i n th e cells of a

bud a nd th e bud i ntu rng rows ou t i nto a b ra nch sh ow
,

in g th is mu t a ti o n .W he neve r a n e w a n d bette r mut a ti o n

i s obse rved i t is mul tiplied by t a ki ng bud s f rom the
mut a ted b ra nch to g ra ft i nt o seedl i ngs As ide from mu t a .

ti ons pl a nts O f this s ort do not ch a nge (excep t i nresp onse

,

to the e n vi ronme nt ) a nd no spe ci a l c a re is necess a ry to

keep them p u re S ome t rees a re m ore p roduc tive th a n
.

o th e rs h oweve r a nd a re no w used by the up to d a te

, ,
- -

nu rse rym a na s th e s o u rce o f his buds .

D I S EA S E R ES I S TA N CE h a s bee n f ou nd to be a he rit a ble

ch a ra cte r i n m a ny species o f pl a nts T wo illust ra ti ons.

will sh ow h ow this ca nbe m a de useful i nc omb a t ing l oss

f rom dise a se .
 FR O S T R E S I S TAN CE 3 95

A few ye a rs a go th e c a bb a ges g rownexte nsively i nWis

c ons inwe re ne a rly wiped ou t by a s oil fu ngus P rofess or .

J ones obse rved i n a field th a t one pl a nt seemed not to

h a ve been i nj u red He s a ved seed fro m it a nd pl a nted
.

them i n the i nfected s oi l a nd a g a i ns a ved seed from the

m ost resist a nt In this wa y h e b red a ra ce Su ffi ci entl y
.

r esist a n t to m a ke it p rofit a ble to g row c a bb a ges a g a i ni n

th a t regi on .

Wh e n th e s outh e rn w a te rmel ons we re a tt a cked by a

wil t fu ngus D r W A Ort on of th e U S D ep a rtment
,
. . . . .

o f Ag ricultu re c rossed them with th e cit ron which i s

, ,

u np a l a t a ble but is resist a nt to this wilt Th e hyb rid .

O ffsp rin g we re self p ollina ted a nd from a m ong this p rog

-

en y h e eve n tu a lly selected new c ombi na ti ons which a re

resist a n t to wilt a nd t a ste like w a te rmel ons a nd i n
, ,

a dditi on st a n d shipp ing bette r th a n th e origi na l mel ons .

FR OS T R S S TA N CE
E I — S ome ye a rs a go a n u nu su a lly
c old wi nte r killed l a rge numbe rs of ora nge t rees i n th e
Fl orid a orch a rds Th e l oss wa s so seve re th a t g rowe rs
!

hesit a ted to invest a g a i n The p roblem wa s eve ntu a lly .

s olved b y D r W T S wi ngle O f th e U S D ep a rtme nt of

. . . . . .

Agricultu re wh o c rossed the cultiv a ted ora nges with th e

h a rdy Ch inese wild species Cit rus t rifoli a t a S electi ons
from th e n
.
,

ume rous a nd v a ried rec ombina ti ons of F a nd 2

l a te r hyb rid ge ne ra ti ons h a ve yielded v a rieties not only

well a d a pted to st a nd th e c old i nth e origi na l cit rus belt
but eve n to exte nd i t ne a rly f ou r hu nd red mi les to the
north .

S U M M AR Y . Pl a nt b reeding c onsists i n
1 Is ol a ting a l re a dy existi ng pu re li nes i nself p olli na t
.
-

ing species .

2 S electi n
. g suit a ble hyb rids f rom p op ul a ti ons O f
na tu ra lly c rossed species a nd inte rb reedi ng th em u ntil
they form p u re or a pp roxim a tely pu re li nes .

3 Crossi n
. g species v a rieties or ra ces to rec omb ine th e
, ,

desired ch a ra cte rs from e a ch i nthe sec ond hyb rid ge ne ra

3 96 PL AN T B R E E DING
ti on from wh ich selecti ons ma y be m a de a nd pu rified by
,

fu rth e r inb reedi ng or self p olli na ti on

-
.

4 Th e selecti on a n
. d p rop a g a ti on by a s exu a l meth od s
O f desi ra ble b ra n ches which h a ve a rise n by veget a tive
mu t a ti ons or bud sp orts .

Th e ch a ra cte rs with wh ich th e pl a nt b reede r w orks a re

'

mut a ti on s a nd no t me re m odific a ti ons or a re rec omb ina

, ,

ti ons of mut a nt ch a ra cte rs Th ey ma y be eithe r Ol d bu t

.

u nno ticed ch a ra cte rs or new ones p roduced by rece nt

mut a ti ons si nce a ll he rit a ble ch a ra cte rs h a ve origi na ted
,

a t s ome time by mu ta ti on .
 C HAPTER XXX III

ANI M AL B R E E DING

R E PR OD U CTI O N HIN A A —
I GH ER Whe re a s the
N IM LS

pl a nt b reede r h a s a c onside ra ble ra nge of rep roductive

p rocesses a t his disp os a l a nd ca n o fte n ch oose th e one
best suited to his pu rp oses th e a nim a l b reede r is limited
,

to a S i ngle one T h e re a re two sexes i na ll h igh e r a n

. im a ls ,

so th a t eve ry in dividu a l h a s two p a rent s a nd is the refore

likely to be hyb rid for a numb er O f ge nes a nd thei r cor
resp on di ng ch a ra cte rs I f th e b reede r m a kes a c ross b e
.

twee n the p u rest O f ra ces h e c a nno t expect th e hyb rids

to be a ll a like a s h e c ould if de a li ng wi th self p olli n
, a ted -

pl a nts bec a use th e p a re nts a re p rob a bly neve r ex a ctly

,

a like i na ll th ei r ge n es .

ON I N TER B R EED I N G S ince the re a re a lw a ys two sexes

'

th e b reede r of h ighe r a nim a ls c a nno t self fe rtilize h is -

fi rst gene ra ti on hyb rids bu t must m a te th em with one

an o the r. In s ome species of a nim a ls i t is p ossible to
m a te b ro ther a nd siste r with out ill e ffects while ino th e rs ,

th is d oes not a ppe a r to be p ra ctic a ble Ina ny eve nt the .

b reede r c omes a s ne a r self fe rtiliz a ti ona s th e na tu re of the

-

species m a kes a dvis a ble i norde r to c ome j ust a s ne a r to

”
b reedi ng a pu re li ne a s p ossible In a ctu a l p ra ctice .

this neve r p ro duces a n a bs olutely pu re ra ce no m a tte r

h ow l ong the pedig ree On the o the r h a nd ra ces a re
.
,

O f te np u re for i n dividu a l ch a ra cte rs which a re rep roduced

g e n e ra ti on a fte r ge ne ra ti o n with r em a rk a ble fidelity .

ON T H E E FFE CT S OF SE LE CT I O N I t must be cle a r from

.

th e re a der s p rese nt knowledge th a t rec ombi na ti ons (p a ge

’

3 6 ) a re c onti nu a lly o ccu rri ng i nrespect to s ome ch a ra c

te rs i na nim a ls S ome o f th ese ma y be sligh tly bette r a nd
.

397
3 98 ANI M AL B R EE D ING
s ome w orse th a n th e p a re nt a l c ombi na ti ons For this .

re a s on c on ti nu o us selecti on of th e best off sp ri ng of e a ch

ge ne ra ti on to bec ome th e p a re nts O f the nex t is a lw a ys
necess a ry to secu re a ny imp roveme nt or eve n to a v oid
dege ne ra t i on o f the s tock I t a ppe a rs to be t rue th a t
.

an im a ls a re especi a lly rich insu p p lementa ry genes (p a ge

3 86 ) wh ich m odify th e qu a n tit a tive exte nt O f a
ch a ra cte r S i nce these a re nume rous the re is excelle nt
.

opp ortu n ity for c onti nued selecti onto isol a te ge notyp es
with m ore a nd m o re O f these ge nes i nb o th membe rs of
the ch rom os ome p a i rs a nd so i nc re as e the qu a nti ta tiv e
-

exp ressi onof th e ch a ra cte r O f c ou rse selecti onis usu a lly

. .

eq u a lly e ff ective in the opp osite di rection .

I N H ER ITAN CE O F CO AT CO LOR I N G U I N EA P I GS Th ese

‘
-
.

sm a ll a nim a ls a re much used for expe rime nt a l w ork i n

bi ol ogy ona cc o u nt O f the s a tisf a ct ory wa y i nwhich th ey
e nd u re c a p tivity T wo ch a ra c te rs of th ei r h a i r will se rve
.

a s illust ra ti on s (Fig 1 1 6 ) o f th e f a ct th a t m a ny t ra its

.

i n a nim a ls beh a ve i n st rict a cc ord a n ce with M e ndel s ’

l a ws o f seg reg a ti on a nd i ndepe nde nt a ss ortme nt A .

sm oo th h a i red bl a ck gui ne a pig c rossed to a ro ugh h a i red

- -

white one gives hyb rid off sp ri ng with rough bl a ck h a ir .

Whe n th ese hyb rids a re inte rb red they yield a sec ond
hyb rid ge ne ra ti on of 9 Bl a ck R ough ; 3 Bl a ck sm oo th ;
3 white R o ugh ; 1 whi te sm oo th H a i r c ol or in m a ny
.

o th e r a n im a ls beh a ves i n esse nti a lly th e s a me wa y ,

a lth o ugh the re a re O fte n m ore p a i rs of ge n es i nv olv ed

an d so th e results a re less simple .

I N H ER ITA N CE O F H O ODED P ATT ER N I N AT S R — In a n

exte nsive se ri es of expe rime nts with ra ts P rofess or C a stle
sh owed th a t a p a tte rn of th e c oa t c ol or ca n re a dily be
m odified by selecti on I f a pu re c ol ored ra ce is c rossed with
.

on e h a vi n g a c ol ored p a tch or h ood ove r th e he a d a nd

sh oulde rs the hyb rid off sp ri ng a re a ll fully c ol ored (Fig .

Wh e nthe hyb rid s a re i nte rb red the sec o nd hyb rid

ge ne ra ti on sh ows 3 c ol ored to 1 h oo ded Th e h ooded .

on es a re of c o u rse recessive a nd pu re for th e h ooded

, ,
 L E THAL G E N E S I N Y ELL OW MI CE 3 99

ge ne Th ey do not h oweve r b re ed wh olly t rue to type

”
.
, , ,

for thei r desce nd a nts S h ow v a ri o us deg rees O f h oodi ng .

By selecti onfor seve ra l ge ne ra ti ons i t is p ossible to b reed

a n e a rly white or ne a rly full c ol ored ra ce This result is -
.

now gene rally believed to be d ue to th e selecti ono f m odi

fy i ng f a ct ors wh ich exte nd the size o f the h ood .

FI G 1 1 6
. . R e sul t s b etweentwo va rieties of gui nea pig
of a C ro ss -

differi ng inth e two u ni t ch a ra cters col or a nd lengt h O f fur Fi g a

-
, . .
,

a co l ore d a n d sh ort h a ir e d gu i n
-
e a pig Fi g b a n a l bi no a nd l ong
-
. .
,

h a ire d gui nea pig T h e P y ou ng w ere co l ore d a nd sh ort h a ired lik e

-
.
I
-

th e p a re n t sh ow ni nFi g a Fi g 0 a co l ore d a nd l ong h a ire d gui nea

. . .
,
-

pig one of th e new F v a rieties Fi g d a na lbi no a nd sh ort h a ire d

, 2 . .
,
-

gu i nea pig th e oth er new F v a riety T h e two oth er F va rieties

-
, 2 . 2

were like th e gra ndp a rents Fi g a a nd b ,

. .

Ca st l e H eredi ty a n
,
d Eu gen i cs Ha rva rd Press . .

LET H A L G E N ES I N Y EL LO W M I CE The re is a t ype of .

yell ow c ol or ch a ra cte ristic of ce rt a i n st ra i ns of hyb rid

mice Wh e nthese hyb rid yell ow mice a re i nte rb red they
.

sh ould p roduce 1 p u re nonyell ow bl a ck ) 2 hyb rid -

yell ow ; 1 pu re yell ow As a m a tte r of f a ct only two .

yell ows (b oth hyb rid ) a re b orn for eve ry 1 nonyell ow -

,

but th e missi ng pu re yell ows ca nbe sh ow nto h a ve p er

i sh ed bef ore bi rth S uch ge nes a s th is a re c a lled le tha ls
.

an d a re of c onside ra ble imp ort a nce C a re h a s to be t a ke n .

4 00 AN I M AL B R E E DING
by th e a nim a l b reede r whe nselecti ng for ce rt a i ndesired
c ombi na ti ons of ch a ra cte rs not t o i nclude a ls o leth a l
genes or othe rs which u nduly diminish the fe rtility or
st a mi na of the ra ce Th e re is a ctu a l rec ord of a su p e ri or
.

ra ce O f c a ttle which pe rished f or th is re a s on .

I N H ERI TA N CE OF W I N T ER EGG P RODU CT I O N BY F OW LS

— I t h a s been f ou nd th a t th e a b ility to p roduce m a ny
.

FI G 1 1 7A In h eri ta nce o f a recessive p a tte rnof whi te sp otting

”
. .

seenin h ooded ra ts
“
T h e p a ren ts a t th e l ef t a re a h o m o zyg o us
.

”
h ooded m other a nd a h etero zygous Irish fa th er (bla ck wi th whi te
“

be ll y ) Anenti re li tte r o f th ei r y ou ng is sh owna t the righ t Fou r

. .

a re h om ozyg ous h oo d e d lik e th e m o th er, fi v e a re h e te rozygou s l ik e

t he f a th er N o te flu ctu a ti on
. s inb oth cla s se s Su ch flu ctu a ti ons a re
.

f ou nd to be inp a rt h eri ta ble .

Cas tl e, Heredi l y a nd Eugen i cs Ha rv a rd Press

. .

FI G 1 1 7 B
. A Gra d ed Seri es o f Pa tte rns Use d i n Gra di ng the
.

C o a ts i na se l ec ti o ne xpe ri m ent B y co n
. ti nu a l sel e cti ono f the p ro g
en y o f the o ff sp ri n g of the cro ss sh o wn a b ove a ny o f th ese types
,

o f p a t te rn
s ca nbe p ro du ce d This res u l t is now supp ose d to be d ue
.

to t h e m o difi ca ti o no f t h e eff ec ts o f o ne m a i n f a cto r fo r h o odi ng ”

by num ero us ot h er fa cto rs whi ch h a ve the eff ect Of m o difyi ng the
e x te nt o f t h e whi te a n d col o re d a rea s .

C as tle .
 W IN TE R E GG P R O D U C TI O N 401

eggs du ri ng th e wi nte r m onth s whe n eggs b ri ng f a ncy

p rices is inhe rited a nd th a t i t depe nds on two p a i rs O f
genes This is a n inst ructive c a se bec a use i t sh ows th a t
.

the a bility to l a y eggs inwi nte r is not a na cqui red ch a r

a cte r int roduced by th e ki nd of c a re t a ke n of th e fl ock .

One Of these ge n e s is in th e sex ch rom os ome of which

Ma l e Pa r e nt . F e ma l e Pa r ent

Ma l e F1 y
h br i d F mle a e PI

P
2
Ma l e F2 Ma l e F2 F ml
e a e P
2
F e ma l e

FI G . 1 18Inh erita nce of Winter Egg Pro d u ction Through th e

.

M a l e O ne f a ctor for wi nte r egg p ro du ct i on is l o ca te d in th e sex

.

ch ro m os o m e of f owls T h e m a l e h a s two se x ch ro m o so m es a n
. d th e
fem a le one Th e di a gra m sh ows t h a t th e pu reb re d m a le tra nsmi ts
.

hi gh egg l a yi ng a bili ty to a ll his d a ugh ters ev en wh en m a te d t o a

-

fem a le wi th ou t this fa ctor (E) Ha lf th e F2 fem a le s inh eri t high

.

a b i li t y a nd h a lf do not All th e m a l e s i n
. h eri t th e gene (E ) b u t only
h a lf of th em a re p ure for i t (EE) a nd t heref ore sui ta ble sires .

the m a le f owl ha s two a ctive or Z ch rom os omes a nd th e

fem a le only one a ctive or Z ch rom os ome a nd one i na ctive
or W ch rom os ome (Fig He nce eve ry d a ughte r re
.

cei v es a na ctive Z f rom th e f a th e r but only a n i na ctive

W from th e m othe r S i nce this ge ne for high egg l a yi ng
.
-

is i n the Z ch rom os ome h e ns i nhe rit th is ch a ra cte ristic

,

from th e fa the r a nd not f rom the m othe r M a nifestly .

4 02 ANI M AL B R E E DI NG
th e f a th e r neve r a cqui red th e a bili ty to l a y eggs a t a ny
se a s on a lth ough h e ca n t ra nsmi t to his d a ughte rs a
m a rked a b i lity t o do so .
I N H ER ITA N CE I N DA I RY CATT LE Th e a bili ty to p ro
n
.

d uce l a rge qu a ntities o f m ilk a nd butte r f a t depe ds on

the p rope r c orrel a ti on o f m a ny st ructu res a nd fu ncti ons
an d is the ref ore no t g o ve rned by a S i n gle p a i r of ge nes .

I t is neve rtheless he rit a ble a s is cle a rly sh ow n by th e

,

supe ri ority O f ce rt a i n b reeds e g J e rseys In b reed i ng

,
. .
, .

for high milk p rod ucti onthe b reede r must keep a l l these
t h in gs i nmi nd M e rely to e nume ra te s ome O f them will
.

suffice to sh ow th e c omplexity o f th e p roblem T O tu rn .

f ood i n t o milk a s ec onomic a lly a s p ossible ce rt a inly re

q ui res a g ood set Of teeth a nd a g ood digestive system
, ,

an d a type o f met a b olism wh ich tu rns f oods int o milk in

ste a d O f i nt o muscle or fa t Th a t m a ny of th e ge nes
.

which c ont rol these th i ngs ca nb e a ssembled in one ra ce

is sh ow n by th e f a c t th a t th e b reede r h a s a ctu a lly p ro
du ced ch a mpi on c ows H olstei ns ) which p rod uce
p o u nds o f milk p er a nnum i npl a ce of th e 3 000
4 000 p ou n ds p roduced by the sc rub cow He re a g a i n it .

sh ould be emph a sized th a t high milk p roducti on is i n

h eri ted f ro m th e m a le p a re n t a s well a s f rom th e fem a le .

I t sh ould be no ted h oweve r th a t th e a dv a nt a ges O f a

, ,

pu re b red si re a re not so g re a t a s i nth e ex a mple o f th e

f owl j ust give n bec a use th e re the ge ne for wi nte r egg
,

p roducti on is sex li nked-

.

IN H E R I TA N C O
E F S P E ED I N H OR S ES — S i nce speed is
equ a lly ch a ra cte ristic of b oth sexes a nd s ince f a s t h orses ,

a re usu a lly desce n ded f ro m p a re nts b o th o f wh om we re

f a st th e i nh e rit a nce o f t rotti ng a bili ty h a s o fte n bee n
,

cited by th e a dv oc a tes of th e i nhe ri t a nce o f a cqui red

ch a ra cte rs a s a c a se i n p o i nt I f a ll c a ses we re O f this
.

s ort it w o uld be ex tremely di fficult to give a na ccept a ble

an swe r to th is a sse rti on for it is a nu nd o ubted f a c t th a t
,

f as t h orses o f te n beget f a st O flsp ri ng The c rux o f the .

q uesti o n is wh ethe r it i s the eflect of tra i n i ng which

 VALU E OF I M P R OVE D B R E E D S 403

is inhe rited or me rely the a bi li ty to be tra i ned Inview .

O f the l a st two ex a mples the l a tte r seems much th e m ore

,

p rob a ble Th e effects of c onti nued selective b reedi ng a re

.
,

the refore to be a sc ribed to th e g ra du a l a ccumul a ti on of

“

nume rou s genes a ll O f wh ich c ont ri bute s omethi ng t ow a rd

speed .

T H E VA L U E O F I M P R OV ED B REED S i s ou t of a ll p rop or
ti onto the c ost of p roduci ng them I t c osts little or no.

m ore to feed a th oroughb red th a n a sc rub cow but ,

the qu a ntity O f milk a nd butte r received ma y i n ex

’

t reme c a ses be ten times a s g re a t I t is ce rt a i nly th e

.

duty the ref ore O f i ndividu a ls a nd g ove rnme nts to p ro

, ,

m ote the b reedi ng o f sup e ri or st ock not only of d a i ry ,

c a ttle a nd f owls but of m a ny othe r d omestic a ted a nim a ls ;

f or wh a t is t rue of milk butte r a nd eggs is a ls o t rue i n
, ,

gene ra l te rms for beef , p ork a nd mutt on w ool h ides a nd

, , , ,

silk.

S U M M AR Y Th e p ra ctic a l a nim a l b reede r h a s ca r

.

ried m a n y species a nd v a rieties to a high st a te of p erfec

ti on Wh e re the knowledge of th e ge nes i nv olved is
.

su fli ci en t th ey a ppe a r to beh a ve in th e s a me ma n
,

ner a s th ose with which we a re a cqu a int ed i n th e

expe rime nt g a rde n a nd b reeding p en Th e meth ods .

empl oyed a re eithe r c ontinued selecti on in a v a riety

t ow a rd a nide a l or selecti ona fte r hyb ridiz a ti on The re
, .

su l ts o f selecti oni nb o th c a ses a ppe a r to be best expl a i ned

on the a ssumpti on of th e g ra du a l a ssembli n g of m a ny

f a v ora ble genes i n a m ore Or less p u re ge notyp e The .

ge nes a re p rob a bly so nume rous th a t no pu re ra ce h a s

yet re a ched a p oi nt wh e re it is p u re f or eve ry p a i r of
ge nes Conseque ntly it ma y be c onfidently h oped th a t
.
,

c ontinued b reeding will imp rove eve nthe best Of b reeds .

SECTI ON 1 0

EVOLU T I ON
 C H APT ER XXX I V

E VID E N CE S O F E VOLUTI O N

EV OLUTI O N For m a ny ce ntu ries it wa s

D E F I N ED .

gene ra lly believed th a t eve ry species of a nim a l a nd pl a nt

wa s a Speci a l a ct o f th e Cre a t or, a nd l ong a go wa s pl a ced
ini ts p rese nt pe rfected st a te a nd i n th e s a me l oc a lity it
now occupies Acc ording to th is V iew a sp ecies is u n
.

ch a nge a ble Th e fish is f oreve r a d a p ted for a life i nth e

.

w a te r a m ole for tu nneli ng i nth e e a rth a nd a bi rd for a n

, ,

existence i nth e a i r This V iew of th e org a nic w orld f a ils

.

to t a ke i n t o a cc ou nt a l a rge b ody of f a cts wh ich is now

th ough t to runc o u nte r to such a belief a nd wh ich is ex ,

pl a i ned by th e m ode rnbi ol ogist onth e p ri nciple of evol u

ti on Acc ordi ng to th is d oct ri ne a species a mid ch a ngi ng
.
,

c onditi ons i s not fixed but on th e c ont ra ry is u nde r

,

g oi ng a lte ra ti ons fro m ge ne ra ti on to ge ne ra ti on Th e .

ch a nges ge ne ra lly a re ve ry sl ow a nd a lm ost impe rceptible

du ri ng a b rief pe ri od bu t wh e re th e rem a i ns of c o u nt less
,

ge ne ra ti ons h a v e bee n p rese rved i n f o ssil f orm du ri ng

milli ons of ye a rs th e t ra nsf orm a ti on is cle a r a nd u n
,

mist a k a ble I t is a ls o t rue th a t th e fu rth e r b a ck th e his

.

t ory of a species is t ra ced th e simple r th e a ncest ra l type

bec omes In o th e r w ords if th e ge ol ogic a l rec ord we re
.
,

c om plete it w o uld be f ou nd th a t e a ch o f th e th ou s a nds

of species of a n im a ls a nd pl a nts is th e desce nd a nt of one
simple f orm of life or a t m ost o f a ve ry few Ev oluti on
, ,
.
,

the ref ore Is th e de riv a ti onof c omplex high a nd speci a l

, , ,

i z ed species from S imple l ow a nd ge n e ra lized a ncest ors

, ,
.

S OUR CE S OF E V I D E N CE Th is gra du a l ch a nge or ev ol u

.

ti onof livi ng th i ngs is to the t ra i ned bi ol ogist a nest a b

li sh ed f a ct T o h im th e evide nce i nsupp ort O f such a b e
.

4 07
408 E VID E N CE S O F E V O LUTI O N

li ef is a s c onvinci ng a s th a t u nde rlyi ng the ph e nome na

O f g ra vit a ti ona n d ch emi c a l a ffin ity A s i ndic a ted i nt h e .

foreg oi ng p a ra g ra ph th e study o f p a le ontol ogy or f o ss i l

, ,

rem a i n s i n th e c rust o f th e e a rth is a m os t imp ort a nt ,

s ou rce O f i nf orm a ti on bu t c ont ra ry to th e p opul a r

,

belief i t is by no me a ns th e on
-
ly one I na dditi on to .

th e p roo f fu rnished by 1 ) P a le ont ol ogy a we a l th o f evi ,

de nce h a s bee nsupplied by studies i n (2) Cl a ssific a ti on ,

( )
3 G e o g ra ph ic a l D ist ribu ti o n (4 ) C o mp a ra tive An at ,

o my , ( )
5 E mb r y o l o gy ( 6 ) A r tifici a l ,
S elec ti o n a nd ( 7 ) ,

G e netics Th ese t opics will now be c onside red i n s om e

.

det a il .

E V I D E N CE FR O M P A L EON T OLOGY I t is O fte nsupp o sed .

th a t if a l l th e fossils i n th e e a rth s c rust c ould be ’

studied a f a i rly c omp lete h ist ory of life c ould be w ritte n

,
.

S uch h o weve r is no t the c a se

, ,
Al l th e a nim a ls a nd .

pl a nts f rom th e fi rst a ppe a ra nce of livi ng th ings d ow nto

pe rh a ps fi fty milli on ye a rs a go h a ve dis a ppe a red com
‘

p l e tel y I f th.ey e ve r we re p r ese rved a s f o ssils these h a ve ,

bee nde st royed by h e a t w a te r or oth e r a ge ncies Wh a t , ,

.

exist s a t th e p rese nt da y is only a sm a ll f ra cti on O f

th e wh ole N eve rth eless this rem na nt is highly illumi na t
.
,

i ng a nd fu rn ish es ex a ctly th o se f a cts wh ich a re to be ex

‘
p eCted a s a resul t O f ev oluti on .

S ome o f th e best k now na n d m ost st riki n g ex a mples o f

a l on g c onti nued ev olu ti ona ry h ist ory h a ve bee n dis
c ove red i nth e U nited S t a tes App roxim a tely th ree mil .

li onye a rs a go i t is estim a t ed th e clim a te of th e c ou nt ry

, ,

wa s simil a r to th a t i nAf ric a a t th e p rese nt time T ropi .

ca l veget a ti onex te n ded a s fa r northw a rd a s th e Ca nad i a n

b orde r c roc odiles i nh a bi ted th e exte nsive m a rsh es wh ile
, ,

h e rd s of c a mels eleph a nts a nd rh i noce roses roa med th e

, ,

l a nd As time p a ssed th e tempe ra tu re gra du a lly l owe red

. ,

the c ou nt ry bec a me d rye r a nd l a rge fo res ted d ist ricts were ,

ch a n ged i nt o t reeless pl a i ns Amid th ese sh ifti ng con .

di ti on s ma n y a nim a ls a nd pl a nts bec a m e exti nct or mi

gra ted i nt o m ore f a v ora ble regi ons O the rs u nde rwe nt .
 HI S T O R Y OF T H E H OR S E 4 09

ch a nge s a nd bei ng a d a pted to th e new c ondi ti ons p er

, ,

sisted M ost f ortu na tely the S kelet ons of m a ny O f these

.
,

an im a ls h a ve bee np rese rved a nd f rom them it h a s bee n ,

p ossible to t ra ce the ev oluti on of v a ri ou s species from

th e a ncest ra l f orm d ow n to th e p rese nt da y .

H I S TOR Y O F T H E H OR S E Th e fi rst u nd oubted mem .

be rs o f th e h orse f a mily wh ich fu rnish es one of th e best

,

k now nex a mples of a l ong ev oluti ona ry h ist ory a ppe a red ,

i n N orth Ame ric a a s sm a ll a nim a ls sc a rcely l a rge r th a n ,

a g ood sized h ouse ca t (Fig

-
I na dditi onto seve ra l.

FI G . 1 19 . R es tora ti onof th e f ou r toe d h ors e ; b a s e d ona m ou nted

-

sk el eton ,
sixteen inch es high , i nth e Ameri ca n M s eum of N a tu ra l u
His tory .
(Af ter C R K nigh t )
. . .

o the r rem a rk a ble ch a ra cte rs th ey h a d f ou r h oofed t oes ,

an d a rudime nt a ry thumb i ne a ch f ore f oot while th e hi nd ,

foot l a cked th e gre a t toe a nd th e fifth digit wa s sm a ll

an d rudime nt a ry D u ri ng th e next milli onye a rs or p er
.
,

h a ps a l onge r pe ri od th ese a ncest ra l f orms g ra du a lly b e

,

c a me l a rge r a nd devel oped or ev olved i nt o (a ) a rel a tively

sl ow goi ng h e a vy type c onfi ned to th e f orests a nd i nt o
-
,

( b ) a g ra ceful ligh t —
b odied,
swift pl a i ns typ e , ,
.

Th e fi rst form bec a me exti nct a fte r u nde rg oi ng ce rt a i n

ch anges while th e pl a i ns a nim a l bec a me di ffe re nti a ted
,
41 0 E VID E N CE S O F E V OLUTI O N

i nt o seve ra l distinc t species S ome O f th ese rem a i ned i n

.

N orth Am e ric a a nd bec a me exti nc t du ri ng th e Ice Age .

O th e rs mig ra ted i nto S o u th Ame ric a wh e re th ey di ed o u t

in ve ry rece nt t imes S till o th e rs t ra velled i nto th e Ol d
.

W orld a nd bec a me th e a nces tors O f th e m ode rn b reed s .

Th e re we re no h orses i nth e N ew W orld whe nColumbus

m a d e h is v oy a ges .

Th rough ou t th is e ntire pe ri od th e b ody wa s u nde rgoi ng

m a ny ch a nges Am ong th e m o st Obvi o us we re th ose
.

a ff ecting the feet Th e mi ddle digit g ra du a lly i nc re a sed

.

in siz e w ,
h ile th e o th e rs bec a me rud ime nt a ry or dis

FI G . 1 20 . Ev ol u ti o n o f t h e Ha nd of th e Ho rse .

a ppe a red c ompletely (Fig Th e resul t is th a t th e

.

m ode rnh orse t ra vels o nth e tip O f a si ngle (th e middle )

digi t th e h oo f rep rese nti ng th e na il wh ile the knee
, ,

is a c tu a lly the w rist or a nkle Th e teeth a ls o bec a me .

m odified a nd a s a result of these a nd oth e r ch a nges th e

,

ge ne ra lized fi ve t oed a ncest or ev olved i nt o one o f th e

-

m ost highly speci a lized o f m ode rn a nim a ls .

E V OLUTI O N O F CA M ELS Alth ough th ese a nim al s a re

.

no t i ncluded inth e list o f na tive N orth Am e ric a nspecies ,

th e f a mily a c tu a lly p a ssed th rough th e gr e a te r p a rt o f

i ts devel opme n t on th is c onti ne nt Th e e a rli est know n .
 CLA S S IFI C ATI O N 41 1

a ncest ors we re sm a ll a nim a ls a b ou t th e size of th e a ve r

,

a ge dog with f o u r h oo fed t o es on e a ch f oo t

,
This type .

bec a me devel oped i nt o m a ny species s ome of which , ,

like th e g ra ceful a nd swift g a zelle c a mels a nd th e exces

si v el y l ong necked gi ra ffe c a mels bec a me exti nct Oth e rs
-
, .

c onti nued to exist a nd mig ra ti ng i nt o S o uth Ame ric a

, ,

ev olved i nt o th e ll a m a gu a na c o a lp a c a a nd rel a ted

, , ,

species S ti l l oth e rs c rossed th e a ncie nt l a nd b ridge int o

.

th e Old W orld a nd bec a me t ra nsformed i nt o th e m ode rn

,

typ e Th ose rem a i ni ng inN orth Ame ric a died ou t l ong

.

a go .

In a dditi on to n i ncre a se i n b odily Si ze a nd seve ra l

a

o th e r ch a ra cte ristic fe a tu res O f th e gro up th e digits dis ,

a ppe a red with th e excep ti ono f th e th i rd a n d f ou rth th e ,

h oo fs bec a me reduced to sm a ll c onic a l na il s a nd th e .

weigh t O f th e b ody wa s supp orted by fleshy p a ds All of .

th ese ev oluti ona ry ch a nges a re evide nced by a we a lth o f

fossil rem a i ns a nd th e hi st ory o f th e c a mel is a cc ord
,

i ngly a s well est a blished a s th a t o f th e h orse .

I t is imp ort a nt to be a r i n mi nd th a t ge ol ogists a re

’

a gr eed th a t th e gre a t ro ck m a sses i n wh ich th ese f o ssil

h orses c a mels a nd oth e r a nim a ls a re e nt ombed form a n

, ,

u nb roke nse ries Th e l owest st ra t a were fi rst f ormed a nd

.
,

th e upp erm ost a re th e m ost rece nt Th e p a le ont ol ogist .

is equ a lly p ositive th a t th e f ossils i n th e oldest dep osits

a re th e m ost p rimitiv e a n d ge ne ra lized a nd th a t th e ,

v a ri ou s species a re m ore a nd m ore c omplex a nd speci a lized

a s th e l a test st ra t a a re a pp roa ch ed T o th e ope nmi nded .
-

stude nt of bi ol o gy th ese f a cts p rese nt u nmist a k a ble evi

de nce of th e re a lity of ev oluti on .

C SS
L A I F I CAT I ON — Th e h ist ory o f th e c a mel f a mily
.

illust ra tes two imp ort a nt f a c ts Inth e fi rst pl a ce th e one.

an d two h umped c a mels th e ll a m a gu a na c o a nd a lp a c a

-
, , ,

a re th e desce n d a nts O f a c omm on a ncest or S ec ondly .

,

th ese a re disti nct sp ecies a t th e p rese nt time owi ng to ,

th e dyi ng ou t O f th e i ntermedi a te st a ges or c onnecting

li nks T o exp ress i t gra ph ic a lly th e f a mily h ist ory of
.
,
41 2 E VID E N CE S OF E V OLUTI O N

th e c a mel i s a t ree wh o se t ru nk is th e a ncest ra l st ock ,

wh ile th e b ra nch es rep rese nt th e ev oluti onof th e rem ote

an cest or i nt o di ffe re nt species Th e gre a te r p a rt o f this
.

tree i s de a d th e tip s of th e b ra n
.

,
ches wh ich c orresp ond ,

to the m ode rn species bei n g th e only livi ng p orti on

, .

O th e r f a mil ies o f a nim a ls a nd pl a nts wh ose a ncest ra l

t rees h a ve bee n c a refully f oll owed i nt o the e a rth a nd ,

th e f ra gme nts o f th ose which ye a rly a re b rough t to light ,

a ll illust ra te these s a me ge n e ra l p ri nciples .

In th e a ttemp t to cl a ssify th ese livi ng species it wa s

l ong a go disc ove red th a t resembl a nces a nd diffe re nces of
stru ctu res a re th e o n ly s a fe guide S h a pe size c ol or .
, , ,

an d o th e r sup e rfici a l ch a ra cte rs we re u sed by th e bi ol ogist

a c en tu ry or m ore a go but th ey we re fo u nd to be p ra cti
,

c a lly w orthless V a ri ous S ch emes o f g ra ph ic a lly rep re

.

Se nti ng rel a ti onsh ips we re a ls o a b a nd oned i nf a v or o f th e

t ree lik e b ra nch i ng pl a n such a s is used to illust ra te the
-
,

ge ne a l ogy or rel a ti onship of v a ri ous h um a nf a milies In .

o th e r w ord s th e e a rlie r scie ntists to wh om ev oluti on

,

wa s u n know nwe re d rive nto a d op t a system O f cl a ssi fica

ti ona pp roxim a ti ng th a t o f th e p rese nt da y Thei r w ork .
,

h oweve r wa s little m ore th a n a n i ntellectu a l exe rcise a

, ,

pige onh oli ng o f org a nisms for c onve nie nce only T o da y .
-

i t rem a i ns a c onve nie nt system yet a t th e s a me time ,

i t exp resses th e bl ood rel a ti onsh ips O f livi ng th i ngs a nd ,

is one O f th e st rong p roo fs th a t ev olu ti onh a s occu rred .

T H E M ODER N S Y S TE M O F C LA S S I F I CAT I O N Acc ordi ng .

to th e p rese nt system o f cl a s sific a ti on a species is defi n ed ,

a s a na ssembl a ge O f simil a r a n im a ls or pl a nts which do ,

not diffe r i nsize sh a p e c ol or e tc bey ond th e limits o f

, , ,
.
,

i ndividu a l v a ri a ti on Fu rth e rm ore th ese ch a ra cte rs a re

.
,

i nh e rited with bu t sligh t m od ifi c a ti on; a nd th e g roup is

no t u nited with a ny oth e r by c onnecti ng li nks N ow let .

th is defi n iti on be a pplied to th e g roup o f c a ts Am ong .

th e d omestic a ted a nim a ls th e re a re m a ny b reeds pe rsi a n , ,

m a ltese m a nx etc bu t th ese do no t c onst itu te disti nct

, ,
.
,

species si nce th e re a re m a ny i nte rmedi a tes

,
Th ey a re .
 S C I E NTI FI C NA M E S 41 3

o nly v a rieties sub species devel oped du ri ng domesti

or -

c a ti onfrom th e wild or P a ll a s ca t O f Af ric a ’

.

Th e li on tige r ocel ot pum a etc a re st ructu ra lly dif

, , , ,
.
,

ferent f rom e a ch o th e r a nd a re disti nct sp eci es a n d e a ch ,

i nh a bits a defi nite ge ogra ph ic a l a re a Fu rth e rm ore e a ch .

,

of these species a n d ne a rly a hu n ,

d red o the rs p ossesses ,

ce rt a i n a na t omic a l ch a ra cte rs i n c omm on a nd a cc ord ,

in gyl they a re u nited i n a l a rge r divisi on th e e n

g us .
, .

Othe r ca t like a nima ls bel ongi ng to o the r ge ne ra a re

-
, ,

g rouped with these i nt o a fa mi ly The ca t f a mily to .

,

gethe r with th ose of the d ogs h ye na s etc a re emb ra ced , ,

.
,

i n a n order This and seve ra l o the r orde rs of a nim a ls

.
,

a ll with h a i r c omp rise th e cla ss o f m a mm a ls

,
Th e sev .

e ra l cl a sses of b a ckb oned a nim a ls f orm th e p hy lu m O f

ve rteb ra tes And fi na lly th e v a ri ou s phyl a (S p onges
.
, , ,

m ollusks etc ) a re subdivisi ons of the a ni ma l ki ngdom

,
. .

Th e pl a nt ki ngd om is subdivided i n th e s a me m a nne r .

Eve ry a nim a l a n d eve ry pl a nt wh ich h a s bee ncl a ssified , ,

o ccupies a d efi n ite pl a ce If th e w ork is a ccu ra tely d one .

its p o siti oni ndic a tes its ne a rest rel a tives a nd th e b roa de r ,

fe a tu res a t le a st of its ev oluti ona ry h ist ory

, , .

S CI E N T I F I C N A M E S Th e qu esti onis o fte na sked why

.
,

is it necess a ry to use ja w b re a ki ng scie ntific na mes ? -

Th e a nswe r a ppe a rs whe n one t a lks a b ou t g oph e rs f or ,

ex a mple In th e So uthe rn S t a tes the w ord refe rs to

.

a t ort o ise i n C a lif orn,

i a to a p o uch ch eek ra t like a ni -
,
-

ma l i n Id a h o to a sp ecies of s na ke a nd i nM ont a na to
, ,

a g rou nd squi rrel V e rna cul a r or p op ul a r na mes a re

.

the ref ore c onfusi ng to sa y th e le a st Th e gre a t w ork o f , .

th e S wed ish scie ntist Li nnaeus wa s th e b ri ngi ng o f orde r , ,

ou t o f th is ch a o s by a d opti n g na mes f rom a de a d l a n

gu a ge eith e r La ti n or Greek a nd h e nce i nc a p a ble o f
'
d
, ,

ch a nge .

Ea ch species of a n im a l a nd pl a nt is give ntwo na mes ,

on e for th e ge n ti s th e o th e r f or th e species ,
Th e ge ne ric .

na me of a ll c a ts is Fe li s Th e specific na me of th e li on .

is leo of th e tige r ti gri s O f th e le op a rd p a rdu s etc Th e

, , , , ,
.
41 4 E VID E N CE S O F E V O LUTI O N

c omplete na m e O f the li onis Felis leo o f th e tige r Felis , ,

ti gris etc ,
na mes th a t a re fixed a n
.
-
d know n by the
e nti re scie ntific world Eve ry species ge nus f a mily a nd .
, , ,

h ighe r divisi on of b oth th e pl a nt a nd a nim a l ki ngd om

h a s its o wn p a rticul a r scie ntific na me .

CO PAR AT V A N AT O Y
M I E M — As i ndic a ted i n a f orme r
.

p a ra g ra ph evide nce f rom supe rfici a l resembl a nces b e

,

twee n th e v a ri ou s species o f a nim a ls a nd pl a nts i s no t

a s a fe guide i ncl a ssific a ti on Onsuch a b a sis th e wh a le .

an d fish w o uld be cl o se rel a tives Actu a lly th ey a re no .

,

m ore cl o sely rel a ted th a nma na nd a sh a rk Th e resem .

bl a nce is me rely one o f a na l o gy a simil a rity o f f orm , .

Wings a re a na l o g ous o rg a ns i nbirds a nd i nsects I nb oth .

g ro ups th e re is a like ness of fu ncti on bu t st ructu ra lly ,

th ey a re ve ry diffe re nt a nd only dist a ntly rel a ted .

On the othe r h a n d th e wi ng of a b a t a nd the a rm O f

,

ma n p rese nt no a na l ogies of eith e r f orm or fu ncti o n .

Th ey a re h oweve r fu nd a me nt a lly a l ike i n st ructu re

, ,

(Fig . B one for b one m uscle f or mu scle ne rve , ,

f or ne rve th ey a re a lm ost ide ntic a l th e diff e re nce

, ,

bei ng me rely th ose O f p rop orti on Exte ndi ng th e com .

p a ris on it is f ou nd th a t th e w i ngs O f bi rds th e p a ddles

, ,

O f wh a les th e diggi ng a rms o f m oles th e legs o f

, ,

h orses zeb ra s etc p rese nt wide di ff e re nces o f f orm

, ,
.
,

an d fu ncti on a n d yet a re st ruct u ra lly ne a rly a like

,
.

O th e r p a rts o f th e b ody illust ra te th e s a me f a ct Th is .

d eep se a ted ide ntity of st ructu re is te rmed h omology

-
,

an d is i nte rp reted a s a S ignO f bl o od rel a ti on sh ip J ust a s .

th e m ode rn o ne fingered h orse h a s bee nde rived f rom a

-

five fin -
g ered a n cest o r so it is believed a ll o f
,
th,
e b a ck ,

b oned a nim a ls o r ve rteb ra tes with th ei r v a ri o us typ es O f ,

limbs a re th e desce nd a nts o f a rem o te a ncest ra l species

,
.

R U DI M E N TA R Y OR V ES T I GIA L ORGA N S In th e se a rch .

for h om ol ogies a m ong th e v a ri o us species o f a n im a ls a nd

pl a nts o rg a ns h a ve c ome to ligh t especi a lly a m ong th e
, ,

high e r species wh ich a re of little or no u se In th e

,
.

m ode rnh orse for ex a mple two sm a ll sp li nt b ones o ccu r

, ,
 R UDI M E NTA R Y OR VE S TIGAL O R GAN S 41 5
‘

j ust bel ow the knee (Fig At the p rese nt time

.

th ey a re Of no service a nd rep rese nt th e l a st t ra ces of two

,

fi nge rs or t oes wh ich once we re p rovided with h oo fs a nd

we re fu ncti ona l .

Inma nthe re a re no t fewe r th a n 1 70 vestigi a l org a n s .

Th e a ppe ndix for ex a mple is a rem na nt ofte na t rouble

, , ,

s ome one whe re a s i n such a nim a ls a s th e ra bbit a nd

,

k a ng a roo it is of V it a l i mp ort a nce The re a re a ls o t ra ces

.

FI G 1 21
. . Hom o l og o us Org a ns : A wing
, of ba t ; B, wi ng of bird .

of a t a il with rudime nt a ry muscles muscles th a t once ,

m oved the e a rs a nd twitched the ski n a rudime nt a ry ,

th i rd eyelid such a s one fi nds well devel op ed i n the

,
-

bi rds a nd nume rou s o the r v esti ges a ssoci a ted wi th the

,

skelet a l muscul a r a nd oth e r systems

, ,
.

E x a mples O f this cl a ss o f org a ns i n o th e r a nim a ls a s

well a s i n m a ny pl a nts c ould be multiplied to a g re a t
le ngth Ineve ry i nst a nce thei r p rese nce ca nbe expl a i ned
.

on th e a ssump ti on th a t e a ch O f th e s e orga ns wa s fu nc
ti on a l i na rem o te a nces tor I ndeed th ey a re defi nitely
.
,
41 6 E VID E N C E S O F E V O LUTI O N

know nto h a ve bee nso i nm a ny inst a nces As the species .

ev olved th e org a n bec a me less imp ort a nt dwi ndled in

, ,

size a nd ultim a t ely ce a sed to fu ncti on

,
.

EV I DE N CE FR O M E M BR YO LOGY —In a f orme r ch a p te r .

(XX V I ) i t h a s bee n sh o w n th a t a l l o f th e m a n y celled -

an im a ls devel op from a si ngle fe rtilized egg cell wh o se -

divisi ons resul t i n a bl a stul a a nd still m ore c omplic a ted

g a st rul a Th is devel opme nt is f ro m a c omp a ra tively
.

si mple to a c omplex c onditi on This i s a l so th e h ist ory .

o f life on th e e a rth Th e a ncie nt simple species h a ve

.

ev olved i nt o th e m ore onless c omplex d esce nd a nts o f

m ode rn times Am oeb a V olv ox a nd Hyd ra resp ec
.
, , ,

ti vel y c orresp o n d to i nc re a si ngly c omplex steps i n th e

ev oluti on of th e a nim a l ki ngd om Th e c orresp ond i ng .

st a ges — egg bl a stul a a nd g a st rul a — a re i nc re a si ngly

, ,

c omplex s ta tes i n th e ev olu ti on or devel opme nt o f a n

i ndividu a l Th e y ou ng st a ges i n th e g rowth o f a h igh e r
.

species th u s resemble th e a dults of l owe r species S pe a k .

in g b r o a dly life h ist o ry,

p a ra llels o r r epe a ts ra ce h isto ry .

D EV ELO P M E N T O F V ERT EBR ATE S Th is p a ra llelism b e .

twee n life a nd ra ce h ist ories is fu rth e r illust ra ted i n a

s triki ng m a nne r by th e devel opme nt o f th e ve rteb ra t es .

Th e fish es f rogs reptiles bi rds a nd m a mm a l s a ll bel ong

, , ,

to this g re a t g ro up a n d a g ree i nth e p ossessi onO f a b a ck

,

b one f ou r limbs a d ors a l ne rv o us system a nd seve ra l

, , ,

oth e r imp ort a n t fe a tu res Th e resembl a nces a re p re .

su ma b l y due to th e f a c t th a t they a re th e desce n d a nts of

a c omm ona n cest or ; th e di ffe re nces a re owi ng to evol u
ti on a ry ch a n ges fitti ng e a ch species f or a defi nite m od e
,

o f life .

I nits devel o pme nt e a ch S pecies t ra vel s esse nti a lly th e

s a me p a th a nd for a l o ng pe ri od th e y o u ng ca nbe dis
,

ti n gu i sh ed o n ly by a n expe r t M o r e o ve r bef o r e defi .n ite ,

cl a sses a ssume th e fi na l a dul t a nd disti nguishi ng ch a ra c

te rs th ey a ll devel op gil ls i nth e th roa t regi o n I nte rna lly .

a s well a s exte rn a lly th ey be a r a st riki ng resembl a nce

,

to fish es I n th e fishes th ese org a n

. s pe rsist th rough ou t
 G E O G R A P HI CAL DI S T R I B UTI O N 41 7

life ; i nth e t oa d liz a rd robi n ra bbit a nd rel a ted species

, , , , ,

th is fi sh like c onditi ong ra du a lly dis a ppe a rs

-
.

F rom th ei r devel opme nt a s well a s from evide nce su p

,

plied by c omp a ra tive a na t omy a nd f ossil rem a i ns i t is ,

ve ry evide nt th a t m ode rnve rteb ra tes a re th e desce nd a nts

of p rimitive b a ckb oned gill be a ri ng a ncest ors wh o se -

ev oluti onh a s resulted i nth e th ous a nds o f m ode rnspecies .

B roa dly spe a ki ng e a ch ve rteb ra te like o the r a nim a ls a s

, ,

well a s pl a nts i nits devel opme nt a s a ni ndividu a l repe a ts

,

its a ncest ra l h ist ory .

G EOGR AP H I CA L D I S TRI B UTI O N Alth ough th e b roa de r

.

fe a tu res o f th e subj ect will be discussed i na no th er ch a p

ter th e evide nce supplied from thi s field i n supp or t o f
,

ev olu ti ondem a nds a p relimi na ry st a teme nt Inth e fi rst .

pl a ce i t is a m ost signific a nt f a ct th a t a lth ough th e a ni

,

m a ls a nd pl a nts Of oce a nic isl a nds a re disti nct th ey neve r ,

th el ess be a r a nu n mist a k a ble resembl a nce to th ose onth e

a dj a ce nt c onti ne nt In s ome i nst a nces th e isl a nds to
.
, ,

gethe r with th ei r imp ris oned i nh a bit a nts we re cu t Off ,

f rom the m a i nl a nd by me a ns of devel opi ng ch a nnels ;

i n o the r c ases they we re p opul a ted by mig ra nts In .

eithe r eve nt th e is ol a ted c ol onies u nde rwe nt ev ol u

,

ti on a ry ch a n ges a nd g ra du a lly bec a me disti nct Th e .

l onge r th ey rem a i ned disti nct the m ore u nlik e they b e

c a me Th e g re a t di ff e re nces betwee n th e a nim a ls a nd
.

pl a nts of Afric a a nd M a d a g a sc a r or betwee n th ose of

Asi a a nd Aust ra li a for ex a mple a re expl a i ned on th e
, ,

b a sis of a sep a ra ti onof grea t a ntiquity

'

As a result of th is ev oluti on with i n a dist rict h edged ,

a b o ut by b a rrie rs seve ra l cl osely rel a ted species ma y

,

ev olve I ndeed it is gene ra lly t rue th a t the m ost cl osely

.

rel a ted species a re f o u n d i n neighb ori ng a re a s For .

ex a mple the re a re ove r f o u r h u nd red species o f hummi ng

,

bi rds a ll c onfi ned to th e N ew W orld ; k a ng a roos O f seve ra l

,

species a re peculi a r to Aust ra li a a nd th e a dj a ce nt isl a nds ;

,

th e Ol d W orld m onkeys a re ve ry disti nct from th ose o f

the Weste rn Hemisphe re Whe re rel a ted species a re
.
41 8 E VID E N CE S O F E V OLU TI O N

widely sep a ra ted a s i nth e c a se O f th e f a mil y of c a mels

, ,

p a st mig ra ti ons with the extincti onO f i nd ividu a ls i nthe

i nte rve ning te rrit ory a re resp onsible In sh ort a ll O f .
,

th e f a cts o f ge og ra ph ic a l dist ributi on a re s a tisf a ct orily

expl a i ned by ev oluti on O the rwise they a re a se nseless
.

j umble .

AR T I F I CI A L S E LE CT I O N Th a t a nim a ls a nd pl a nts a re
.

no t rigidly fixed but onthe c ont ra ry a re c a p a ble O f ch a nge

is k now nto eve ry b reede r (Ch a p XXX I I I ) H e decides . .

up ons ome ch a ra cte r a speci a l f orm c ol or or qu a l ity , ,

an d selects th ose i ndividu a ls wh ich m a nifest i t in the

high est deg ree Th e su rviv ors i n th e selecti on p rocess
.

a re the n b red ,
th e m ost desi ra ble membe rs a re a g a in
selected a nd b red a nd so on ge ne ra ti ona fte r ge ne ra tion
, ,
.

“
I t w ould seem a s if th ey h a d j ust ch a lked on th e w a ll
a f orm pe rfect i nitself a n d the nh a d give nit existe nce .

Th e two h u nd red a nd m ore b reeds of th e d omestic ,

pige onh a ve bee nde rived fro m th e rock pige onof E u rope
an d Asi a by a rtifici a l selecti on The m a n y v a rieties of
.

d omestic a ted fowls a re th e a rtifici a lly selected desce nd

an t s O f th e j u ngle fowl o f I ndi a wh ich h a s bee nb red for ,

m ore th a nth ree th ous a nd ye a rs S heep c a ttle a nd h ogs .

, , ,

p a nsies tulips d a hli as a nd m a ny o th e r cultiv a ted fl owe rs

, , ,

a s well a s n ume rou s g a rde n veget a bles a re likewise ,

g re a tly a l te red desce nd a nt s o f th e origi na l wild st ocks .

B ut while ma nis th u s a ble to m old a species int o m a ny

distinct v a rieties the re is a wide diffe re nce i none respect
,

betwee nh is results a nd th ose obt a i ned u nde r na tu ra l con

di ti on .s I f th ese d omes tic a ted b reeds a re a ll owed to es
c a pe a nd c ome i nt o c ompetiti onwith wild species they ,

a re u su a ll y exte rmi n a ted E vide n tly th e ch a ra cte rs de

.
,

si red by ma na re no t necess a rily th ose th a t best fit their

p o ss ess ors to c ompete successfully with wild f orms
'

Th e re is h oweve r a ge ne ra l simil a rity betwee nth e a rti

, ,

fi ci a l a n d na tu ra l p rod ucti o n o f species a s will a ppe a r ,

i n the next ch a pte r An d fu rthe rm ore the evide n

. ce ,

s uppli ed by these b reedi n g ope ra ti ons no t only dem on

 G E N E TI C S 41 9

st ra tes th a t species a re c a p a ble of ch a nge but a ls o p ro

vides a clue to the meth od wh e reby species ma y h a ve
devel oped in a st a te of na t u re
G EN ETI CS Inrece nt ye a rs b reedi ng expe rime nts h a ve
.

beenc a rried onby the t ra i ned bi ol ogist not so much for,

the pu rp o se of b ri ngi ng to a h i gh st a te of devel opme nt

s ome p a rticul a r ch a ra cte r i na norg a nism a s to dete rmi ne
th e u nde rlyi ng c a uses of v a ri a ti ons a nd th e m a nne r of
th ei r inhe rit a nce Th is new scie nce of ge netics a s it is
.
,

te rmed is thus l a rgely a study of he redity c ont rolled a nd

,

an a lyzed a t eve ry st a ge .

A sp ecl es of a nim a l or pl a nt 1 s selected for th e exp eri

me nt wh ose a ncest ra l h ist ory or p edig ree is well
k now n I ts devel op me nt a t le a st i n a ge ne ra l wa y is
.
, ,

t ra ced ou t a nd th e beh a vi or a nd st ructu re O f th e ge rm

,

cells i s dete rm ined with a high deg ree of a ccu ra cy Th e .

di ffe re nt ki nds of v a ri a ti ons which occu r du ri ng th e

c ou rse of the expe rime nts a re a ls o r ec orded a nd whe re ,

th ese a re i nh e rited th e mech a nism dete rmi ni ng th ei r i n

h erit a nce is a na lyzed by th e m o st h ighly refi ned meth ods
)

k now nto th e scie ntist Th e dete rmi na ti onof sex a nd th e

.
,

rel a ti on th is ph e n o me n on be a rs to h e redity a re a ls o

ph a ses O f th e ge ne ra l subj ect of ge net i cs P a le ontol ogy

.
,

ge ogra phic a l dist ribu ti on cl a ssific a ti on a nd th e othe r sci

, ,

en ces p revi o usly me n ti oned supply p roof th a t ev oluti on

h a s t a ke n pl a ce A rtifici a l selecti on dem onst ra tes th a t
.

ev oluti on is still p ossible a nd ca n be guided by th e

b reede r a nd the results of studies i nge netics h a ve e na bled
,

th e bi ol ogist to form a fa r cle a re r p ictu re of th e mech a n

ism of ev oluti onth a nwa s k nowni nD a rwi ns time ’
.
 C H APT E R XXX V

TH E O R I E S O F E V OLUTI O N

PR E -
DAR W I N I A N S CI E N CE . Al th o ugh D a rwi nism
nd a

ev oluti ona re p op ul a rly believed to be sy nonym ous te rms ,

the ge ne ra l no ti on is a ctu a lly olde r th a n the C h risti a n

Era . Inthe w riti ngs o f the G reek phil os ophe rs t h e ide a
is occ a si ona lly exp ressed usu a lly i nve ry v a gue l a ngu a ge
,

u nsupp orted by f a cts to a cc ou nt for the origi n a nd

,

devel opment of th e u nive rse i ncludi ng th e org a nic w orld

,
.

A no t a ble excepti ona ppe a rs i nthe c a se o f A rist otle (384

3 22 B C ) . In th e re a lm O f bi ol o gy h e cl a ssified o ve r five
h u nd red a nim a ls u nde rst ood a d a p t a ti on rec og nized the
,
-
,

me a ni ng o f physi ol ogic a l divisi on of l a b or studied em ,

b ry ol ogy a nd a dv a nced th e scie nce O f livi ng thi ngs to th e

,

p o siti oni t h eld for sixtee nh u nd red ye a rs A rist o tle a ls o .

beli eved th a t i norg a nic m a t te r ev olved int o a s o ft m a ss ,

livi ng bu t u norg a nized f rom which highe r species g ra du

,

a lly devel oped a s a result o f a n i n te rna l impelli ng force ,

I n ce rt a i n imp o rt a nt respects his th e o ry resembles th a t

h eld by th e m ode rnbi ol ogist .

Afte r th e time of A rist o tle b o th scie nce a nd th e desi re

,

to fi n d a na tu ra l expl a na ti on for th e origi n o f th i n gs

u nde rwe nt a decli ne l a sti ng u nt il th e 1 2th ce ntu ry .

T ow a rd the cl ose o f th is pe ri od th e w riti ngs of the G reek

phil os oph e rs we re t ra nsl a ted by th e A ra bs wh o u nde r , ,

th ei r i nfl ue nce fo u nded th e fi rst u nive rsities th a t h a d

,

existed for ce ntu ries As a result O f th is i ntellectu a l

.

a w a ke n i ng a l a rge numbe r o f s tude nts a rose wh o p ro

,

fo u n dly i nflue nced scie ntific th ough t a nd di rectly o r in ,

di rectly l a id th e f o u nd a ti ons o f m ode rn scie nce .

T H E I N F LU E N CE O F DA R W I N S W OR E — D u ri ng the
’

4 20
 FA C T O R S OF DA R WIN ’
S TH E OR Y 42 1

f ou r hu nd red ye a rs a fte r the begi nni ng O f the reviv a l of

le a rning na tu ra lists h a d a ccumul a ted a v a st a m o u nt O f
u nrel a ted i nf orm a ti on i n m a ny diffe re nt fields a nd h a d ,

devel oped to a Conside ra ble deg ree th e m ode rns cie nces o f
an a t omy emb ry ol ogy physi ol ogy ge og ra ph ic a l di stri
, , ,

b u ti on cl a ssific a ti on a nd mic ro sc opy

, ,
In the m ore re .

m o te dep a rtments of hist ory l a ngu a ge religi on p olitics , , , ,

etc the re wa s a ls o a h uge m a ss O f u nrel a t ed Obse rv a ti ons

.
, ,

u nrel a ted for th e re a s onth a t the u nde rlyi ng fa ct O f ev ol u

ti onh a d not bee np roved a nd a cc ordi ngly wa s not ge ne r
a lly con side red to be eve na rem o t e p o ssibility .

At the p rese nt time it is difficult to pictu re th e i nflu

e nce up on hum a n th ough t of D a rwi ns th e ory of ev ol u ’

ti on His fi rst w ork

. Th e Origi nof Species a pp ea red
,
“
,

i n 1 859 I t wa s b a sed up on m or e th a n twe nty ye a rs O f

.

c a reful obse rv a ti on wa s cle a rly a nd f a i rly p resented with

, ,

p roo fs a t eve ry st a ge a nd i nth e f a ce O f bitte r opp ositi on

,

—
wa s a ccep ted by th e ope mi nded stude nt
n U nde r its i n .

flu ence ve ry m a ny questi ons o f the m ost p uzzli ng ch a ra c

ter we re m a d e pl a i n a n d th e fl oo d o f ligh t which wa s let
,

i n on th e i ntellectu a l w orld a t th a t p e ri o d h a s i nc re a sed

with e a ch successive ye a r The time wa s m ore th a nripe
.

for the f a c t of ev oluti on to b e rec og nized a nd it is no t

'

to be m a rvelled a t th a t D a rwi n is h a iled a s th e g re a test

scientist the w orld h a s eve r k nown .

FA CT OR S OF DAR W I N S T H EOR Y ’
Th e D a rwi ni a nth e .

ory O f ev oluti on whi ch is a l s o te rmed N a tu ra l S electi on

,

or th e S u rviv a l o f th e Fittest i n v olves seve ra l di ffe re nt

,

fa ct ors Th e fi rst of these is (a ) o ve r p ro ducti on All

.
-
.

an im a ls a nd pl a nts p roduce a fa r g re a te r numbe r O f off

sp ri ng th a nca n su rvive owi ng to l a ck o f sp a ce or f ood
,
.

Si nce (b ) th e numbe r o f i ndividu a ls rem a i ns a b out the

s a me from ce ntu ry to ce ntu ry it f oll ows th a t (c ) the re ,

is a st ruggle for existe nce Fu rthe rm ore th e re is (d) .

,

v a ri a ti on mut a ti on) a nd (e ) h e redity N O two or .

a n i sm s a e ex c tly like nd m a n y o f th ese di ffe re nces

g r a a a ,

a re t ra n smitted from one ge ne ra ti on to the next The .

4 22 TH E O R I E S O F E V O LUTI O N

r esult is th a t s ome a re bette r fitted to thei r su rro u nd i ngs

th a noth e rs which resul ts i n(f) the su rviv a l O f the fi ttest
,

i nth e st ruggle f or existe nce .

All these f a ct ors with one excepti on we re rec og nized

, ,

by seve ra l bi ol ogis ts before th e time of D a rwi n I ndeed .

,

D a rwins g ra ndf a the r E ra smus D a rwi n specific a l ly men

’
, ,

ti ons them i n h is w ritings Th e sup reme w ork o f the .

g ra nds onwa s supplyi ng the evide nce for the su rviv a l o f

th e fi t test wh ich ma y be c on
,
side red th e c orne rst one O f
th e m ost ge ne ra lly a ccepted th e orie s o f ev oluti on a t th e
p rese nt time Th ese v a ri ous f a ct ors w ill now be con
.

si dered inm ore det a il

O VER PR OD U CT I O N The re is no excep ti onto the rule

.

th a t eve ry species of org a nic being a nim a l or pl a nt , ,

na tu ra lly i nc re a ses a t so h igh a ra te th a t if none we re

dest royed th e e a rth w o uld S oonbe c ove red by th e p ro g
,

en y o f a si ngle p a i r I f f o r ex
. a mple eve,
ry h um a n bei
,
ng
we re to c ome to ma turi ty a nd we re to b reed a t th e a ve ra g e
ra te th e n
,
umbe rs O f i ndivid u a ls w o uld be d oubled e ve ry
fifty ye a rs A t th e end o f a few th ous a nd ye a rs if
.
,

th ese c onditi ons p rev a iled th e re w ould not be st a ndi ng ,

room for th e desce n d a nts An a nnu a l pl a nt p rod ucing

.

two seeds on ly w o uld h a ve O ffsp ri n g i ntwe nty

on e ye a rs if e a ch i n dividu a l a tt a i ned m a tu rity a nd rep ro
duced Th e c o nge r eel l a ys
. eggs e a ch ye a r th e ,

codfi sh th e V i rgi ni a oyste r not fewe r th a n

on th e a ve ra ge I f th ese we re to c ome to
.

m a tu ri ty a nd b reed thei r p roge ny w ould fill th e sea i n

,

a ve ry b rief pe ri o d This f a ct or th eref ore i nth e th e ory

.
, ,

o f ev oluti on is a ge n e ra lly rec ognized a nd fi rmly est a b

l i sh ed f a c t .

STAT I O N AR Y N U M BER S OF I N DI V IDUA LS I t is a ls o wel l .

est a blish ed th a t th e numbe r of i ndividu a ls o f the ma

j o ri ty o f species r em a i n s f a i rly c onst a nt o ve r l on g pe ri ods .

The re ma y be a ni nc re a se a t one time a nd a dec re a se a t

an oth e r but i nthe l on
,
g runthe a ve ra ge is a pp roxim a tely
u nif o rm Th is is to be expected whe nit is remem b e red
.
 T HE S T R UGGL E FO R E X I S T E N CE 423

th a t eve ry species o f a nim a l a nd pl a nt is fitted to a

defi nite set of c onditi ons a nd th a t withi nits ra nge e a ch
,

S p ecies fi n ds a limited f ood supply e nemies O f m a ny ,

ki nds a nd u nfa v ora ble a s well a s fa v ora ble c onditi ons O f

,

s oil a nd clim a te All of th ese a ge ncies tend to h old th e

.

numbe r O f i ndividu a ls a t a give n level .

T H E S TR U GGLE FOR EX I S TEN CE — I t is a ls o m a n ifest

th a t si nce m a ny m ore i ndividu a ls a re p roduced th a nca n
c ome to m a tu rity a st ruggle for existe nce is th e i nevit a ble
,

resul t B roa dly spe a ki ng this is a b a ttle a g a i nst th e sur

.
,

ro u n di ng c onditi ons a n e ffort c onsci ous or othe rwise on

, , ,

the p a rt of th e org a nism to pl a ce itself i nth e m ost f a v or

a ble rel a ti on s to th e e nvi ronment In th is c omb a t a ni .

m a ls pl a y a m ore obvi o us p a rt bu t it is none the less t rue ,

th a t pl a nts a re a t wa r with one a no the r a nd V ict ory ,

c omes to a ve ry few This st ruggle a g a inst u nf a v ora ble

.

c onditi ons p rese nts a th reef old a spect Th e i ndividu a ls .

of the s a me species ma y c ompete with on e an o the r A .

pl a nt ma y bec ome o ve rsh a d owed by its fell ows a nd pe rish

for l a ck O f su nligh t A ra t ma y figh t with its own ki nd
.

an d eve n dev o u r its y o u ng Or membe rs of one species.

ma y ente r i nt o c ompetiti on with th ose of a nothe r The .

e a gle st rikes d own a f a w n a nd l ocusts c onsume th e h a r

,

vest c rops Fi na lly the re is a c onti nu a l st rivi ng on the

.
,

p a rt of a n org a nism to withst a nd ext remes of he a t a nd

c old a l a ck or a supe ra bu nd ance of m o istu re a nd o the r
, ,

u nf a v ora ble physic a l fe a tu res of its su rrou ndi ngs .

I N TERRELATI O N S OF OR GA N I S M S Th e e nvironme nt of .

a na n im a l or a pla nt is m ore th a n a S imple mixtu re of

light he a t a i r m oistu re a nd s oil plus o the r org a nisms
, , , , ,
.

These elements a re p a rts O f a c omplex system whe re

i n a rel a tively sm a ll m odific a ti on a t one p oi nt ma y
p roduce h ighly imp ort a nt ch a nges i n a dist a nt a nd fre
quently u nexpected qu a rte r In this N a tu re resembles
“

.
,

a p o ol whe re a p ebble d ropped i n t o th e dep th s c a uses

fa r re a ch i ng effects
-
.

D a rwi n c a lls a tte nti on to the f a ct th a t bumbl e bees

4 24 TH E O R I E S OF E V OLUTI O N

c a rry p olle n f ro m one cl ove r pl a nt to a noth e r a nd the ,

fl owe rs th us f e rtilized be a r seeds B umble bees a re de .

st royed by mice which i n tu rn a re e a te n by c a ts The

, .

m ore c a ts th e fewer mice a nd th e g re a te r numbe r o f bees

,

an d cl o ver seeds E a rthw orm s by tu n

. nel ing th e s oil a nd
,

c a rryi ng org a nic f ood f ro m the su rf a ce i nt o thei r b ur

ro ws a re high ly imp o rta n
, t a ge nts i n l oose ni ng up a nd
fe rtilizi ng th e s oil Th ese a nim a ls th e ref o re a re a ni m
.
, ,

p ort a nt f a ct o r i n dete rmi ni ng the size of c rops a nd the

numbe r o f h um a ni nh a bit a nts i nthe regi on The h ist ory .

o f th e ra bbi t i n Aust ra li a th e p o t a t o b eetle i n I rel a nd

, ,

th e c odli ng m o th sc a le bugs th e w a te r hy a ci nth thistle

n
d O f m a y o the r a ni m
, , , ,

an al an d pl a nt pests a ls o illust ra te ,

h ow the b a l a nce o f na tu re ma y be se ri o usly dest royed

by insig nific a nt c a uses .

VAR I AT I O N AN D H ER ED I TY — While i t is t r ue th a t .

org a n isms te nd to resemble thei r p a re nts i t is equ a lly ,

ce rta i n th a t th ey a re ra rely if eve r th eir ex a ct c ou nte r

p a r ts I n th
. e h um a n r a ce f or ex a mple th
,
e re o fte n a re ,

st riki ng f a mily resembl a nces a nd yet i nv a ri a bly i ndivid

u a l di ff e re n ces i n f orm a nd st ructu re ca n be sh ow n to
exist In s ome i nst a nces this u nlike ness or v a ri a ti on is
.

th e resul t of exe rcise or s ome d i rect effect o f th e e nvi ron

me nt An a bu nd a nce or l a ck of o u rish me nt th e exer
. n ,

cise o f a m uscle th e eff ec t o f su nligh t a nd m a ny o th e r

, ,

a ge n cies ma y p rod uce m a rked ch a nges i n a n a nim a l or

pl a nt du ring th e c ou rse of its lifetim e S uch e ff ects so .
,

fa r a s is k now n a re no t t ra nsmitted to th e o ff sp ri ng ; th ey
,

a re n onhe ri ta ble v a ri a ti o n
-
s or m odific a ti ons O n the .

o the r h a n d the re a re di ff e re nces O f c ol or f o rm p rop orti on , , ,

size a nd th ou s a nd s o f o the r ch a ra cte rs k now na s mut a

, ,

t i ons whi ch a re t ra nsmi tted f rom ge ne ra ti onto ge ne ra

,

ti on Th e p a rt wh ich th e ge rm cells p l a y i nthis p rocess

.

wa s e n ti rely u nk now ni n D a rwi ns time Si nce h is da y ’

.

a g re a t a m o u n t o f i nform a ti onh a s a ccumul a ted a nd th e ,

me th od o f th e t ra nsmissi o n of h erit a ble v a ri a ti ons is

a lm o st a s well k n o w n a s th e st ructu re of a n i ma ls a nd
 T HE S U R VI VAL OF T HE FI TT E S T 4 25

pl a nts For the fi rst time the bi ol ogist h a s re a ched the

.

p o int wh e re the mech a nism o f he redity is f a i rly well

u nde rst ood a nd the ca uses a nd meth ods O f ev oluti on
,

st a nd ou t i na cle a re r ligh t .

T H E S U R V I V A L O F H E F I TTE S T
T — This te rm sig nifies
.

th a t i n th e st ruggle for exis te nce a m ong a nim a ls a nd

pl a nts th ose i ndividu a ls with the g re a test Vig or th e ,

kee nest senses th e highest p owe rs O f speed or th o se

, ,

o the rwise best a d a p ted to thei r su rro u n di ngs a re the ones ,

which tend to o utl a st th ei r c omp a ni ons Th ey a céord .

in gyl a re the o nes m ost likely t o p r o duce o ffsp ri ng And .

these i ndividu a ls a re likely to t ra nsmit th e s a me h e rit a ble

v a ri a ti ons to s ome of thei r p roge ny wh ile th e o the rs less,

fit will die i nth e st ruggle F ro m time to time mu t a ti ons

.

o ccu r i non e or m ore i n dividu a ls Whe re these a re be ne

.

fi ci a l the p o ssess or is m ore ce rt a i nto be p rese rved a n d to

t ra nsmit this new ch a ra cte r to s ome o f its desce nd a nts .

I f th e mut a ti onre nde rs a n org a nism less fit less a ble to ,

withst a nd the rig ors of its su rrou ndi ngs i t pe rish es

n
.
,

N a tu re thus c o ti nu a lly p uts a p remium on the fit test

i ndividu a ls while th e impe rfect ill a d a p ted ones a re v a n
,
-

q u i sh ed i n th e c o mb a.t Th e species is t h us g r a du a lly

ch a nged or i n o the r words i t ev olves by the i ncorp o “

‘

ra ti oni n t o the ra ce of th ose mut a ti ons th a t a re be nefici a l

to the life a n d rep ro ducti onO f th e org a nism
” .

A PP LI CATI O N OF T H E T H EOR Y —The re is c onclusive

.

evide nce th a t the ex cessively l ong necked gi ra ffe c a mels -

ev olved from a rel a tively S h ort necked a ncest or (see p a ge

-

It is a ls o highly p rob a ble th a t th e m ode rn gi ra ffe

‘

likewise descended from c omp a ra tively sh ort ne cked dee r -

like p roge nit ors Acc ordi ng to th e th eory O f N a tu ra l

.
j

S electi on s ome of th e i ndividu a ls of th is a ncie nt s tock

p ossessed s omewh a t l onge r ne cks th a n thei r fell ows .

They we re th us e na bled to b rowse i nthei r semi a rid su r -

ro u n di ngs on foli a ge ou t of re a ch o f th e o the rs Thi s .

a dv a n t a ge seemi ngly i nsignifi ca nt neve rtheless e na bled

, ,

m ore l ong necked th a n sh ort necked i ndividu a ls to c ome

- -
 6 T H EOR I E S O F E V O LUTI O N

to m a tu rity a nd re p rod uce th eir kind N ew v a ri a ti ons .

,

i nthe f o rm o f mu t a ti on s a ppe a red du ring th e l ong h i s

,

t o ry o f th e ra ce Wh e re these resulted i n l onge r necks

'

th a n before they we re i nc orp ora ted int o the ra ce Th e .

gre a t h eigh t a nd exceedi ngly l ong neck of the gi ra ff e is

th e ref ore n o t th e result o f o ne v a ri a ti oni nth e h ist o ry o f

a dee r l i ke a n cest or On the c ont ra ry th e ev oluti on O f

'

-
.

th e species is the result of th e g ra du a l a c cumul a ti on O f

m a ny c omp a ra tively sligh t v a ri a ti ons or mut a ti ons wh ich
no t Only m odified th e ra ce in respect to th e neck but
e na bled it to withst a nd th e a tt a cks of e nemies a nd th e
seve re c onditi ons of its su rrou nd ings This s a me li ne o f .

re a s on i ng i s a pplic a ble i n the c a se of eve ry o th e r species

of a n im a l a s well a s pl a nt .

S EX UA L S ELE CT I O N — I h a n o th e r c on
. necti on (Ch a p .

XX V I ) i t h a s bee n sh ow n th a t a m ong m a ny ki nds of

im a ls m a rked diff e re n
,

an ,
ces exist be twee nthe m a les a nd
fem a les Th e re a s onfor this f a ct a ppe a red to D a rwinto
.

lie m ore or less bey ond th e re a lm o f N a tu ra l S electi on ,

an d w a s ra th e r to be expl a i ned on th e b a sis o f S exu a l

S electi on Acc o rding to this th e o ry th e m a les eith e r
.

e ng a ge i n a st ruggle a m o ng themselves for a p a rtic ul a r

m a te o r a m ong o the r spe cies they a re ch ose n by th e
, , ,

opp o site sex Whe n a b a ttle e nsues it is a well k now n

.

f a ct th a t i nm a ny i nst a nces th o se m a les with th e g re a test

st re ng th a nd th e m o st highly devel oped we a p ons c ome
o ff vict o ri o us a n d le a ve desce nd a nts Th is is eve nm ore .

a pp a re n t a m ong p olyg a m o us a nim a ls whe re one m a le ,

figh ts for th e p ossessi onO f seve ra l fem a les a s i nth e c a se

o f th e f u r se a l sea li o n d omesti c f owl a n
, ,
d seve ra l o th e r ,

species I n a l l O f these c a ses th e re is a su rviv a l o f th e

.

fittest a nd a t th e p rese nt time this ph a se of sexu a l sel ec

,

ti on is expl a i n ed on th e b a sis O f N a tu ra l S electi on .

Am ong the bi rds i nse cts th e c ra bs a nd rel a t ed species

, , ,

whe re th e m a le is O f te n highly c ol o red o r dis ti nguish ed

by o th e r se co nd a ry sexu a l ch a ra cte ris tics the fem a le , ,

a cc ord i n g to the the ory o f S exu a l S electi on is believed to ,

 L AM AR CKI S M 4 27

ch oose a m
a le wh o se be a u ty g ra ce or o the r c on , spicu ou s
fe a tu re s a tisfies h er esthetic se nse Th e su rviv a l O f the .

fittest is thus dete rmi ned by the fem a le AS a m a tte r .

O f f a ct it h a s bee n sh o w n th a t a m on g m a ny i nsects th e
wings ma y be dyed or evencu t off with out distu rb ing the
m a ti ng i nsti nct Th e sexes a re a tt ra cted by speci a l od ors
.
,

an d no t by c ol or C e rt a i n c ra bs a n
. d rel a ted species a s, ,

well a s th e Spide rs m a te by me a ns of the se nse of t ouch

, .

Am ong th e bi rds th e fem a le usu a lly p a ys no a tte nti on

to the m a le wh o i s S h o win g O ff ” The rem a rk a ble bi rd
.

‘
d a nces u nde rg one by ce rt a i n species o ccu r a fte r m a ti ng
h a s a l re a d t a ke n pl a ce F rom these a n d m a ny o th e r
y .

ex a mples which ca nbe cited the re is little evidence th a t

, ,

th e fem a le ch ooses h er m a te On th e othe r h a nd it is .

not cle a r th a t such st ructu res a s th e g re a t t a il of th e

pe a c o ck the tuft of h a i r like fe a the rs onthe b re a st O f th e
,
-

tu rkey th e w a tt les plumes spi nes k nobs a nd consp i cu

,
-
, , , ,

ou s c ol ors p o ssessed by the m a les of m a n y species a re Of

sup reme imp ort a nce i n th e life a nd de a th st ruggle for
existe nce In sh ort th e re is a s yet no s a tisf a ct ory a ll
.
,

emb ra ci n g e x
'

p l a na t i o n o f th e S ig nific a nce o f sec ond a ry

sexu a l ch a ra cte rs .

L A M AR CK I S M . I t is a nu nc ontested f a ct th a t a nim a ls
an d pl a nts a re c onti nu a lly bei ng m olded u nde r the i n
flu ence of th ei r su rro u ndi ngs F ood a nd clim a te a re i m
.

p ort a nt a ge nts i nthis respect E xe rcise i nc re a ses th e size

.

o f muscles a n d disuse c a uses thei r sh ri nk a ge D rugs such .

,

a s m orphi n e a nd a lc oh ol m odify th e org a nism a nd m a ny

, ,

o the r f orces c a use m ore or less exte n sive a lte ra ti ons .

Acc ordi ng to th e F re nch scie ntist La m a rck (1 74 4 1 829) — ,

th e ch a ra cte rs or m odific a ti ons thus a cqui red by the

org a n ism du ri ng its lifetime a re supp osed to be i nhe rited
by the Off sp ri ng This simple pl a usible the ory wa s a c
.

cep ted by D a rwi n a s a f a ct or i n ev oluti on a ry p rocesses ,

an d wa slikewise a ccep ted a s a m a tte r o f c ou rse by seie n

ti sts a n d l a ity a like u ntil rece nt ye a rs .

Th e l ong neck o f the gi ra ffe a cc ordi ng to th e L a m a rck ,

4 28 TH E O R I E S O F E V O L UTI O N

i a nthe ory , is due to th e f a ct th a t the dee r lik e a ncest ors -

st retched a nd so le ngth e ned thei r necks to a sligh t ex tent .

Th e immedi a te desce nd a nts i nhe rited l onge r necks i ncon

seque nce o f th ei r p a re nts a c tivi ty G e ne ra ti ona fte r gen
’
.

era ti o n th is p rocess O f s tretchi n g a nd t ra nsmi tti ng th e

e ff ec ts we nt on a nd th e i nc re a si ngly l ong necked s p ecies
,
-

ev olved int o the m ode rn gira ff e .

S cie ntific j o u rna ls a nd th e public p ress rec o u nted in

st a nces of th e (supp o sed ) i nhe ri ta nce o f m util a ti ons th e ,

t ra nsmissi onof a p owe rful physique d ue to exe rcise a nd ,

implicitly or e xplici tly th e w ork o f the sch ools ch u rches

, , ,

a sylums h o spit a ls
,
an d o the r ph il a nth ropic i n
,
stituti ons
wa s b a sed onth e belief th a t the be nefi ts c onfe rred up on
on e ge n e ra ti on a re i nhe rited by the next .

T H E N E W L A M AR C K I A N S CH OOL — T ow a rd th e cl ose
of th e l a st ce n tu ry Weism a nnch a lle nged this th e ory o f
,

the i nhe rit a nce of a cqui red ch a ra cte rs a nd dem a nd ed ,

m ore c onclusive evide nce th a nh a d hith e rt o s a tisfied th e

scientifi c w orld He a ls o di rected a tte nti on to th e f a ct
.

th a t a m odific a ti on such a s a ne ck le ngthe ned by st retch

,

in g
,
ca n o nly be t ra nsmitted to th e succeedi n g ge n e ra ti o n
by me a ns of th e ge rm cells And no t only th is but th e .
,

L a m a rcki a n th e ory requi res th a t p recisely th a t p orti on

o f e a ch ge rm cell wh i ch c on t rols th e devel opme nt of th e
neck m ust be ch a nged to th e ex a ct exte nt requi red for
th e i nhe rit a nce o f th e m odific a ti o n As a resul t of th is .

ch a lle nge old evide nce wa s c a refully sc ru ti nized a nd

,

f o u nd to be i nc onclusive N ew evide nce wa s s ough t bu t

.
,

wi th o ut a v a il T od a y th e re a re no k now n c a ses wh e re
.

defi nite specific ch a nges i na norg a nism d ue to th e i nflu

,

e nce of th e e nvi ronme nt onthe b ody a re i nh e rited by the

O ff sp ri ng .

Th e re is evid e nce th a t th e f orm c ol or size vit a lity , , , ,

etc o f a ni nd ividu a l ma y be m odified by e nvi ronme nt a l

.
,

i nflue nces a nd th e des ce nd a nts o f such p a re nts ma y

,

v a ry m o re or less widely f ro m th e norm a l type For ex .

a mple th e size O f seeds st a n

,
d s i n cl ose rel a ti on to the
 O R T H O GE N E S I S 4 29

a m ou nt of nou rishme nt received by th e p a re nt If . .

st a rved a nd highly nou rish ed seeds are pl a nted i n the

s a me envi ronme nt they respectively produce sm a ll a nd
l a rge pl a nts I t is imp ort a nt to note h oweve r th a t such
.
, ,

e ffects p a ss a w a y i n a few ge ne ra ti ons Th e ge rm cells .

h a ve not bee n a ffected a nd th e m odific a ti on is no t i n

,

h eri ted Th e s a me f a cts h a ve bee n sh ow n to be t rue i n

.

th e c a se O f ce rt a i norg a nisms subj ected to di ffe re nt deg rees

o f tempe ra tu re m oistu re light d rugs a n
, , ,
d seve ra l o the r ,

m odifyi ng a ge nts Th e e ffects of th e e nvi ronme nt i n

.

such c a ses a re not pe rm a ne nt a nd he nce a re not h e rit a ble

, .

On the o the r h a nd i t is bey ond questi on th a t mut a

ti ons do a ppe a r from time to time I t i i s a ls o well est a b .

l i sh ed th a t this is due to th e re a rra nge m e nt of th e he redi

t a ry ch rom a ti ni n th e ge rm cell In s ome c a ses the re is
.

evidence th a t such ch a nges a re cl osely rel a ted to tempe ra

tu e r — a n en vi ronme nt a l f a ct or whic h th us a ppe a rs to
m odify h e redity S i nce th e ge rm cell s a re p rot opl a sm ;
.

an d p rot opl a sm is c a p a ble O f ch a nge zi t ma y i ndeed be

“

t rue th a t e nvi ronme nt a l influe nces h a ve a ne ff ect They .

a re believed to h a ve by the l a te r La m a rcki a nsch ool but ,

th e questi onis a s yet a h otly deb a ted and u nsettled one .

E ve nif it we re settled i n th e a ffi rm a tive h oweve r it is , ,

p rob a ble th a t i t w o uld do no m ore th a nS hed ligh t up on

the origi n of h e rit a ble v a ri a ti ons a nd w o uld i nno wise,

u nse a t the the ory of N a tu ra l S electi on .

ORT H OGE N ES I S OR D ETER M I N ATE E V OL UTI O N Th e .

ev oluti on o f the org a nic w orld ma y be c omp a red to the

growth of a t ree the tips of th e b ra nch es c orresp ondi ng
,

to th e m ode rn species o f a n im a ls a nd pl a nts And j us t .

a s the t ree is c a used to g ro w by f orces withi n it so a , ,

c onside ra ble numbe r of scie ntists a re c onvi nced livi ng ,

thi ngs a re c onst ra i ned by thei r st ructu re or by i nte rna l

or exte rn a l f orces or by these i n c ombi n a ti on to ev olve

a l on g fa i rly defi nitely fixed p a ths S uch i ngene ra l is th e

.

me a ning of Orth oge nesis .

Acc ordi ng to th e m ost ext reme V iew p rot opl a sm is en ,

4 30 TH E O R I E S O F E V O LUTI O N

d owed wi th a peculi a r f orce which not only impels th e or

,

g a n i sm to ev o lve i n a defi n ite di recti o n but a

, ls o a d a pts it
to its su rrou n di ngs A less ra dic a l p ositi on is t a ke n by
.

ce rt a in o the r scie ntists wh o m a int a i n th a t a t ce rt a i n

defi ni te pe ri ods th e st ructu re o f a n a nim a l or pl a nt de
te rm ines th e ge ne ra l t re nd of ev oluti on a fte r which th e
,

su rrou ndi ng c ond iti ons m old the species to fit th e e nvi ron
me nt S eve ra l o the r the ories m ore or less L a m a rcki a n
.
,

i np ri nciple h a ve a ls o bee np rop o sed a s a ll suffi cie nt ex

,
-

pl a na ti ons o f ev oluti on or a s subsidi a ry th e ories to a c

,

c o u nt for th e orde rly a nd seemi ngly p redete rmi ned c ou rse

O f species devel opme n t .

A t the p rese nt time th e m a j ority of bi ol ogists a re a g reed

th a t h e rit a ble v a ri a ti ons a re to a ce rt a i n exte nt limited ,

th e ra nge bei ng dete rm ined by th e st ructu re of th e org a n

ism c once rned As Huxley w rote wh a les neve r p roduce
.

fe a the rs nor bi rds wh a leb one

,
N eve rtheless e a ch
.
,

species of wh a le bi rd or o the r species o f a nim a l or pl a nt

, ,

ma y v a ry i na ny one o f th ous a nd s of d i ffe re nt di recti ons .

P rovided the mut a ti onh a rm onizes with th e e nvi ronme nt ,

ev oluti on ma y a cc ordi ngly t a ke pl a ce a l ong a ny one O f

a multitude o f di ffe re n t li nes a nd not a l ong th e limit ed
,

numbe r dem a nded by th e usu a l th e ory o f Orth oge nesis .

 C HAPTER XX X VI

T HE R E S UL T S O F E V O LUTI O N

ADAPTAT I O N Th e cu rre nt o f life a s it h a s fl owed

.
-
,

th rough the a ges resembles a mighty rive r or bette r, , ,

pe rh a p s the sl ow m ovi ng gl a cie r which a t v a ryi ng dis

,
-
,

t a nces from its s ou rce b re a ks up or ev olves i nt o nume ro us

b ra nches And i nmuch th e s a me m a nne r th a t the gl a cie r
.

fits i nt o th e i nequ a lities o f i ts bed th e v a ri o us species of ,

an im a ls a nd pl a nts fi t i nt o or a re a d a pted to thei r su r

ro u ndi ngs This is th e g re a t o utst a ndi ng ch a ra cte ris ti c
.

of livi n g thi ngs Fu rthe rm ore a d a pt a ti ond oe s not st op

.
,

with the org a nism a s a wh ole On the c ont ra ry eve ry .

,

system org a n tissue cell a nd th e elements of th e cell

, , ,

itself a re a ll a d a p ted p a rts O f th e V it a l m a chi ne Th e .

r esults of ev oluti on the ref or e a re l a rgely summed up i n

the one w ord a d a pt a ti on ,

.

CO N VER G N T A APTAT ON S N A N M A LS
E D I I I — V iewed i n .

its b roa de r a spects one st riki ng fe a tu re o f a d a pt a ti on

,

a m on g org a nisms is th e so c a lled c onvergence or p a ra llel -

ism of f orm Th a t is to sa y m a ny species o f a nim a ls a nd

.
,

pl a nts su rrou nded by esse nti a lly th e s a me condi ti ons,bea r

,

a supe rfici a l resembl a n ce to e a ch othe r T h e wh a le for .

ex a mple h a s bee n c a lled a fish since the d a ys of J ona h .

Inre a lity th is i s a c a se Of c on ve rge nce wh e reby a h a i ry ,

w a rm bl ooded ve rteb ra te or m a mm a l h a s ev olved a f orm

-
, ,

o f b o dy a d a pted for cle a vi n g th e w a te r with the le a st

p ossible resist a nce Its resembl a nce to c old bl ooded .
-

fish is e nti rely supe rfici a l Th e s a me is t rue of the ex .

tinct fi sh like reptiles wh ich l ong a go i nh a bited the sea

'

-
.
,

A nothe r f a mili a r ex a mple O f conve rge nce occu rs a m ong

the bi rds b a ts ce rt a in a ncie nt rep tiles a nd to a less ex
, ,

431
4 32 T HE R E S UL T S O F E V OLUTI O N

te nt a m ong th e flyi ng squ i rrels flyi ng lemu rs a nd ce rt ai n ,

species of liz a rds He re a g a i nth e wi ngs a nd pl a nes a re

.

a d a pt a ti on s wit h th e s a me ge ne ra l fu nc ti on th o ugh the

o rg a n s th emselves a re st ructu ra lly q uite diffe re nt .

On the high se a s m a ny ve rteb ra tes s na ils c ra bs , , ,

sh rimp s j elly fi sh es a nd th e y o u ng O f m a ny oth e r equ a lly

,
-

dist a ntly rel a t ed species a re t ra nsp a re nt a s gl a ss a con ,

ve rge nt a d a p ta ti on e na bli ng such a nim a ls to esc a pe th e

,

notice o f m a ny sh a rp sigh ted e nemies N ume rous a qu a tic

-
.

org a n isms a re p ro vided with O i l a nd fa t which dec re a ses

th ei r specific g ra vi ty or th ey a re fu rnish ed wi th ga s b a gs
,

o r fe a th e r like o utg ro wth s which p reve n

-
t thei r ra pid si nk
i ng Th e h e a vy f ore limbs O f diggi n
. g a nim a ls th e cl a ws ,

O f th o se which climb the l ong legs a nd te nde ncy to go

,

a b o u t o n tip t o e c o mm on to m a n y ru nni ng species th e ,

m ore or less bli nd a nd c ol orless c onditi onO f ca ve a nim a ls ,

th e ph o sph ores ce nt o rg a n s o f nume ro us deep sea org a n

isms a nd m a ny o th e r ex a mples k nown to th e obse rv a nt
s tude nt illus tra te c onve rge nt a d a pt a ti on .

CO N V R G N T A D APTAT O N S A M O N G P LA N T S
E E I — In th e
pl a nt ki ngd om no less th a na m ong a nim a ls the re a re m a ny
ex a mples o f c onve rge nt ev oluti on N ume ro us species o f .

widely di ffe re nt rel a ti onships h a ve ev olved th e climbi ng

h a bit i n th e st ruggle f or existe nce By m e a ns o f ten
, .

d rils h oldfa s ts or by twi ni ng a b ou t s ome supp o rt they a re

, ,

a d a p t ed t o s ca le a w a ll o r f o rest t ree a nd exp ose the i r

le a ves to th e light In th e m ost ext reme c a se ce rt a i n
.
,

p a lms c omm only k now na s ra t ta ns f orm gre a t fest oons

, ,

i n t ropic a l f ores ts a n d n ot i nf reque ntly a tta i n a le ngth

,

o f 60 0 to 900 feet th e g re a t est k now n le ng th o f a ny

,

o rg a n ism .

I nth e life of pl a n ts of a rid dist ricts it is o f the utm ost

imp o rt a nce th a t the w a te r a bs o rbed by the roots be co n
se rved a s fa r a s p ossible T 0 this end th e le a f su rf a ce is
.

g re a tly reduced thus p reve nti ng ra pid ev a p ora ti on; a nd

,

v a ri ous spe cies h a ve ev olved w a te r st ora ge tissue givi ng ,

the le a ves or stems a thicke ned fleshy a ppe a ra nce These .

 OTH E R CLA S SE S OF ADAPTATI O N S 4 33

a re c onve rge nt a d a p t a ti ons ch a ra cte ristic of nume rous

dese rt pl a nts the w orld ove r I ncide nt a ll y it ma y be s a id
.

th a t a s a result of th is a dj ustme nt to th e e nvi ronme nt

th e a g a ve rep rese nted by such species a s th e ce ntu ry
,

pl a nt O f this c o u nt ry be a rs a cl ose resembl a nce to th e

,

ol d w orld a l oes ,
an d th e c a ctus cl osely duplic a tes the
Afric a nspu rges .

Th e pl a nts of high m o u nt a i ns a nd of th e t reeless re

gi ons of th e fa r north wh e re deep s nows p rev a il th rough
,

ou t th e g re a te r p a rt o f th e ye a r f orm l ow de n
,
se m a ts or
c a rpets C onve rgent a d a p t a ti ons a pp e a r i n th e c a se of
.

m a ny species whe re the ove r l a ppi ng le a ves chiefly rest

-

on th e g ro u n d a nd dimi nish th e l o ss of h e a t wh ile th e ,

roots a re ge n e ra lly l oa ded with st a rch a nd fu rnish th e

f ood necess a ry for th e ra pid growth of th e pl a nt du ri g n
th e sh ort summe r .

Th e pl a nts a d a pted to a nex iste nce i nw a te r or to the ,

s a ndy st retches b orde ri ng th e sea a nd the l a rge r l a kes ,

or to a n y o th e r r el a tively l a rge te rr it ory whe r e esse nti a lly

th e s a me c onditi ons p rev a il p ossess ce rt a i n fe a tu res i n
,

c omm on; but c onside ra ti ons of sp a ce p revent th ei r

“

det a iled descripti on .

OT H ER C LA S S E S OF A DAPTATI O N S While the a nim a ls

.

an d pl a nts O f a give nregi on or i ndi ff e re nt dist ricts wh e re

,

the s a me ge ne ra l c onditi ons p rev a il ma y m a nifest this ,

c onve rge nce or p a ra llelism O f f orm it is equ a lly t rue th a t

,

eve ry species p ossesses its own p a rticul a r a d a p t a ti ons

which e na ble it to live a mid su rro u ndi ngs m ore or less
diffe re nt from th o se of a ny o th e r species I ndeed it is .

these mi nor a d a pt a ti ons wh ich i nl a rge me a su re se rve to

disti nguish one species fro m a no th e r Th e rel a ti ons of .

pl a nts to thei r e nvi ronme nt a re usu a lly less c omplex th a n

i na nim a ls with h ighly devel oped ne rv o us systems se nse ,

org a n s a nd a ctive p owe rs O f l oc om o ti on Acc ordi ngly

n
,
.
,

pl a nt a d a p t a ti ons a re less Obvi o us a d no t ge ne ra lly so

st riki ng a s th o se i nthe a nim a l kingd om th ough they a re ,

a s un ive rs a l a nd bel ong to th e s a me ge ne ra l cl a sses .

434 T HE R E S UL T S O F E V O LUTI O N

Th ese lesse r specific v a ri a ti ons h a ve bee n v a ri ously

cl a s sified Acc ord ing to one rece nt V iew the re a re five
.

ge ne ral c a teg ories : (a ) food secu ri ng ; (b ) self defe nse ;

“
- -

( ) defe n se f n g d
; ( ) iv l y ; ( ) a dj ustme n t to
”
c o y ou r a r e

su rro u ndi ngs .

F OOD S EC UR I N G AD APTAT O N S
-
I — Teeth a re p rese nt i n
th e g re a t m a j ority of th e h igh e r a nim a ls a nd a re v a ri ,

ou sl y a d a p ted fo r seiz in g a nd killi ng p rey or biti ng off

veget a ble m a te ri a l or te a ring th rough the p ro tective
,

husks a nd sh ells of nuts a s well a s for c rush i ng or cutti ng

‘

the food p rep a ra t ory to sw a ll owi ng S o cl ose is th e f orm .

o f th e teeth rel a ted to thei r fu n cti onth a t th e p a l eontol o

gist is f reque ntly a ble not only to dete rmi ne f rom a
si ngle t oo th the rel a ti onships o f the species but a ls o to
form a fa i rly cle a r p ictu re of its f ood a nd feedi ng h a bits
»

Th e s a m e p ri nciple a pplies to a n a lm ost equ a l exte nt i n

th e c a se of bi rd s C e rt a i np a rrots h a ve a rel a tively l ong
.

an d cu rved be a k a d a pted for diggi ng up roo ts O the r .

species p ossess a he a vie r sh orte r bill se rvi ng for th e ,

—
c ra cki ng of nuts a nd o the r thick shelled seeds The .

Sp a rrows a nd o th e r seed e a ti ng bi rd s h a ve he a vy c onic a l

-

bills while th ose of the w a rble rs a nd ce rt a i nothe r i nsect

,

e a te rs a re m ore sle nde r .

Inm a n y of th e l owe r a nima ls such a s th e i nsects a nd ,

c ra bs a nd rel a ted species th e m outh is u np rovided with ,

teeth a nd ce rt a i n o f th e legs h a ve bec ome a d a pted to

,

form cutti ng or pie rci ng org a ns Th e bill of th e mos .

“

qui t o a nd th e l ong p robo scis o f th e bu tte rfly a re org a ns

of this type Fi na lly the re a re m a ny a tt a ch ed or st a ti on
.
,

a ry a n im a ls such a s sp onges c ora ls nume rous species o f

, , ,

w orms b a rna cles a nd v a ri ous cl a ms which obvi ously

, , ,

mu s t subsist up on f ood b ro ugh t with i n re ac h In th e .

b a rna cles nume rou s legs cre a te cu rre nts i n th e w a te r

,

an d gra sp th e f ood a s i t p a sses by I n m ost O f the re .

m a i ni ng species m enti oned my ri a ds O f cil i a on th e su r ,

f a ce o f th e b ody d rive fl oa ti ng nut ritive p a rticles i nto the

m outh .
 P LANT S -
ADA P T E D TO A B S OR B GA S E S 435

P LA N TS A DAPTED To A B S OR B GA S E S W ATER AN D SA LTS , , .

-
S i nce these ra w m a te ri a ls a re a bs orbed th rough th e
le a ves a nd roots it f oll ows th a t the f orme r must be
p rope rly exp osed to the a i r a nd th e l a tte r to the s oil .

Le a ve s must not only a bs orb oxyge n f rom th e a i r a nd

disch a rge c a rb ondi oxid i nt o it i nthe d a rk bu t they must
a ls o r eve rse this orde r i n su n light a nd a bs orb c a rb on
di oxid a nd ligh t e ne rgy a nd disch a rge oxyge n T o do .

this efficie ntly requi res m a ny sh a pes a nd p ositi ons of

le a ves .

Th e roo ts likewise p rese nt a wide ra nge of v a ri a ti on .

P rim a rily they a re a d a pted to a bs orb w a te r a nd diss olved

s a lts a nywh e re from th e su rf a ce o f th e e a rth to a dep th O f
a t le a st seve n ty fi ve feet i n the c a se o f ce rt a i n dese rt
-
,

species At the s a me time it is highly imp ort a nt th a t

n
.

they fu ncti o a s h oldf a sts to a nch or th e pl a nt i na secu re

p ositi on .

In p a ra sitic species such a s th e mistlet oe a n

,
d d odde r
( C h a p X
. V I I I ) speci a ll y a d a p ted r oo ts se rve to a b s orb
nut ritive m a te ri a ls from th e h ost pl a nt Inm a ny l ower .

species of pl a nts such a s fu ngi a nd b a cte ri a th e i ndivid

, ,

u a l is fitted to dec omp o se a n d a bs orb org a nic sub

st a nces S till oth e r species ra nging from rel a tively simple
.
,

to highly c omplex t ypes a re a d a p ted to a n a qu a tic life

,

where delic a te ti ssues few if a ny roots but a h ighly de

, ,

vel op ed a bs orbi n g b ody su rf a ce a re th e m ore ge ne ra l

a d a p t a ti on s .

I N S E CTI V OR OU S P LA N T S S eve ra l species of fl owe ri ng

.

pl a nts h a ve bee ndesc ribed which a dd to thei r cust om a ry

f ood supply th e d igested rem a i ns of a ni m a ls ch iefly i n ,

sects a nd i nthis wa y a re a d a pted to a life i n s oils p oor

,

i nnit roge n In th e pitch e r pl a nt which a ff ords a g ood

.
,

ex a mple the le a ves a re sle nde r v a se sh a ped st ructu res

,
-

wh ose rims secrete a sweet fluid which sl owly oozes d own

thei r oute r su rf a ces In th e i nte ri or immedi a tely bel ow
.
,

the ri m is a sm ooth slippe ry z one ; be ne a th this the re

is a sec ond z one p rovided with i nw a rdly di rected h a i rs ;
436 T HE R E S ULT S O F E V OLU TI O N

wh ile a t the b o tt om o f the u rnor pitche r i s a fl uid con

t a ini ng a digestive e nzym I f a n insec t a tt ra cted by .
,

th e nect a r a ttemp ts to c ra wl i nto th e u rni t Slips onth e

,

sm oo th z on e fa lls i nt o th e liquid a nd u na ble to esc a pe

, ,

by re a s ono f th e h a i rs it d row ns is digested a nd a bs orb ed , , .

Th e V e nus fly t ra p of the s outhe a ste rnp a rt O f the U nited

-

S t a tes is a no the r o f th ese rem a rk a ble i nsect t ra ppi ng -

species Th e te rmi na l p orti on O f e a ch o f its le a ves is

.

devel op ed i nt o two fla t highly se nsitive l obes with nume r ,

o u s h a ir s l o c a ted a ro u nd thei r m a rgi ns Whe na n i nsect .

set tles up on this p orti on o f th e le a f th e l obes cl ose

like th e j a ws of a steel t ra p a nd th e c a ptive is digested .

Th is ope ra ti onove r the l obes a g a i nope na nd the p rocess ,

is repe a ted .

Th is cu ri ous h a bit o f e ns na ri ng a nd digest ing i nsects is

c omm on to seve ra l o the r species o ccu rri ng i n th is a nd
o the r c o u n t ries Th e i nte rested stude nt wi ll fi nd th e sub
.

je c t e nte rt a i n i ngly discussed i n D a rwi ns I nsectiv orous ’

Pl a nts ” .

ADAPTAT I O N S FOR S EL F D EFE N S E — Anim a ls wh en in -

d a nge r o f a tt a ck t a ke fligh t a nd esc a p e by vi rtue o f fleet »

ness or th e a bili ty to h ide or th ey ma y defe nd th emselves ,

by me a ns of cl a ws h orns teeth a nd v a ri ou s o the r org a ns

, , ,
.

Th e ra bbit for ex a mple whe ni nd a nge r d a sh es i nt o its

, , ,

bu rro w v a ri ou s species o f bi rd s esc a pe i nt o thick veget a

,

ti on ce rt a i n fish es bu ry th emselves i n th e mud while

, ,

h o sts o f i nsects spide rs a nd c ra bs c once a l themselves

, ,

u nde r sticks a nd st ones a nd i no the r s a fe ret re a ts The re .

a re a ls o m a n y species wh ose p rotective c ove ri ng re nde rs

th em c omp a ra tively f ree f rom a t ta ck Th e S pi nes of .

m a ny c ra bs fish es p orcupi nes th e a rm or O f th e a rm a

, , ,

dill o the shells O f cl a ms a nd s na ils a re f a mi li a r ex a mples

, ,

of this In seve ra l species a n a dditi on

. al p rotecti on
is a ff ord ed by p o is on wh ich i n s ome o f th e sea u rch i ns ,

an d fishes is situ a ted i n sm a ll c a vities i n o r ne a r the

spi nes P ois on s a c s a re a ls o a ch a ra cte ristic fe a tu re of
.

m a ny s na kes a ll o f th e spide rs ce ntipedes th e sti ngs o f

, , ,
 D E FE N S E OF Y O UNG 4 37

m a ny i nsects a nd th e sti ngi ng cells of th e c ora ls j elly

, ,

fishes a nd th ei r nume ro us rel a tives I n seve ra l o f th e

,
.

fishes (for ex a mple ce rt a i nof the eels a nd ra ys ) the re a re

,

elect ric org a ns sufficie ntly p owe rful i na l a rge i ndividu a l

to p a ra lyze a ma ntemp ora rily Th e devil fi sh es squids .
-
, ,

an d rel a ted species th row ou t a ni nky cl oud wh e nm olested

an d esc a p e a mid th e gl oom F rom such ex a mples it is
.

bu t a sh ort step to th ose c a ses whe re p rotecti onis a ff orded

to a n a nim a l c ove red with f oreig n subst a nces a s de ,

sc ribed i n th e secti on on P rotective R esembl a nces .

D E F E N S E OF Y O U N G — Ad a pt a ti ons u sed i n th e de
fense of the y o u ng a re i n m ost i n st a nces the s a me a s
th ose se rvi ng for self defe nse -
.

S F D EFE N S E N P A N T S
EL -
I L — M a ny species of pl a nts
.
,

a s well a s n ume rous fishes sh rimps a nd o the r p rolific,

an im a ls pe rsist by S hee r f orce of numbe rs If a g ra ss

,
.

pl a nt or SO c a lle d weed is dest royed by s ome e nemy

m
-
,

enough rem a i nto m a i nt a i nth e ra ce Innu e rous o the r .

species th e p rese nce of th orns on stem or le a f r e nders

them c omp a ra tively f ree f rom a tt a ck This is especi a lly .

t rue of dese rt pl a nts whe re a mid seve re c onditi ons th e

, ,

numbe r O f i ndividu a ls is rel a tively sm a ll a nd a d a p t a

ti ons a l ong th e li ne of self defe nse a re of sup reme imp or
-

t a nce In th e s a me or m ore f a v ora ble situ a ti ons ce rt a in

.

o the r species a re c o ve red with stiff p rickles or with sti n g ,

in g cells a s i n
,
the c a se O f th e nettle o r a s i nthe J a ck l n , ,
-

th e pulpit a re p a cked with needle —

-
,
like c ryst a ls which i n
su re them a g a i nst the a tt a cks O f i nsects s na ils a nd o the r
i ma ls S eve ra l species su ch a s th e ox a lis devel op
, ,

an
“

.
, ,

a cids i nthei r tissues while m a n y o the rs f orm u nple a s a nt

,

an d eve n p o is ono u s subst a nces which p rob a bly e na ble

such individu a ls to esc a pe c omplete dest ructi on
A DAPTATI O N S FOR R I V A LR Y B a ttles a m ong th e m a le
.

i ndividu a ls of a species for the p ossessi on O f a fem a le

Occu r i np ra ctic a lly eve ry cl a ss O f a n im a ls f rom th e c ra bs
an d i nsects to ma n Fu rthe rmore it p rob a bly is self
.
,

evident th a t th e org a ns a d a pted for defe nce a re chiefly

.
4 38 T HE R E S UL T S OF E V OLUTI O N

c once rned in this m a tte r O f riv a lry Am ong ce rt a i n O f .

th e fish es such a s th e s a lm on t ro u t a n
,
d stickleb a ck
, , ,

the re a re c omb a ts ra ngi ng f rom sudde nrushes with out ,

an y m a t e ri a l d a m a ge r esul ti n g to bi t te rly c onte s ted

,

s truggles wh e re de a th ma y resul t teeth bei ng th e org a ns ,

ch iefly c once rned M a ny o f th e tu rtles li z a rds a nd c roc o

.
, ,

diles a ls o e ng a ge i nlively tilts d u ring the b reeding se a s on ,

using tee th cl a ws or be a ks a s we a p ons Am ong th e bird s

, ,
.

th e re a re m a ny species chiefly th ose rel a ted to th e

,

phe a s a nts a nd d omestic f owl whe re spu rs a nd wings a nd ,

be a ks a re a t times used with de a dly e ff ect Am ong th e .

hi ghe r a nim a ls especi a lly th ose wh e re one m a le st rives

,

to g a i n p o ssessi on of seve ra l fem a les th e st ruggles a re ,

eve n m ore seve re Th is is especi a ll y t rue of ce rt a i n

.

species of wild sh eep a nd swi ne a nd of s ome of th e dee r

an d a ntel op e t ribe wh e re teeth a n ,
d h oofs l a y th e v a n
q u i sh ed l ow o r d r ive h im f ro m th e field .

R I VA LR Y A M O N G P LA N T S I f a ny one is i nd oubt as to
.

th e existe nce O f riv a l ry a m ong pl a nts let h im repe a t th e

exp e rime nt O f D a rwi n i nwh ich a sma ll p l ot o f gr ou nd
'

wa s cle a red a nd a ce nsu s t a ke n o f th e l a rge numbe r o f

i ndividu al pl a nts which ge rmi na ted a nd of the sm a ll
numbe r wh ich c a me to m a tu rity A s inth e c a se of a ni .

m a ls th e st ruggle for sup rem a cy is m ost seve re a m ong th e

membe rs of th e s a me species so th a t th rough ou t its ra nge ,

m a ny m ore i ndividu al s a re b ornth a na re a ble to live a nd

rep roduce . Bu t while it i s t rue th a t only a limited nu m
b er o f O ffsp ri ng o f a si ngle species ca nlive inone l oc a l i ty ,

th is d oes no t p reve nt membe rs o f othe r species f rom flo u r

i sh i n g i n th e s a me regi o n A gl a n.ce a t th e veget a ti o n o f
an y field o r f o rest is sufficie n t to p ro ve this f a c t I n .

eithe r eve nt h oweve r wh eth e r th e riv a l ry is betwee nth e

, ,

i ndivid u a ls O f th e s a me or di ff ere nt species th e ill a d a p ted ,

-

die i nth e st ruggle J ust wh a t a d a p t a ti ons a re the m o st

.

v a lu a ble a ids i n a ch ievi ng success is difficult to decide .

Obvi o usly a f a v ora ble l oc a ti on is o f p rime imp ort a n

,
ce ,

an d a vig o rous c on stitu ti onis sc a rcely l ess so I t is a l s o .

 ADA P TATI O N S I N ANI M AL S 4 39

ve ry ce rt a i n th a t mut a ti ons resulti ng i n new a nd m ore

f a v ora ble a d a p t a ti ons or i nth e i nc re a sed v a lue o f a l re a dy
existi ng ones a re of the high est imp ort a nce i np e rpetu a
,

ti on Of th e ra ce .

AD J U S T M E N T A DAPTATI ON S I N A N I M A LS In a b ro a d .

se nse eve ry a d a pt a ti on wh ich e na bles a n org a nism to

m a i nt a in itself beget y ou ng a nd i n s ome i nst a nces to
, ,

c a re for them is i n th e na tu re O f a n a dj ustme nt to the

,

e nvi ronme nt Th e gills a nd fi ns o f a qu a tic a nim a ls the

.
,

lu ngs a nd legs o f l and dwelle rs the fu r sh ells spi nes , , , ,

sc a les a nd othe r p rotective st ructu res diggi ng climbi ng

, , , ,

an d flyi ng org a ns a nd i nnume ra ble o the r a d a p t a ti ons a re

,

i neve ry i nst a nce of se rvice to th e species i n a dj usti ng it

to th e su rro u n di ng c onditi ons Inm a ny regi ons a na ni .

m a l s su rrou ndi ngs are subj ect to m a rked ch a nges with

’

pe ri ods of h igh a nd l ow tempe ra tu re a lte rna ti ng In .

‘

such a l oc a ti onm a ny a nim a ls u nde rg o a wi nte r or sum

mer sleep (Ch a p XX I I ) Wh e re the c onditi ons a re less
. .

seve re a c onside ra ble p rop orti ono f these s a me species a s ,

well a s nume rous o the rs a re a ctive th rough ou t th e ye a r

,
.

At th e s a me time this l a st na med cl a ss usu a lly displ a ys

di ffe re nt deg rees of a ctivity i nresp onse to da y a nd night
c onditi ons Th e gre a te r numbe r of a nim a ls a re a ctive
.

d u ri ng th e d a ytime a nd sleep or rem a i n c omp a ra tively

,

quiet a t nigh t ; oth e r species reve rse the p ro cess Inb rief .
,

eve ry species displ a ys a ch a ra cteri sti c seri es of a ctivities

i n resp onse to he a t ligh t c old or moi stu re or to th e
, , ,
'

na tu re O f its f ood supply a nd is c onti nu a lly a dj usti ng ,

itself to th e ch a ngi ng c onditi ons of its e nvi ronme nt .

P LA N T A D J U S T M E N T S N ot only a re pl a nts a nd a ni
.

m a ls a d a pted to live i n a defi nite t e rrit ory but a ls o , ,

withi nthis ra nge they a re c onti nu a lly u nde rg oi ng minor

,

ch a nges which se rve to a dj ust them to shifti ng c ondi

ti ons M os t pl a nts i n m ode ra te ligh t exp o se to the
.
, ,

fullest p ossible exte nt thei r le a ves a nd fl owe rs to th e

su nlight a nd s ome eve n shift thei r p ositi ons th rough
,

ou t th e da y a s the di recti on o f the ligh t ch a n ges On .

440 T HE R E S ULT S O F E V O LU TI O N

the o the r h a nd wh e re th e ligh t a nd h e a t a re i nte nse

, ,

a s in the t ro pics o r dese rts the le a ves O f m a ny pl a nts

,

cl ose a nd be nd d ow nw a rd th us esc a pi ng th e d a nge r o f

,

excessive illumi na ti on a nd ev a p o ra ti on O th e r pl a nts .

u nde rg o s omewh a t sim il a r noc tu rna l m oveme nts a nd

”

p reve nt a nu ndue l oss o f h e a t .

A nothe r ph a se o f a d j ustme nt a ppe a rs in the c a se O f

th ose pl a nts c a p a ble o f existi ng a mid m a rkedly diffe re nt
su rro u ndings I ndividu a ls eve n f ro m the s a me p a re nt
.
, ,

if g row n in ve ry m o ist h a bit a ts diffe r m a te ri a lly f rom ,

th ose i nc omp a ra tively dry s oil Th ose o f c old elev a ted .

regi on s a re decidedly u nlike thei r b roth e rs o f th e l owl a nds .

An d fu rthe rm ore while th ese pl a nts o f the m o u nt a i ntop

,

o r c old c ou n t ries die d ow n i n the f a ll the ve ry s a me ,

pl a nts i n w a rme r regi ons i n m a ny i nst a nces ma y li ve

, ,

f or Seve ra l ye a rs .

E v ery p l a nt a nd eve ry o rg a n o f th e pl a nt is th us a
d elic a tely a d j usted mech a nism u nde r th e sw a y o f m a ny
c ont rolli ng i nflue nces He a t ligh t m o istu re s oil c ond i
.
, , ,

ti ons g ra vity etc a re c onti nu a lly stimul a ti ng the livi ng

, ,
.
,

m a te ri a l wh ich resp ond s by m oldi ng its le a ves stems or

, , ,

roo ts to h a rm o nize with or fit i nt o th e shifti ng c onditi ons .

CO LOR S O F A N I M A LS One o f th e m ost u nive rs a l fe a t

.

u res o f livi ng thi ngs i s th ei r c ol or In m a ny i nst a nces .

this O bvi ously is a n a d a p t a ti o n; i n ce rt a i n o th e r c a ses

its sig nific a nce a t th e p rese nt t ime is by no me a ns
cle a r The red ness o f th e bl ood for ex a mple is
.
, ,

seemi ngly a n i ncide nt a l resul t o f its ch emic a l com

p ositi o n a nd is of no m ore u se to th e org a nism th a n
,

c ol o r is to a b rick or a te n ce nt piece O n th e oth e r .

h a nd the re a re ve ry m a ny S pecies whe re th e c ol or p a tte rn

,

is believed to be o f th e h ighest v a lue i nthe st ruggle for

existe nce I t ma y se rve so to ble nd a na nim a l with its
.

su rro u ndi ngs th a t i t esc a pes th e no ti ce o f e nemies or i t ,

ma y e na ble it to ste a l u nobse rved up o nits p rey Ino th e r .

an im a ls it a pp a re ntly is o f se rvice i n givi ng w a rni ng to

f rie nd s or i n still o th e r species to f oes And fina lly it
, ,
.
 P R O T E C TIV E R E S E M B LAN CE 4 41

ma y se rve to reflect or a bs orb he a t or p rotect th e tissues

o f th e b o dy f rom i n j u ri ous e ffects O f th e su ns ra ys A few ’
.

ex a mples of these v a ri ous cl a sses o f a nim a l c ol ora ti onwill

illust ra te the v a ri o us w a ys whe reby it se rves to a d a p t a
’

Species to its su rro u n di ngs a nd th e st ruggle for existence .

PR OTE CT I VE R ES E M B LA N CE Th e c ol or p a tte rnO f this

.

cl a ss of a d a p t a ti ons e na bles a na nim a l to esc a pe recogni '

ti ononth e p a rt o f its e nemies I t ma y be of a ge ne ra l.

ch a ra cter (Fig . an d h a rm onize th e a nim a l with the

FI G . 1 22 — Th e mmon E a stern qu a il
co , or B ob - whi te, Coli nu s
vi rgi nia nus .
(P h oto gra ph by J R S l.ona k er )
. .

Kell o gg a nd Doa n e E con Z ool

,
.Hol t . .

c ol or a s a wh ole O f th e su rrou ndi ngs ; or i t ma y be m ore

specific a nd c a use th e org a nis mto resemble a le a f pebble
, ,

, ,

or o s me o the r r el a tively sm a ll O bj ect T o th e fi rst cl a ss .

b el ong su ch a nima l s a s th e p ol a r be a r a rctic fox a nd h a re

'

, ,

wh ose wh ite c oa ts ble nd with th e snow ; or th e b row n

t ones O f dese rt a nim a ls wh ich h a rm onize with s a ndy
w a stes ; or the g ree ns g ra ys a nd o the r ti nts O f m a ny
, ,
 4 42 T HE R E S UL T S O F E VO LUTI O N

insects spide rs liz a rds a nd bi rds livi ng a m ong t rees

, , , , ,

g ra ss a nd othe r pl a nts
, .

I nth e sec on d cl a ss the c ol or resembl a nce o f a na nim a l

to a p a rticul a r O bj ect is usu a lly h eigh te n ed by a simil a r
ity o f f orm Th e f a mili a r w a lki ng stick i nsect a nd the
.
-
,

c a te rp il l a rs O f m a ny species O f butte rflies a nd m oth s

cl osely resemble th e twigs on wh ich they feed A con .

si dera bl e n umbe r O f butte rflies k a tydids a nd o the r i nsects

, ,

be a r a rem a rk a ble resembl a nce to le a ves d uplic a ti ng ,

eve nth e w orm h oles a nd fu ngus sp o ts N ume rous oth e r .

i nsects cl osely resemble th orns g ra i ns o f s a nd bits o f , ,

e a rth a nd o the r Obj ects E ve ry obse rv a nt stude nt of

.

na tu re ca na dd othe r ex a mples to th e list of this wide

'

Sp re a d type O f a d a pt a ti on .

S eve ra l a nim a ls a re c a p a ble O f a lte ring thei r c ol or with

a ch a n ge i nthe su rrou ndings D evil fish es squids m a ny
.
, ,

fish es t ree fro gs a nd liz a rds a re ex a mples O f th is well

,
-
,

k now n cl a ss Fi na lly th ere a re o th e r Species wh ich g a i n

.

a dditi on a l p ro tecti onby c o ve ri n g th emselves with obj ects

u na ppetizi ng to th ei r e nemies S eve ra l c ra bs for ex .
,

a mple ,
a tt a ch se a weeds to th ei r b a cks and th us co n ,

cea l ed esc a pe d a n
,
ge r O th e r species t ra vel a b ou t with
.

cl a m sh ells h eld ove r th ei r b a cks In a l l of th ese a nd .

th e o th e r c a ses o f p rotective resembl a nce i t is di fficul t

for h um a nsigh t to rec og nize th e p rese nce o f such a n im a ls ,

an d it a ppe a rs to be a n equ a lly a ssu red f a ct th a t th ei r

e nemies a re likely to ove rl ook them .

CO OR S OF H S I OLO G C
L P Y I A L VA L U E — Th e re a re m a ny
an im a ls wh ose b odies be a r c ol or c oa ts wh ich ma y or ma y -

no t be p rotective i nth e se nse th a t th ey c once a l th em f rom

e nemies bu t wh ich neve rtheless ma y se rve to p ro tect
,

th em from th e i nj u ri ou s eff ects o f th e su ns ra ys M a ny ’

.

subte rra ne a n a nd w ood b ori ng a nim a ls eith e r th rough

-

o u t life o r i na nimm a tu re c on diti on a re with ou t pigme nt ,

.

Inth e c a se O f the te rmites or wh ite a nts wh ich se rve a s

, ,

a g o o d illust ra ti o n the wi n ged o r rep roductive i ndivid u a ls

,

a re c ol orless up to th e time whe nth ey a re a b o u t to le a ve

 W A R NING COL O R ATI O N 4 43

th e nest to f ou nd new c ol onies Th e c ol or which the n .

devel ops i n th e ski n d oes not p rotect th em a g a i nst t h e

a tt a cks of e n emies bu t it d oes a ppe a r to sc ree nthe inte r
,

na l org a ns a g a inst su nbu rn or th e i nj u ri ous e ffects o f

ligh t T a nis a temp ora ry p ro tective c oa t a nd th e b row n
.
,

an d yell ow pigments which a re pe rm a ne nt fe a tu res o f

the ski n of v a ri ous ra ces of men a re p rotective a d a p t a
ti ons of the s a me ch a ra cte r Inseve ra l species of a nim a ls
.

the ne rves a re m ore or less c o ve red with p igme nt while ,

i no th e rs a h e a vy pigme nted c o a t li nes th e visce ra l c a vity .

In m a n y Othe rwise t ra nsp a re nt m a ri ne org a nisms the

eyes a nd i n ce rt a i n species th e rep roductive org a ns a s
, , ,

well a re c oa ted with v a ri ously c ol ored subst a nces In

m
.
,

these a nd a ny o the r species i t is a ssumed th a t c ol or ,

sc ree ns a re p rotective or i n ce rt a i n i nst a nces they ma y

,

a bs orb or r eflec t th e ligh t a n d th us keep a n a nim a l

w a rm or c ool .

W AR N I N G COLOR ATI ON — M a ny a nim a ls a re so b ril

l i a ntly a nd c onsp icu o usly c ol ored th a t it a ppe a rs su rp ris
i ng th a t th ey h a ve esc a ped a nnihil a tio n on the p a rt o f
nume rous e nemies Th e expl a na ti on a t le a st i nce rt a i n
.
,

defi nitely est a blished i nst a nces is th a t the c ol or a dv er

.

tises the f a ct th a t they a re d a nge ro us ona cc ou nt O f sti ngs ,

p ois ons or a na use ous fl a v or For ex a mple th e re is a

, .
,

g orge ously bl ue a nd red frog i nN ic a ra gu a which fl a u nts

its c ol ors i n th e b rilli a nt s unsh ine c omp a ra tively s a fe
from a tt a ck M r Belt wh o c onducted th e expe rime nt
. .
, ,

st a tes th a t h e off e red one O f th ese f rogs to ducks a nd

chicke ns a l l of wh ich refused it excep t one y ou ng duck
,

wh ich sna tch ed i t up p romp tly d ropped it ,

an d we nt ,
“

a b ou t j e rki ng i ts h e a d a s if t ryi ng to th row Off s ome u n

ple a s a nt t a ste ” I t h a s fu rth e r b ee n sh ow n th a t th e
.

mem ory of such exp erie nces e ndu res for a ve ry c onside r
a ble pe ri od a n d th e V ictims c a nno t be i nduced to repe a t
,

th e expe rime nt .

Th e yell ow a nd bl a ck b a nded b odies o f h ornets a nd

w a sps a re a l so w a rni ng or sig na lli ng c ol ors remi ndi ng ,
444 T HE R E S U LT S O F E V OLUTI O N

th ei r e nemies o f d a nge r a h e a d Th e st ri king c ol ors o f

.

ce rt a in butte rflies bugs bee tles sku nks a nd o th e r a ni

, , ,

m a ls wi th u nple a s a nt odor or t a s te b ri ng sh a rply to th e ,

a tte n ti on of s ome w ould be a ss a il a nt th e pe na l ty of a n

-

a tt a ck I t ma y i ndeed h a ppe n th a t a n e nemy ma y de

.

s troy a w a rni ngly c ol o red a nim a l bu t i t is a ls o p rob a ble , ,

a s h a s bee np ro ve nf or a c on side ra ble numbe r o f species ,

th a t it wi ll ra rely repe a t th e ope ra ti on Th e dest ructi on

’

o f a few i n dividu a ls th u s i nsu res th e ra c e a g a inst con

ti n u ed a tt a cks a n d e na bles i t to pe rsist .

M I M I CRY I f w a rning c ol ora ti onis a v a lu a ble a d a pt a

ti onto one species of a nim a l it must foll ow th a t a nothe r
,

FI G 1 23. . M imi cry . A ,

B, b umble -
bee a nd w a sp mimi cked by
,

flies .

one wh ich resembles i t is likewise be nefi ted A t le a st .

th is is th e ge ne ra l expl a na ti on of th o se c a ses wh e re a
h a rmless a nd i no ff e nsive species m ore or less cl osely
duplic a tes a na u se ous one I nst a nces o f th i s type o f p ro
.

tecti v e c ol ora ti on a re nume rou s a m ong i nsects a nd a re ,

a ls o k now n to o ccu r i n seve ra l o th e r g ro ups For ex .

a mple the re a re seve ra l species of flies which so cl o sely

,

resemble h on ey bees or bumble bees th a t it requi res cl ose

- -

sc ruti ny to detect th e f ra ud (Fig S ome of the .

 FL O W E R S AND I N S E C T S 4 45

w a sps a re mimicked by beetles ; ce rt a i n c onspicu ously

m a rked beetles such a s the p ot a t o beetle a re mimicked
, ,

i ntu rnby g ra ssh oppe rs ; a well k n ow nle a f h oppe r i nsect

be a rs a st riki ng resembl a nce to the p owe rful a nd a ggres
sive le a f cu tti ng a nt ; a nd the re a re m a ny spide rs which
-

signific a ntly a re ve ry simil a r to the flies up onwhich th ey

feed Am ong the butte rflies th e re a re a ls o m a ny species
.

wh ich mimic dist a steful ones a nd a few which resemble ,

bees or w a sps .

F LOW ER S N D I N S CT S
A E — Alth ough pl a nts i n ge ne ra l
do not p ossess a s definite or a t le a st a s rem a rk a ble a d a pt
,

a ti on s a s ch a ra cte rize a n im a ls the re a re m a ny S pecies

,

which p rove a nexcepti onto the rule This is especi a ll y .

the c a se with pl a nts i nr el a ti on to i nsects As is well .

k now n the g re a t m a j ority of fl owe ri ng pl a nts p roduce

,

p ollen a nd ovules wh ose u ni on is necess ary to p roduce

seeds Th e sex cells ma y be th e p roduct o f th e s a me
.
-

i ndividu a l or m ore freque ntly O f d iff e re nt pl a nts Fur .

th ermore the p olle n ma y be c a rried by the wi n

,
d a s in ,

th e c a se of m o st f orest t rees a nd g ra s ses or it ma y be ,

t ra nsp orted by i nsects The wi nd p olli na ted fl owe rs a re

.
-

sm a ll sce ntless a nd i nc onspicu ously c ol ored

, ,
Th ose .

p olli na ted by i nsects a re l a rge r c onspicuo usly c ol ored

'

, ,

an d a re f reque ntly p rovided with nect a r a nd di sti nc

tive O d ors Th e p resumpti on i s th a t th e fl owe rs of the
.

l a tte r cl a ss a re th us a d a pted to a tt ra ct i nsects Th is .

h a s a ctu a lly bee n p roved to be th e c a se with m a n y

S pecies .

Inth e fi rst pl a ce i t ca n e a sily b e seen th a t bees flies , ,

butte rflies m oth s a nd even h ummi ng bi rd s i n thei r

, , ,

se a rch for nect a r bec ome m ore or less c oa ted with p ollen
,

a s they j o u rn ey from fl owe r to fl owe r I t h a s beencon .

elusively dem onst ra ted th a t du ri ng these V isits the p olle n

o f on e fl owe r is disl odged i n a n o th e r M ore ove r it h a s .
,

bee n expe riment al ly p roved th a t the h oney bee is espe -

ci a l l y a tt ra cted by c ol ors W a sps h a ve a ls o bee nS h own

.

to be a tt ra cted by c ol or a n d it is p rob a ble th a t m a n

,
y
4 46 THE R E S ULT S O F E V OLU TI O N

o th e r i nsects p osse ss a c ol or se nse a nd a re a tt ra cted by

fl owe rs in c onseque nce .

I t is a ls o p rob a ble th a t O d ors a tt ra ct i nsects This is

.

ce rt a inly th e c a se wi th seve ra l species O f pl a nts rel a ted

to th e c omm on sku n k c a bb a ge wh ose fl owe rs p o sse ss a
-
,

st rong Od or of dec a yi ng me a t a nd a re p olli na ted by mul

ti tudes o f flesh fli es wh ich c ra wl ove r th em a n
-
d eve n
dep o si t eggs Needless to st a te the i nsti ncts O f th e flies
.
,

i n this c a se a re f a ulty si nce the y ou n

,
g s oon st a rve to
de a th wh ile the pl a nt p rofits by th e decepti ona nd p ro
,

duces seeds .

In a dditi on to a llu ri n g c ol ors a nd sce nts th e re a re

m a ny mech a nic a l a d a pt a ti ons i n fl owe rs wh ich fur the r
in su re p olli na ti on Th e nect a r is u su a lly l oc a ted deep
.

with in th e fl owe r so th a t to r e a ch it the i nsect must

,

inevit a bly b rush a g a i nst th e p olle n In the D utchm a ns

.
’

pipe a nd rel a ted pl a nts the re a re i nw a rdly di rected h a irs

,

which p reve nt th e esc a p e of th e vi siti ng flies u nt i l th ey

h a ve c oa ted themselves with p olle n With th e withe ring
.

o f th e h a i rs th e c a p ti ves esc a p e to a no th e r bl oss om The .

fl owe rs of th e s a ge a re so a d a p ted th a t th e st a me ns a re
c a used by th e e nte ri ng i nsect to be nd d ow n a nd da b a
m a ss O f p olle nonits b a ck In the orch ids a d a pt a ti ons
.
,

o f this cl a ss re a ch a h igh deg ree o f devel opme n t In .

a dditi on to a tt ra ctively c ol ored f oo tp a th s up which the

insec t is i nvited to w a lk th e re a re delic a tely a d j usted

,

c ont riv a nces for a tt a ch ing p olle n to th e visit or or for

a ctu a lly disch a rgi n g it a t its h e a d Th e d et a ils of the
.

p rocess a l ong with m a ny o th er ex a mpl es a re give n i n

,

D a rwi ns w ork onth e Fe rtili z a ti onO f Orchids

’ “
,
” .
 SECT ION 1 1

THE D I ST R IB U TI ON OF OR GA N I SM S
 C HAPTER XXXVII

PL AN T D I S T R I B UTI ON
“

TH E F A CT S OF D I S TRI B UTI O N Almost every one h a s

.
-

t ra velled f a r e nough to re a lize th a t the a ssembl a ges of

pl a nts a nd a nim a ls i n di ff e rent pl a ces a re m ore or le ss
di ff e re nt The ch ief u nde rlyi ng c a uses of this f a ct b e
.

c ome m ore obvi ous wh e ndist a nt a re a s a re c onside red a s , ,

for ex a mple N orth Am e ric a S o uth Ame ri c a Asi a a n

, , , ,
d
Afric a . I f one we re to fi nd a n a re a i n e a ch of these
c ontine nts in which th e c onditi ons o f s oil a nd clim a te
a re so n e a rly th e s a me th a t the s a me pl a nts ca nbe culti
v a ted equ a lly well i na l l h e w o uld n
,
eve rtheless fi nd th a t
th e pl a nts na tive i n e a ch a re a di ffe r from th ose i n a ll
th e oth e rs bu t th a t th e re exists a l a rge me a su re of supe r
,

fi ci a l rese mbl a n ce i n th e ge ne ra l a ppe a ra nce or a spect O f

the a ssembl a ges of pl a nts Whe n the i ndividu al pl a nts
.

a re c on side red it i s f o u nd th a t the d iff e re nce is much

gre a te r th a n the ge ne ra l a pp e a ra nce i n the m a ss w ould
suggest From these obse rv a ti ons we ma y deduce two
.

c onclusi ons : fi rst th a t me re dist a nce h a s s omethi ng to

do with th e dist ribu ti on o f pl a nts a nd sec ond th a t
simil a rity of h a bit a t h as a g ood de a l to do with S imil a r
ity of a pp e a ra nce i n th e m a ss For ex a mple if one of
.
,

th e supp osed a re a s is a f orested one it is quite ce rt a i n

th a t a ll the o th ers will be a ls o .

T H E G E N ER A L CAU S E S OF D I S TR IB UT I O N a re a l re a dy
f a mili a r to the re a de r for h e k nows th a t org a nisms u nde r
g o mut a ti ons or h e r it a ble v a ri a ti on s a n d th a t they a re

a ble to S p re a d th emselves ove r c on side ra ble a re a s eith er

bec a use th ey th emselves a re m o tile or bec a use their seeds
or sp ores ca n be c a rried by wi n d w a te r or a nim a ls
, ,
.

4 49
4 50 P LAN T DI S T R I B UTI O N
M ore ove r h e is a w a re th a t a l l org a nisms h a ve v a ri ous
,

me a ns O f a dj usti ng th emselves to thei r e nvi ronme nt .

These th ree f a ct ors a c cou nt i n a ge ne ra l wa y for the .

fu nd a me nt a l f a cts o f dist ributi on na mely : (1 ) th a t

,

the re a re diff e re nt a ssembl a ges O f org a nisms i n d ist a nt

regi on s (2) th a t species a re m ore or less widely di stri
,

buted ove r the w orld a nd (3 ) th a t pl a nts a nd a nim a ls

,

a re a gg reg a ted in t o g roups a d a pted to live u nde r simil a r

e nvi ronme nt a l c onditi ons E a ch of these f a c t ors may
.

now receive a s omewh a t fulle r c onside ra ti on in orde r

to see j ust h ow they a ctu a lly w o rk ou t .

EV OLUT O N AN D D S TR B UT O N
I I I I — Wh e n new sp ecies
.

origin a te by ev oluti on i t is p rob a ble th a t i n th e v a st

m a j ority of c a ses they do so i n one p a rticul a r l o c a lity

on ly F rom this th ey sp re ad m ore or less widely a cc ord
.

in g to ci rcumst a nces I.nth e g re a t m a j ority O f c a ses they

fin d some thi ng to p reve nt th eir S p re a d ove r th e whole
w orld f rom which i t foll ows na tu ra lly th a t ev oluti on
,

te nds to c re a te dive rsity a m ong th e pl a nts a nd a nim a ls

O f widely sep a ra ted regi o n s I t ca nbe re a d ily see nth a t
.

this w ould be t rue eve n if a ll p a rts O f the e a rth h a d

bee noriginally p opul a ted with ex a ctly th e s a me species ;
ev oluti on of new a nd di ff e re nt species i nregi ons dist a nt
f rom one a nothe r w ould i nth e c ou rse of time h a ve given
them widely diffe re nt a ssembl a ges of pl a nts a nd a nim a ls .

M I GRATI O N AN D D I S TR I B UT I O N — Wh e n new species

‘

o f pl a n ts a nd a nim a ls h a ve once ev olved th ey sp re a d

gra d u a lly aw a y f ro m th ei r pl a ce of origi n S ome s orts o f .

an im a ls a nd pl a nts a re a bl e to sp re a d ra pidly a s for , ,

ex a mple bi rds a nd d a ndeli ons Onthe o the r h a nd a ngle

,
.

w orms a nd pl a nts with l a rge h e a vy seeds h a ve i nna tu re

ve ry much less efficie nt me a ns of dispe rs a l F rom ill us .

tra ti o n s O f th is s ort it is e a sy to u n de rst a nd th a t th e

me a ns o f dispe rs a l g ove rni ns ome me a s u re th e p rob a b i l
i ty o f a p a rticul a r org a nism sp re a di ng to th e li mits of its
p oss ible ra nge H oweve r i t must be noted th a t eve n
.
,

th e org a nisms with th e le a st efficie nt me a ns o f dispe rs a l

 ADA P TATI O N AND DI S T R I B UTI O N 4 51

w ould ultim a tely occupy a ll suit a ble a re a s u nless the re

we re s ome a ctu a l b a rrie r to p reve nt thei r re a ch ing them .

ADAPTATI O N AN D D I S TR I B UT I O N — A no the r f a c t O f
.

g re a t signific a nce i s th a t pl a nts a nd a nim a ls do not oc

cup y a ll the a re a s wh ich they ca nre a d ily r e a ch A study .

of a n y a re a o f a few squ a re miles will o rdi na r ily S h o w

th a t th e re a re seve ra l diffe re nt a ssembl a ges of pl a nts
with init ea ch occu p y i ng a h a bit a t o f a speci a l s ort Th e
,
.

p l a nts i ne a ch o f these f or th e m ost p a rt c ould re a dily

, ,

re a ch a n y o f th e o the rs a n d l ikewis e i n m ost c a ses th ey

,

do so but do no t est a blish th emselves bec a use the eu ,

vi ron ment i s not su it a ble For illus t ra ti on it is not

.
,

u nu su a l to fi nd a l a ke a st re a m a pl a in a nd a m ou nt a i n
, , ,

wi th incl ose p roximity bu t with ou t a ny te nde ncy for th e

p l a nts of th e l a ke to sp re a d to the m ou nt a i ntop Wh ile .

the m ou nt a i n a nd th e p l a i n a re not so di ff e re nt a s th e
m ou nt a i na nd th e l a ke yet e a ch is likely to h a v e a dis
,

ti n cti v e a ssembl a ge o f pl a n ts a nd to Sh ow little te nde ncy

for th is diff e re nc e to b e lesse ned by migra ti on f rom one
h a bit a t to th e oth e r This s ort of phe nome non sh ows
.

th a t s ome p l a nts a re a d a pted to liv e u nde r one set of

c onditi ons a nd not u nde r othe rs Th e s a me th i ng is .

t rue of a nim a ls but to a lesse r exte nt ona cc ou nt of thei r

-

gre a te r i ndepende nce o f certa i n fe a tu res o f th e e nvi ron

ment A p ol a r b ea r w o uld p rob a bly find no i nsupe ra ble
o

p h ysi ca l obst a cle to p reve n t h is migra ti on from Al a sk a

to P a na m a bu t h e w ould a lm ost ce r t a i n ly fi nd th e c ondi
ti ons th e re so little a d a pted to h is fu r cl a d b ody a nd -

h a bits O f life th a t h e w ould le a d a sa d life of no l ong

du ra ti on. N or w ould th e c oug a r o f th e M exic a n j u ngle
find h imself bette r situ a ted sh o uld h e p e rch a nce mig ra te
’
to th e p ol a r be a r s a rc tic h ome .

F LOR A S AN D FAU N A S Inorde r to e n

. a ble u s to spe a k

a ccu ra tely th e bi ol o gist h a s c o i ned two w ords to exp ress

th e a ssembl a ge of p l a nts or a nim a ls ch a ra cte ristic of a
p a rticul a r regi on All th e pl a nts f ou nd na tive i nM a i ne
.

or Ca lif orn ,
i a th e U nited S t a tes or E u rop e c onstitu te
4 52 P LAN T DI S T R I B UT I O N
th e flora o f th a t regi on The a nim a ls na tive to a regi on
.

c onside red a l l t ogethe r c onstitute i ts fa u na A p a rticul a r

.

species of pl a nt or a nim a l ma y occ ur i nthe fl ora or f a u na

respectively o f m a n y regi ons a nd be a ss oci a ted with a
di ff e re nt set O f oth e r species i n e a ch This illust ra tes
.

the u nive rs a l f a c t th a t the ra nge of a ny p a rticul a r species

is distinctive of i t bu t ra rely c orresp onds a ccu ra tely with
th e ra nges o f o the r species with which i t ma y be
a ss oci a ted
—
.

V E GE TAT N
I O . Wh e none spe a ks of th e fl ora or f a u na
o f a regi on h e refe rs to th e a ctu a l S pecies i n h a biti ng it ,

bu t wh e nhe spe a ks o f a f orest p ra i rie or dese rt h e refe rs

, ,

not to the species c ons tituti ng this a ssembl a ge o f pl a nts

bu t to th e ge ne ra l a ppe a ra nce or a spect o f th e regi on .

A s a m a tte r O f f a c t ve ry di ff e re nt a s sembl a ges of species

ma y c ombi ne to p roduce f orests O f ve ry simil a r a ppe a r
an . n
ce Pi ne f orests occu r i M a ine M ich ig a n a nd Ca li
, ,

forni a for ex a mple which l ook much a like b u t wh ich

, ,

h a ve few species i nc omm on a nd no o ne species O f pi ne

,

is f ou nd i n a ll Any of th ese w ould h a ve still fewe r

.

species ide ntic a l with a p i ne f orest i n E u rope or Asi a

an d p rob a bly w ould not h a ve a si n gle species th a t c o uld
”
be f ou nd i n a p i ne f orest i n Chile ; in f a ct the so
“

c a lled pi ne f orest o f th e l a tte r w ould not h a ve eve na t rue

pi ne of a ny ki nd but me rely a no th e r c onife rous t ree
m ore or less resembli ng a pi ne One w ould fi nd th is s a me
.

p ri nciple to be t rue of a p ra i rie i nthe M i ssissippi V a lley

c omp a red with one i ns outh e rn B ra zil or o f th e dese rts ,

of ou r a rid S o uthwest a n d the S a h a ra .

Illust ra ti ons o f th is s ort migh t be multiplied to a lmost

an y ex te nt to illust ra te th e imp o rt a nt f a ct th a t s i m i l ar

ha bi ta ts i na n y p a r t o f th e w orld a re o ccu p i ed by a si m i
la r typ e of vege ta ti on but th a t th e species a re d iff e re n
,
t
an d i n ge n e ra l m ore diff e re nt th e fu rth e r rem oved f rom
on e an oth e r a re th e regi on s u nde r c onside ra ti on Th us .

i t is to be u nde rst ood th a t te rms su ch a s forest dese rt , ,

p ra i rie refe r to vege ta ti ontyp es with ou t refe re nce to the

BAR R IE R S TO UNIV E R S AL DI S T R I B UTI O N 4 53

p a rticul a r species th a t m a ke up th e pl a nt a ssembl a ges ,

wh ile flora a pplies to the a gg reg a te o f species with o ut

refe re n ce to the veget a ti on types which th ey ma y com
bi ne to p roduce .

T H E B A RR I ER S T O U N I V ERS A L D I S TR I B UT I O N
. I n th e
p receding t opics th e c onditi ons p osi ti vely c once rned i n
dive rsifyi ng th e a s s embl a ges o f pl a nts a nd a nim a l s i n
diffe re nt regi ons of the w orld h a ve bee n emph a sized a nd
on ly b a re menti on m a de O f ce rt a i n nega ti ve f a ct ors of
ex t reme imp ort a nce One ma y exp ress th e ide a i n th is
.

wa y : whe n pl a nts a nd a nim a ls ev olve i nt o new S pecies

these w ould sp re a d to a ll p a rts of th e e a r th u nless th e re
were s ome s ort of b a rrie r to st op th em or th e c onditi ons
of the e n vi ronme nt we re u nsuited to thei r h a bits O f life .

To pu t this p o sitively one ma y sa y th a t i n th e a bse nce

of b a rrie rs o f a n y ki n d a ll Species of pl a nts w o ul d ulti
m a tely sp re a d a ll ove r th e h a bit a ble w orld bu t w o uld be
rest ricted to th o se p a r ticul a r l oc a l a re a s or h a bi ta ts which

w o uld be suit a ble to e a ch As a result of this the re w ould

.

be much th e sa me sort of vege ta ti on typ es a s now exist ,

bu t the spec i es i n e a ch w o uld b e the s a m e a ll the world

o ver . Onth e o the r h a nd i f a l l pl a n
,
ts we re suited to th e
s a me e nvi ronme nt a l c onditi ons bu t th e p re se nt b a rrie rs
still existed th e re w o uld be a u ni form vege ta ti onwi thi n
“

ea ch ba rri er li mi t (E xcep t o f c o u rse f or the f a ct th a t th e

s a me b a rrie rs do not a pply to a l l species ) but the di ffer

,

en ces i n th e a ctu a l sp ecies n ow p rese n t i n e a ch g re a t

divisi on of th e w orld w ould rem a i n j ust a s they a re .

From c onside ra ti ons of this s ort it is Obvi ous th a t b a rrie rs

pl a y a n imp ort a nt rOl e in dete rmi ni ng th e Geogra p hi ca l
D is tri bu ti onO f species a nd it will be pe rt in ent to i nqui re
i nt o th e na tu re of b a rrie rs a nd to c onside r h ow they
ope ra te .

T A N GI B LE B ARR IER S a re l itera l l y th ose which ca n be

t ouched bu t the te rm is used p ra c tic a lly for a l l th o se o f
a m a te ri a l a n d nonlivi ng ch a ra cte r . T o a ny a nim a l
-

which ca n w a lk but c a nnot fly or S w i m w a te r c onsti

, ,
4 54 PLAN T DI S T R I B UTI O N
tu tes b a rrie r difficult but no t a lw a ys imp ossible to p a ss
a .

T o such a rive r ma y be a ve ry e ffi cie nt check to mig ra

,

ti on p rovided th a t i t is no t p ra ctic a ble to go rou nd its

h e a d w a te rs Anoce a nc ons titutes a p ra c tic a lly i mp a ssa
-
.

ble b a rrie r a rou nd a nisl a nd or a c onti ne nt to su ch a nim a ls

or to pl a n ts wh o se seeds c a nno t be c a rried by w i nd or
w a te r Fish es a nd st rong fly i ng bi rds on the c ont ra ry
.
-

a s well a s pl a n ts with thistle d ow nlike seeds re a d ily

- -

c ross ve ry l a rge b odies O f w a te r L a rge l a nd m a sses o f

.

c ou rse c onstitu te efficient b a rrie rs a g a i nst fishes Inf a ct .

i nth e I nte r M o u nt a inR egi ono f the U nited S t a tes the re

-

a re st re a m s which n ow l o se thei r w a te rs i n the dese rt

s a nds withi n a few miles o f e a ch o th e r a nd yet h a ve

diffe re nt species of fish i nthem a lth ough th ey we re b o th
,

c onnected i nth e no t dist a nt ge ol ogic a l p a st by the w a te rs

O f th e a n cie nt a nd now d ried u p l a ke i nt o wh i ch they
-

emptied A sid e f rom pl a nts with sm a ll seeds e a sily dis

.

p ersed by wi n d pl a nts i n gene ra l a re less ra pid i n ex

,

te ndi ng thei r range th a n th e h ighe r a nim a ls They too .

fin d oce a ns a nd h igh s n o w cl a d m o u n
-
t a i ns a nd such
like physic a l forms se ri o u s a nd s ometimes imp a s sa ble
b a rrie rs I t is not i nte nded th a t this p a ra gra ph S h ould
.

a ttemp t to list a ll th e s orts o f th i n gs which ma y c onsti

tute t a ngible b a rrie rs bu t me rely to i ndic a te th e f a ct by
,

illust ra ti onth a t th e re a re m a ny s orts of physic a l eleme nts

which c onstitute efficie nt b a rrie rs for p a rticul a r species
an d th a t wh a t i s a b a rrie r for one ma y no t be for a noth e r .

I N TA N GIBLE B ARR IERS I t is re a dily see nth a t pl a nts

.

O fte n f a il to sp re a d i n t o a d j a ce nt a re a s whe re the re a re

no t a ngible b a rrie rs to st op th em For illust ra ti o n th e .

pi ne t ree s of M i ch ig a n w o uld h a ve nei th e r m o u nt a i n ,

d ese rt nor oce a nto ch eck th ei r sp re a d s o u th th rough th e

,

p ra i rie regi on to th e G ulf o f M exic o bu t neve rth eless , ,

they do not do so N ei th e r do th e p ra i ries nor the i nd i

.

vidu a l species wh ich g row i nth em sp re a d north i nt o th e

f ores t c o u nt ry to a ny g re a t exte nt or west i nt o th e d rie r
pl a i ns a nd m o u nt a ins a lth ough m a ny spe cies h a ve seeds
,
 IN TANGI B L E B A R R I E R S 455
e dily t ra nsp orted for l ong dist a nces Endless ex a mples
r a .

o f this s a me s ort o f rest ra i n t o f species wi thi n limits

not est a blished by m a te ri a l b a rrie rs migh t be cited F rom .

this one ma y c onclude th a t they do not S p re a d bec a use

the c ondi ti ons of s oil a nd clim a te a re not suit a ble to them
bey ond the limits of th ei r na tu ra l ra nges . A c a reful
c onside ra ti onof the f a ct ors sh ows th a t wh e re no t a ngible
b a rrie rs e xist a nd the regi onh as a n equ a lly dist ributed
,

ra i n f a ll species sp re a d ove r a f a r g re a te r extent e a st a nd

,

west th a nnorth a nd s outh Th is suggests th a t tempe ra

.

tu re ma y be one O f th e c onditi oni ng f a c tors a supp o siti on

,

which ca n re a dily be ve rified expe rime nt a lly i n m a ny

c a ses There will thus be a te ndency for the w orld to be
.

divided i nt o tempe ra tu re belts or z ones e a ch of wh i ch is

'

a d a p ted to p a rticul a r species B u t S ince th e ra i nf a ll is

u nlikely to be u nif orm th rough ou t th e e a st a nd west
.

exte nt of such z ones th ey will a g a i n te nd to be divided

,

i nt o sm a lle r a re a s on th e ge ne ra l b a sis O f th e m oistu re

f a ct or of clim a te ; O f c ou rse it is well k now n to eve ry
-

on e th a t on a cc o u n t of th e p o siti on O f m o u nt a i ns a nd
l a rge b odies O f w a ter a nd th e di recti on O f th e wi nds ,

these z ones O f tempe ra tu re a re not st ra igh t h oriz ont a l

belts exte ndi ng u nif ormly a ro u nd th e e a rth but a re i n
f a ct be nt fa r to th e north or s outh wide ned ou t or g re a tly
,

na rrowed a cc ordi ng to th e v a ri ous p ositi ons of th ese

physic a l f a ct ors Inf a ct one ma y O ftengo s o uth a nd up
.

a m ou n t a i n a nd find th e s a me ch a nges i n tempe ra tu re

an d veget a ti on th a t h e w o uld fi nd i n g oi ng north ove r
a ve ry much l on ge r dist a nce on the level This is clue
.
,

o f c o u rse ,
to th e well k now n f a ct th a t tempe ra tu re
dec re a s es with a ltitude a s well a s with l a ti tude N or .

a re the divisi on s d epe nde nt on m oistu re a ny m ore regu

l a r S i nce ra i nfa ll a ls o usu a lly i nc re a ses with a ltitude
, ,

esp eci a lly onth e wi ndw a rd sl opes o f m o u nt a i ns Incon .

si deri n f a cts O f th is s or t it S h o uld no t be f org o tte nth a t

g
th ese di ffe re nces o f clim a te re a lly a ct a s condi ti ons for
f a v o ra b l e d e ve l op m e nt a nd o n ly o cc a si ona lly a s ge nuin e
456 P LAN T DI S T R I B U TI O N
b a rrie rs I t i s t rue th a t th e re a re nume ro us pl a nts i n
.

N orth Ame ric a which c o uld re a dily g row i nS o uth Ame r

i ca bu t do no t sp re a d f rom th e one pl a ce to the o the r b e
c a use th e w a rm m oist clim a te of th e i nte rve ni ng z ones ,

t ogethe r with th e pl a nts which i nh a bi t them m a ke i t ,

imp ossible to c ro ss .

B I O LO GI CA L B AR R I ER S In c on
. side ri ng th e t reeless
p ra i rie a nd th e a dj a ce nt f orested a re a s one migh t supp ose
th a t th ey a re dis ti n ct bec a use the species f o u nd i n e a ch
cou ld n o t live i n th e o the r bu t this is d a ily p ro ved u n t rue
by t h e th rivi ng g roves pl a nted a b ou t the f a rms of th e
p ra i rie st a tes a nd by th e me a d ows O f the f orest regi on .

Th e tru th — i n p a rt a t le a s t — seems to be th a t th e
,

pl a nts of one regi on ca n a ctu a lly grow i n the o the r i n

m a ny i nst a nces but they a re not so well a d a p ted to s o il
an d clim a te a s th e na tives a nd so a re u n a ble to co mp e te

su ccessfu l ly wi th them He re a g a i n i t is to be no ted

.

th a t it is not a questi on O f a ctu a l b a rrie rs bu t ra th e r

on e o f a f a ilu re i ncompetiti on du e to impe rfec t a d a pt a

ti on O f c ou rse a g ra ssy p ra i rie lyi ng be twee n two

.

simil a r forested a re a s w ould a ct a s a ba rri er to hi nde r

species sp re a di ng f rom one to the othe r ; for if they we re
u na ble to live i n th e g ra ss l a nd they w ould be u nlikely
-

to re a ch the oth e r a re a suit a ble to th ei r g rowth .

T H E G R EAT P LA N T R EA L M S —Wh e n th e w orld a s a

.

wh ole i s reg a rd ed on a b a sis o f th e p ri nciples set f orth

a b o ve a n d th e a c tu a l dist ribu ti ono f th e species of pl a nts
an d an im a ls is c onsid e red it is f o u nd th a t i t ca n be
,

divided i nt o five m a j or d ivisi o ns or R e a lms a s foll ows : ,

(1 ) Th e H ol a rctic R e a lm i ncludi ng th e wh ole O f Eu rope

, ,

m o st O f N o rth Ame ric a a nd Asi a a nd Af ric a north O f ,

th e S a h a ra D ese rt ; (2) Th e N eo t ropic a l R e a lm i n

-
clud ,

in g S outh Ame ric a ,

C e nt ra l Ame ric a a n d the West ,

I ndies ; (3 ) The Orie nt a l R e a lm i nclud i ng Asi a s outh ,

O f th e Him a l a y as ,
B orne o J a v a a nd th e Ph ilippi nes ;
, ,

(4 ) The A ust ra li a n R e a lm i ncludi ng Aust ra li a New

, ,

G ui ne a T a sm a ni a a nd seve ra l sm a lle r isl a nds ; (5 ) Th e

, ,
 T HE H OLA R C TI C R E AL M 7

E thi opi a n R e a lm , i ncludi ng Af ric a s o u th of the S a h a ra ,

M a d a g a sc a r, a nd p a rt of A ra bi a I t w o uld be neithe r
.

p ossible nor desi ra ble to set f orth i na b oo k of this ch a r

a cte r th e det a ils o f th e flora s or f a u n a s O f th ese g re a t

r e a lms The p u rp ose is ra the r to exhibit ge ne ra l bi ol ogi

.

ca l p ri nciples a nd me rely to illust ra te th em with sufficie nt

de ta il to m a ke them u nde rst a nd a ble This end ma y be .

a cc omplished by c on side ra ti on i n the rem a i ni n

, g p a ra

g ra ph s of th is ch a p te r o f th e chief veget a ti on types

,

f o u nd i na limited regi on such a s the U nited S t a tes a nd

Ca na d a a nd i n th e f oll owi ng ch a pte r a b rief p rese nt a
,

ti on Of th e ch a ra cte ristic a ni ma l sp eci es of th e v a ri ous

re a lms .

T H E H OLAR CTI C R EA L M c on stitutes a ne a rly c onti nu

ou s l a n d m a ss bei ng b roke n Only by th e ve ry na rrow
‘

B e ri ng S t ra it M ore ove r th e re is a b u nd a nt evide nce

.

th a t i n recent ge ol ogic tim e th e c onnecti ons h a ve bee n

f a r m ore exte nsive b o th i n th a t regi on a nd V i a G ree n
l a nd so th a t c ommu nic a ti ona nd dist ributi onwe re much
,

e a sier th a nnow This wa s f a cilit a ted a ls o by the w a rme r

.

clim a te i nth e f a r nor th a t th a t time H oweve r a con .

,

si dera bl e ge ol ogic time h a s el a psed S i n ce th e p rese nt i m

pe rfect c ommu nic a ti on wa s est a blish ed a nd th e clim a te
bec a me c olde r with the result th a t th e sp ecies wh ich
,

h a ve ev olved si nc e th a t time h a ve bee n m ore l a rgely

c onfi ned to th e c onti ne nt of th ei r origin Th e conse .

q u en ce is th a t N orth Ame ric a and E u r a si a resemble

e a ch oth e r fa r m ore th a neith e r of them resembles a ny
o th e r re a lm bu t still h a ve ch a ra cte ristic diffe re nces
, .

N orth Ame ric a ma y th e ref ore be th ough t o f a s one o f

th e v a st regi ons of th is e norm ou s re a lm Th e s a me p ri n .

ci p le s, h oweve r a pply eve rywh e re a nd ma y be illus

,

tra ted s a tisf a ct orily by c onside ri ng a still m ore limited

‘

regi on .

V EGETAT I O N OF T H E U N I TED STATES AN D CA N ADA (see

a cc omp a n yi ng ma p ) With i nthis a re a a re to be f ou nd
.

the p ri ncip a l veget a ti on types of th e Tempe ra te a nd

458 P LAN T DI S T R I B UTI O N
A rc tic z ones S ome or a l l o f th em a re fa mili a r to th e
.

re a de rs o f th is b o o k R efe re nce to th e a cc omp a n

. y i ng
ma p will Sh o w th e a re a s ove r wh ich the seve ra l veget a
ti ontyp es a re rep rese nted They ma y be e nume ra ted i n.

th e f oll owing b rief sy nO ps i s : (1 ) D ecidu ous F orest ,

o ccupyi n g m ost O f th e e a ste rnU nited S t a tes s o uth e a ste rn ,

Ca na d a p a rt s O f th e M issis si p pi V a lley a n
,
d exte n di ng ,

westw a rd a l ong th e st re a ms i nt o th e p ra i rie regi on .

(2) C onife ro us F orest o ccupyi ng two ra th e r distinc t but

,

c onnec ted a re a s th e e a ste rnone lyi ng north of the deci du

,

o u s f o rest ,
exte ndi ng north a s f a r a s s outhe rn L a b ra d or
an d northwestw a rd to j oi n with th e w este rn c onife ro us
f orest This weste rn a re a i ncludes fi rst th e e nti re R ocky
.

M o u nt a i nregionf rom Y uk on Te rrit ory s outh i nt o N e w

M exic o a nd Ar iz ona a nd sec ond th e m ou nta i n ra nges
,

lyi ng ne a r th e coa st from Al a sk a to s ou th e rn C a lif orni a

w

( )
3 Th e Tu n d r a o r th e B a rr e n G r o u nd n o rth o f th e limit ,

O f t rees a n d wh a t is ne a rly th e s a me th e Alpi ne veget a

, ,

ti ona b ove th e t ree li ne onth e h igh e r m ou nt a i nsummits

b oth e a st a nd west (4 ). Th e G ra ssl a nd s o ccupyi n g th e
i nte ri or o f the c onti ne nt betwee n th e decidu ous f o rests
an d th e R ocky M o u nt a i ns (5 ) The D ese rt occupyi ng
.

th e G re a t B a s i n a nd th e a rid pl a i ns a nd pl a te a us o f
th e s outhweste rnU nited S t a tes (6 ) M ixed
. C o nife ro us
an d D ecidu ous F orests o ccupyi n g th e s outh e a ste rnc oa st a l
pl a i nf rom N ew J e rsey to Fl orid a a nd west a l ong th e gulf
c oa st i nt o Tex a s (7 Ch a p a rra l a sh rubby growth
.
,

O f eve rgree npl a n ts occ u pyi ng th e footh ills o f th e S ie rra

N ev a d a a nd Coa st R a nge M ou nt a i ns of Ca lif orni a .

Ea ch of th ese l a rge regi o ns i s o f c o u rse s u scep ti , ,

ble of a na lysis i nt o sm a lle r a re a s e a ch o f wh ich h a s

its own ch a ra cte ristics For ex a mple th e s ou th e rn ex
.
,

te nsi onof th e Conife rous F orest h a s a f ri nge c ove red with

a d en se g rowth o f eve rg ree nb roa d le a ved pl a nts p roduc -

in g a vege t a ti o n type c a lled C h a p a rra l Th e ce n t ra l .

G ra s sl a nd a ls o na tu ra lly f a lls i nt o two p a rts o n th e b a sis

o f ra i n f a ll The e a s te rn p a rt o f it is fa r m oiste r a nd
.
rt

ti
ti
0!

Ct

fc

ly
 D E CID U O U S FOR E S T 4 59

c ove red with luxu ri a nt p ra i rie i nits na tu ra l st a te wh e re ,

a s th e so c a lled G re a t Pl a i ns i n th e weste rn p a rt h a ve
-

a much sp a rse r c ove ri n g O f bu nch g ra ss .

D E CI D U O U S F OR ES T (Fig — In this regi on the

.

clim a te is tempe ra te with p recipit a ti on well dist ributed

,

th rough th e ye a r Th e wi nte rs a re col d s now ofte nlies

.
'

on the g ro u n d for c onside ra ble p e ri ods a nd the g rou nd ,

FI G 1 24
. . Fore st of Whi te P in
th e m o st ch a ra cterist i c
e, o ne of

tree s of th e N ort hw e s tern C oniferous forest T he l ow er b ra nch es .

h a ve b een kill e d by th e dense sh a de ca s t by th e upper f oli a ge .

freezes to a c onside ra ble dep th All of these clim a tic con .

di ti ons ma ke it a regi onwhe re g rowth is g re a tly f a v ored

i nsumme r a nd a t a ve ry l ow ebb i nwi nte r Inge ne ra l .

th e s oil is fe rtile a nd s uited to th e g rowth of the h ighest

type O f veget a ti on Th e f orests c onsist of beech m a ple
.
, ,

oa k tulip t ree a n
, d m a ny oth e rs ch a ra cte rized by h a vi ng
,

b roa d th i n le a ves a ble to do a m a ximum a m ou nt o f

ph ot osynth esis du ri ng th e w a rm m oist summers but u nfit
to e n du re th e wi nte r (Fig Th e te rm decidu ou s re
.

fe rs to th e f a ct th a t th ey sh ed thei r le a ves on th e a p
p roa ch of wi nte r a nd thu s p rese nt only b a re b ra nches du r
in g th e i nclement se a s on I t is not to be u nde rst ood th a t
.
4 60 P LAN T DI S T R I B UTI O N
this reg i on i s exclusively occupied by decidu ous f orest
t ree s Inth e sp ring bef ore th e new c rop O f le a ves c omes
.

FI G . 1 25 — D e idu ou s
c Fores t sh a dy w oods in a ra vi ne
.R i ch
nea r the Illi nois R iv er centra l
,
R e d o a k is the p ri n
Illi noi s
. cip a l

fores t tree a l t h o u gh t here a re num ero us oth er kinds (Ves ta l )

, . .
 W E S TE R N CO NI FE R O U S F OR E S T 4 61

ou t th e f orests a re b ri gh t wi th sp ri ng fl owe rs L a te r i n .

the se a s onthe le a ves c a st so much sh a de th a t the f orest

fl oor is eithe r free from sm a ll pl a nts or ca nsupp ort th ose
on ly with b roa d thi nle a ves a ble to w ork with ve ry we a k
ligh t This i llust ra tes a p ri nciple O f c om mona pplic a ti on
.

i n th a t th e chief veget a ti on typ e i s th e m ost p owe rful

f a ct or i ndete rmi ning the na tu re o f the pl a nts a ss oci a ted
with i t M ore ove r the na tu re of the p la nt c ove r a ls o
.

exe rts a p owe rful i nfl ue nce on th e type of a nim a ls i n

FI G . 1 26 .
-
T h e H igh e st
T y pe of D e cidu ous Forest A b eech .

m a pl e w oods i n M i chiga n wi th h eml ock a lso This f orest d evel ops

,
.

into a typ e i nwhi ch th e b ee ch b y rea s ono f i ts d ense sh a de b ecomes

, _ ,

th e o nl y tree (V esta l ) . .

h a biti ng th e s a me regi on A f orest O i beech es a nd oa ks

”
-
.

a ff ords b o th a b u nd a nt f ood a nd suit a ble h omes f or such

typ es of a nim a ls a s squi rrels wh ich c ould su rvive with ,

difficulty i na p ra i rie or dese rt ,

.

TH E W ES TER N CO N I F ER O U S F ORES T (Fig 1 27 ) occupies .

a r egi on th a t is on th e wh ole c o ole r i n summe r a nd i n

some p a rts w a rme r i nwi nte r th a nth a t of th e D ecidu ous
Forest M ore ove r th e ra i ns a re m ore Copi ous i nwi nte r
.
,
4 62 PL AN T DI S T R I B UTI O N

( Fig . 1 28) a n d i n s ome p a rts of it a lm ost l a cki ng i n

summe r (Figs 1 29 Obvi o usly t rees which shed
.
,

thei r le a ve s i n winte r w o uld be ill suited to this regi on -

.

Th e c onife rou s trees on th e c ont ra ry bec a use they re

, ,

t a i n th eir le a ves for l onge r th a n a ye a r a re eve rg reen

an d S O c onstituted a s to t a ke a dv a n t a ge of opp ortu nity
to m a ke f ood a t a n y se a s on T rees with b roa
. d le a ves ,

F IG . Th1 27
e C on.if e ro us R a i n-
Fore s t o f th e P a cifi c N orthwest,
i nW a shi ngt o n Th e h ea vy ra i nf a ll supp o rts a d e n
. se fores t O f l a rge
tree s T h e sm a ll er tru n
. ks inthe fore gro u nd a re th o se of the gi a nt
a lder T he l a rge s tripe d tru n
. k a t the righ t is gia nt ced a r Th ere .

a re a lso D ou gl a s fi r, o n e or two t ru e fi rs , a n d th e S i tk a sp ru ce .

(Vesta l ) .

sh ed i nwi nte r or du ri ng th e dry h o t summe r, a re fou nd

i n this regi on bu t on ly a l ong st re a ms wh e re th e w a te r
supply i n th e s oil is sufficie nt to repl a ce th a t l ost f rom
th e le a ves Aw a y f rom th e st re a ms th e m o istu re con
.

te nt of the s oil is so much less th a t i nge ne ra l decidu ous , ,

t rees a re u na ble to secu re a supply a dequ a te to th ei r

mi nimum need s i nsumme r The needle le a ved c onife rs .
-
,

h oweve r do not l ose w a te r so ra pidly a nd find such

,

h a bit a ts suit a ble A no th e r f a c tor of s ome c onseque nce

.

i n th is c on necti on is th e p oore r s oil of this regi on ove r

 THE G R A S S LAND S 4 63

th e p a rticul a r a re a s wh e re f orests a re fou nd S imil a r .

p oor a re a s withi n th e ge ne ra l limits of th e D ecidu ous

F orest regi on a re a ls o O fte n occupied by l oc a l coni f

FI G 1 28. . T he C oa st R e dw oo d (S equ oi a mp ervi ren

se s ) i n centra l
Ca l if orn
ia . s grow to a di a m eter nea rly e qu a lli ng th e
Th ese tree
gi a nt s e qu o i a of th e S i erra N e v a d a an d t o a so m ewh a t grea ter
,

h eigh t Th ey a re rigidly confi ne d to th e z one of sea fo gs nea r th e -

d a re b e st d ev el o p e d ont h e sea w a r d mp e of th e o u ter l i n

.

coa s t a n
,
e

of co a s t ra n ge s (Cra ndal l ) .

erou s f orests , as f or e x a mple th e Pi ne B a rre ns

,
of N ew
J e rsey .

TH E G R A S S LA N DS a re c ove red with a veget a ti on of

h e rbs a nd gra sses h a vi ng t rees only i nspeci a lly f a v ored
,

l o c a lities I nthe summe r they O fte np rese nt a ma gnifi

.
4 64 P LAN T DI S TR I BU TI ON
ce nt p a nora m a of fl owe rs (Figs 1 3 1 . which v a ries,

f rom m onth to m onth with the a d v a nce of the se a s on .

Th e s oil of p a rts of this regi onis a m ong the m ost fert i le to

FI G 1 29
. . A V iew inth e S o u th ernR ocky M o u nta i ns i nC ol ora d o,

wi th w estern yell ow pine i n middl e gro u nd D o u gl a s fi r o n th e two

,

ridges a n
, d a tru e fir b et w eent h e ridge s . S cru b oa k occu rs i nrigh t
fore gro u nd a nd on dista nt slo p e a t l ef t Fores ts of this typ e a re
.

ch a ra c te ris t i c o f t h e high er d ri er a n
, , d co ld er p a rt s of th e W es tern
C on i f ero us Fores t
.
(Ves ta l )
.

FI G 1 30
. Whi te Pi ne inthe High er Pa rt of the S ierra Nev a da
. ,

Ce ntra l C a lifo rnia This is nea r the uppe r limi ts of tree s

.
.
 THE G R A S S LAN D S 4 65

be fou nd a nywhe re a nd c a p a ble of supp orti ng a v a ried

an d h ighly successful a g ricultu re The summe rs a re .

w a rm a nd ofte n h ot bu t ge ne ra lly spe a ki ng h a ve less

, ,

ra i n th a ni n th e f orested regi ons Th e winte rs a re c old .

an d th e g ro u nd deeply f roze nor s now c ove red I t is a ls o -

.

a regi on O f st ron g wi nds I t is still h oweve r a m a tte r

.
, ,

o f c on t rove rsy a s to p recisely wh a t a re th e c ont rolli ng

f a ct ors i n dete rmi ni ng th is veget a ti on type T rees ca n .

an d do g row i n this regi on whe n p l a nted a n d c a red for

'

FI G 1 3 1
. . An E xtensiv e V i ew o f P ra irie , wit h th e b l a zing sta r ,

a co mp osite ,
i n bl o o m
.
(F C Ga tes )
. . .

bu t they do not na tu ra lly i nv a de it with success Al ong .

st re a ms the re i s usu a lly a fri nge of t rees or eve na st rip .

O f f orest These a re a lm os t a lw a ys decidu o us a nd no t

.

c onife rou s f orest U nde r cultiv a ti on it h a s p roved es

.

p eci a llv fitted f or th e p roducti on o f wh e a t a n d c orn a nd

to a lesse r exte n t O f oth e r ce re a ls a nd he rb a ce ous c rops ,

p a rticul a rly fibe r pl a nts And m ore over j ust a s the food ‘

supply a nd pl a nt c ove r inna tu re On

.
,

ce supp orted th e i m
me nse h erds of bis on a nd othe r gra z mg a nim a ls of th is
regi on so n ,ow th e c rops o f c orn g ra i n a n d p a stu ra ge
, , ,
4 66 P L AN T DI S T R I B UTI O N
‘

m a ke i t a g re a t ce nte r for th e p roducti onof h ogs , c a ttle ,

an d d a i ry produce .

T H E D ES ER T i s l a rgely ch a ra cte rized by a h ot dry

cli m a te This te rm is used to c ove r a ve ry wide ra nge
.

o f clima tic a n d s o il c onditi ons a s well a s for a nequ a lly

,

wide ra nge O f veget a ti on In a ll regions h oweve r th e

.
, ,

ra i nf a ll is sc a nty a nd the ref ore i na dequ a te for the needs

o f the veget a ti on types so f a r discussed In s ome p a rts .

th e a nnu a l ra inf a ll is not m ore th a n a n i nch or two

FI G 1 3 2.
'

Pr a i ri e Ve ge t a t i o n o f th e Ea s tern P a rt o f th e Gras s
.

la nd R e gi on T h e b ro a d - l e a ve d pl a nt is ro si n
.
-
w eed The d a isy -like .

pl a nt is the pur ple cone- flower N umerous gra s ses a nd o th er

.

h erb a ceo s pl a nts a re a lso p rese nt (Ves ta l )

u . .

a nd c omes with g re a t i rregul a rity M uch of th e s oil is .

st ony thi n or s a ndy a lth ough h e re a nd the re a re v a lleys

, , ,

with s oil of g re a t fe rtility needing only w a te r to m a ke

a ve rit a ble pl a n t p a ra d ise M a ny such v a lleys a re now
.

i rrig a ted a nd yield imme nse c rops of c ot t on mel ons a nd , ,

a gre a t v a riety O f o th e r p rodu ce The na tu ra l veget a ti on .

v a ries f rom sc rubby bush es (Fig 1 33 ) onth e Sa ge B rush .

pl a i ns to th e sc a tte red c a cti (Fig 1 34 ) of th e m ore a rid .

p a rts The c a ctus exh ibits a cu ri ous a d a pt a ti on to the

.
 T HE DE SER T 4 67

s oil i nwhich it grows B ec a use the ra i ns a re infreque nt

.

the only w a te r a v a il a ble to the p l a nt is to be fou nd ne a r

FI G 1 33
. . Th e U ta h D e sert (e a sternU ta h ) wit h sca ttere d d esert
sh ru bs i na lk a line soil i nth e fl oo d pl a inwhi ch f orms th e f oregro und
- .

Th e cli ffs a re p ra ct i ca lly d ev o id o f pl a nt li f e

. (Ves ta l )
.

FI G 1 3 4
. . S u ccu lent Ve geta ti ono f th e s ou th ern A riz ona desert .

T he gi a nt ca ctus or Sa h u a ro (Sh reve )

‘

the su rf a ce immedi a tely a fte r ra i n As we h a ve see ni n

.

th e ch a pter on T ropisms a nd Correl a ti ons i n Pl a nts

4 68 P LAN T DI S T R I B UTI O N

(C h a p X I.I ) ,
m o st roo ts a re se nsi t ive to m o istu re , gra v
ity , a nd d iss olved s a l t s i n the s oil , a nd res p ond to
.

them i na wa y to secu re a f a v ora ble p ositi on A c a c tus .

root , h oweve r, m ust a l re a dy be i n p ositi on i f it is to

ta k e a dv a nt a ge of th e occ a si ona l sh owe rs This is .

brough t a b o u t th rou gh th e resp onse of th e roo t to h e a t .

R esp on di ng p osi tively to a w a rm tempe ra tu re kee p s th e

roo ts i n th e uppe r a n d wa rmer l a yers o f s oil wh e re th e
w a te r will be wh e n i t d oes c ome C e rt a i n o th e r dese rt
.

FI G 1 3 5
. . C h a p a rra l Ve ge ta ti ononthe i n l a nd sid e of one of th e
C a liforn i a C o a s t R a nge s T he pl a nts a re m o s tly eve rg ree nsh rubs of
.

m a nz a ni ta , sm a ll oa ks , a nd C e a no thus Wh ere l o ca l co ndi tio ns

.

a re fa v o ra bl e t re e gr o w th is w e ll d e ve l o pe d Liv e o a k , C a lifo rnia

—
.

ba y, a n d a few o th er b ro a d l e a v e d e v er gree nt ree s , a s w e ll a s a fe w

d ecidu o us t ree s , m a k e up the tree grow th i n the pi ctu re D ryer .

hills i n S ou th e rn C a li f orni a h a ve only the sh ru bs , a nd th ese a re

sm a lle r a nd se p a ra te d (C ra nd a ll )
. .

pl a nts g row wh e re th e re is subte rra ne a n w a te r to be

h a d by se ndi ng e norm ously l ong roo ts d ow nto i t .

C H APARRA L (Fig 1 3 5 ) is a na me of a veget a ti on type

.

exh ibi ted ing re a t pe rfecti oni nC a lif orni a a nd i np a rts of

the M edite rra ne a nregi onO f Eu rope I t occu rs i nregi ons
.
 C HA P A R R AL 4 69

with dry a nd s ometimes h o t summe rs bu t with c omp a ra

ti v el y mild wet wi n te rs Th e pl a nts t h emselves a re
.

sh rubs or sm a ll t rees m o stly equipped with thick shi ny

le a ves a ble to e ndu re th e he a t a nd d rough t o f summe r
wi th ou t l o si ng so much w a te r from th e le a ves a s to do
.
.

them se ri ous i nj u ry Th ey usu a lly p ut ou t fresh g reen

.

le a ves du ri ng th e wet se a s ona nd so a re a ble to m a ke

the best use of them d u ri ng th e time whe n c onditi ons

FI G 1 36 . A qu a ti c Ve geta ti on i na p rotecte d b a y of a M i chiga n

.

la k e R ush es i n sh a ll ow w a ter i n f ore grou nd T h e floa ting lea v es

. .

fa rth er ou t a re th ose of Wa ter lilies a nd th e w a ter sm a rtweed Th ere

-
.

a re a ls o se v en or e igh t ki nds o f submerse d a qu a ti c pl a nt s in pl a ces

,

wh ere th e d ep th is f rom two to ten feet T h e f ores t o nth e f a rth er

.

sh ore cons ists of b eech h a rd m a ple h eml ock elm a nd p a p er birch

, , , ,
.

(H A Glea s on)
. . .

for ph ot osynthesis a re m ost f a v ora ble As the se a s on .

a dv a nces a nd the s oil a nd a i r be come d rye r a nd d rye r ,

th e le a ves dep osit ove r th ei r su rf a ces a sh i ny l a ye r O f

w a te rp roofi ng cu ti n Th ey ca nstill do s ome ph o t o
.

synthesis a nd yet h a ve restri cted w a te r l oss to a s a fe

,

ra te This veget a ti on type me rges i nt o t ru e forest i n

.

m oiste r situ a ti ons onnorth h illsides a nd i nt o gra ssl a nd

on the d rye r sides }
470 P LAN T DI S T R I B UTI O N
S WA M P S se rve ve ry well to illust ra te on a sm a ll sc a l e
th e e ff ec t of w a te r on veget a ti on types I f one fi nds a .

p ond with dee p w a te r inth e middl e a nd g ra d u a lly Sh el v

in g t o w a rd th e sh o re with a b o tt o m a nd b a nks c omp o sed
O f deep rich s o il h e will fi n ,
d th a t th e veget a ti on is dis
t ributed in m ore or less regul a r c once nt ric z ones (Fig .

The ope n w a te r i n the m iddle i f d eep e nough , ,

ca nsupp ort o n l yfl oa ting pl a nts O utsid e th is th e re wi ll

.

p rob a bly be a z one of w a te r l ilies with thei r roots a n

FI G 1 3 7
. .
A Typi ca l S ubmerse d A qu a tic Fl ow eri ng Pla nt Cera ,

tO p h yll um . Th e v ery na rr o w divisi o ns o f the l e a v es a re e spec ia lly

ch a ra cte ris ti c o f t h e a qu a ti c h a bi t
. B o t h l ea v e s a nd s tems co nta i n
l a rge a ir sp a ces .

ch ored i n th e b ott om of th e p ond but with th ei r le a ves

exte ndi ng up to th e top a nd the re fl oa ti ng m ore or less
f reely (Fig Inthis z one ma y a ls o be f ou n
. d o th e r
pl a nts with l ong stem s a nd subme rged le a ves All th ese .

pl a nts h a ve we a k b odies a nd depe nd l a rgely o nthe w a te r

to supp ort thei r u pp ei f ree p a rts N e a re r sh ore is likely
.

to be a z on e o f t a ll ru sh like pl a nts such a s bul rushes

-
,

tule ca t t a ils etc Th ey exte nd to j ust a b ou t the li ne

,
-
,
.

whe re s oil a ppe a rs a b ove th e w a te r Th e nc omes a z one

.

O f sw a mp pl a n ts of v a ri ous s orts such a s sedges Thi s .

 S U MM A R Y 471

z one me rges m ore or less g ra du a lly a s th e g rou nd rises ,

an d bec omes d rie r with the su rro u n ding veget a ti ontype

, ,

f orest (Fig 1 38) or me a d ow for ex a mple Withi n e a ch

.
,
.

z one the re a re usu a lly m ore th a n one species Of pl a nt ;

an d m ore o ve r the species f o u nd
,
i ne a ch v a ry a cc ordi ng
to th e p a rticul a r regi oni nwhich the p on d occu rs .

S U M M AR Y S pecies O f pl a nts a nd a n
.
i m a ls h a ve c ome

i nt o existe nce th rough ev o lu ti on i n al l p a rts O f th e

h a bit a ble e a r th Th ey h a ve i nthe c ou rse Of time s p re a d
.

FI G 1 3 8
. . Ve geta ti on of th e N orth ea s ternC on
B og i fero us f ore s t
regi on . S urrou ndi ng o p en w a ter is a z o ne o f a n e ver green sh rub
o f t h e h e a t h f a m i ly ; i n th e b a ckgro u nd is th e e a s tern l a rch or
t a m a ra ck (H A Glea s on)
. . . .

a w a y f rom the pl a ces O f ori g i n i n wh a teve r di recti ons

they e nc o u nte red suit a ble c onditi ons Th ey h a ve not .

r e a ch ed a ll th e pl a ces suit a ble to them bec a use th ey h a ve

usu a l ly e nc o u nte red b a rrie rs O f one s ort or oth e r th a t

check or st op th em a lt ogethe r Wh a t is a b a rrie r to one .

org a n ism ma y be the me a ns of t ra vel to a no the r or a t

le a st p rese nt little difficulty i n bei ng p a ssed Bec a use .

a ll o rg a n isms do not fi nd th e s a me c onditi ons equ a lly

suit a ble for th ei r h a bit a ti on the re a rises a seg reg a ti onO f
,

th ose with like a d a pt a bilities i n the s a me or simil a r

regi on s S i nce th ose pl a nts a d a pted to simil a r h a bit a ts
.

gen e ra lly p ossess s im il a r a ppe a ra nce , the re rises the

472 P LAN T DI S T R I B UT I O N
ph e nome non O f vege ta ti on typ es a s distinct f rom the
ge og ra phic a l dist ributi on o f species or flora Th e l a tte r
.

is m ore intim a tely rel a ted to the p ossibili ties o f mig ra

ti on o r sp re a di n g whe re a s th e forme r depe nds m ore
,

l a rgely on the p ossibilities o f a d a pt a ti on T o put it

.

an o the r wa y
,
the one is l a rgely dete rm i ned by wh a t
species ca nget to a pl a ce a nd the othe r by the a rriv a ls
,

which ca na ctu a lly c ompete successfully a nd so m a i nt a in

themselves .
 C HAPTER XXX V III

D I S T R I B UTI O N OF LAND ANI M AL S

E X TE N T OF TH E —
S U B J E CT
Th e ge ogra p h ic a l di stri
.

b u ti on of a nim a ls de a ls with th e dist ributi on o f species

o ve r th e e a rth s su rf a ce th e c onditi ons which g ove rnthei r
’

existe nce i n th e regi on th ey occupy a nd th e a ge ncies ,

f a v ori ng or hi nde ri ng thei r dispe rs a l i nt o o the r a re a s .

It th us t re a ts of dist ribu ti oni nsp a ce a t the p resent time

an d a ppe a rs to be disti nct f rom p a le ont ol ogy or the dis ,

tri b u ti on o f species i n p a st times b a sed on the study of

f o ssil rem a i ns e nt o mbed i n th e ro cks a nd f rom ge ol ogy ,

de a li ng with th e physic a l c onditi ons a nd ch a nges of th e

e a rth Inre a li ty these th ree scie nces a re cl o sely li nked
.
, ,

an d m a ny p uzzli ng fe a tu res of ge og ra phic a l dist ribut i on

h a ve bee ne xpl a i ned only a fte r th e mig ra ti ons o f extinct
species a nd th e rel a ti ons o f a ncie nt l a nd m a sses we re
u nde rst ood Th e f a mily o f c a mels for ex a mple is rep
.
, ,

rese n ted only i n S o uth Ame ric a Asi a and N orth e rn , ,

Afric a From th ei r f o ssil rem a i ns it is k now n th a t i n

.

an cie nt times th ey i nh a bited N orth Ame ri c a th a t s ome ,

i ndividu a ls migra ted to S ou th Ame ric a while oth e rs ,

c rossed a l a nd b ridge i nt o Asi a a nd th us with the ex

ti ncti onO f th e N orth Ame ric a np op ul a ti on th ei r p rese nt
,

dist ributi onis m a de cle a r M ore ove r eve ry othe r species

.
,

o f livi n g a nim a l c omes from a l ong li ne o f a ncest ors

wh ose mig ra ti ons i n byg one times h a ve dete rmined its
p ositi on on th e e a rth t o a y .

-
MEA N S OF D I S P ER S A L —Th e mig ra ti ons Of a nim a ls
.

a re th e results O f a ge n cies of v a ried ch a ra cte r The .

high est te rrest ri a l spe cies such a s h orses c a mels el e

, , ,

ph a n ts be a rs dee r a n
, , d m a ny m ore or less cl osely r el a ted
,

4 73
47 4 D I S T R I B UTI O N OF LAND ANI MAL S
f orms depe nd up onth ei r ownp owe rs o f l oc om oti on In
,
.

sects bi rds a nd b a ts a re a c tively flyi ng c re a tu res and as

, , ,

a resul t o f th ei r eff orts ma y be dist ributed o ve r wide

a re a s .Am ong the l a st na med a nim a ls a nd a l a rge num

b er o f l owe r f orms wi nd f reque ntly pl a ys a n imp ort a nt
,

p a rt a s a dist ributi ng a ge ncy Th e re a re nume rous .

rec ord s o f bi rds a n d i nsects wh ich h a ve bee n c a ptu red

a b oa rd ship f a r o u t a t sea and it is a lt ogethe r p rob a ble
,

th a t m a ny o f th e oce a nic isl a nds h a ve bee n p op ul a ted

by wi nd swep t st ra ggle rs An eve n gre a te r numbe r O f
-
.

an im a ls of widely diff e re nt rel a ti onsh ips ma y be ca r

, ,

ried by ice or fl o a ti n g veget a ti on M ice ra ts squ i rrels .

, , ,

insects s na ils w orms a nd a h ost o f o th e r species a re

, , ,
‘

know n in m a ny i nst a nces to h a ve bee n t ra nsp orted by

such me a ns for l ong dist a nces Fi na lly th e re a re l a rge .
,

numbe rs o f a nim a ls wh ich h a ve bee n widely sc a t te red

ove r the e a rth th ro ugh the a ge n cy o f ma n In s ome .

i nst a nces whe re a species h a s bee n i nt roduced to com

, ,

ba t a n insec t pes t fo r ex a mple th e results a re highly ,

s a ti sf a ct ory but th e ra bbit i n Aust ra li a a nd th e English

,

S p a rro w i n Am e ric a o r th e N orw a y ra t o f th e w orld a t

,

l a rge a nd th e c o ck roa ch i nAme ric a a re i nt roduced species

wh ich h a ve multiplied a bu nd a ntly a nd bec ome a nnoy
in g to sa y th e le a st
,
.

TA N G I B LE BA R R I ER S — I t is a well k n ow n f a ct th a t

e a ch species o f a nim a l o ccupies a defi nite a re a up onth e

e a rth s su rfa ce a nd it is equ a lly t ru e a s p oi nted ou t
’

, , ,

th a t m a ny types wh e n t ra nsp o rted i nt o a new te rrit ory

th rive eve n bette r th a n i n th ei r origi na l h omes S i nce .

eve ry species b roa de ns i ts ra nge a s f a r a s i t ca n it is ,

evide nt th a t b a rrie rs te nd to ch eck its m oveme nts These .

b a rrie rs a re ve ry evide nt o r t a ngible i n s ome c a ses ; i n

o the rs th ey a re i n t a ngible or not so a pp a re nt M ou nt a i n
, .

ra n ges especi a lly th ose wh ose summits a re c ove red with

,

s now a re imp ort a nt t a ngible b a rrie rs which hi nde r the

,

w a nd e ri ng o f m a ny o f the sm a lle r a nim a ls D ese rts a re .

likewise Obst a cles to th e p rog ress o f d eli c a te o rg a nisms

 IN TANGI B L E B A R R I E R S 475

or to th ose requi ri ng c onside ra ble a m ou nts of m oistu re .

L a rge b odies o f w a te r set a n eff ec tu a l limit to th e sc a t

te ri ng o f l a rge te rrest ri a l a nim a ls a nd rive rs st re a ms
, , ,

an d l a kes a re a s imp a ssa ble i nth e c a se o f m a n y sm a lle r

species On th e othe r h a nd it is equ a lly m a nifest th a t
.
,

th e m oveme nts of a qu a tic f orms a re ch ecked by i nte r

v en in g t r acts o f l a n d .P ra ir ies a n d me a d ows f orests ,

an d b rushy c o u nt ry ro cky o utc rops di ffe re nces i n th e

, ,

ch a ra cte r of th e s o il a nd m a ny oth e r l oc a l fe a tu res a ct a s

a dditi on a l b a rrie rs to th e wide dispe rsa l of m a n y sp ecies .

I N TA N GI B LE B ARR R S I E — Adul t a nim a ls ma y mig ra te

or be t ra n sp orted i nto a new te rrit ory th a t suits th ei r i n
dividu a l needs a dmi ra bly a nd yet be u na ble to exist for
,

an y le n gth o f time on a cc o u n t O f th e l a ck O f suit a ble

shelte r from th e a tt a cks O f e nemies Bi rds for ex a mple .
, ,

ma y be u na ble to re a r thei r off sp ri ng owi ng to the a bsence

of t rees i n wh ich to build s a fe h omes Clim a tic c ondi
n
.

ti o s ma y a ls o re nde r a l o c a lity u nsuit a ble a s a h a bit a

ti onfor nume rous a nim a ls yet evenh e re a few a re o fte n
,

a ble to exist i f o th e r c on diti ons a re f a v ora ble The .

c a mel delights i na w a rm c ou nt ry yet it exists i nM on ,

g oli a a nd tige rs cert a i n a pes a nd th e eleph a nt which

, , , ,

we ma y thi nk of a s only t ropic a l a nim a ls a re not i nfre ,

que ntly f o u nd i n regi ons wh ere th e wi nte rs a re com

p a r a t i v el y seve r e B. u t a l l o f th ese i n t a ngible b a rrie r s
e nemi es l a ck of shelte r or r efuges a nd cl i m a tic c onditi ons
, ,
—a re o f less imp ort a nce th a n f ood An a nim a l or a .

S pecies with o u t f o o d bu t ina no the rwise f a v ora ble situ a

,

ti on bec omes ex t i nct ; with su fli ci ent no u rishme nt e a ch

,

ma y pe rsist eve nth o ugh c onditi ons a re seve re

,
.

LAW S AN D R ES U LT S OF D I S TR IB UT I O N It now f oll ows

.

from th e f a cts noted i nthe p recedi ng p a ra g ra ph th a t (a )

eve ry species of anim a l i s f ou nd i n eve ry qu a rte r of the
gl obe u n l ess i t h a s bee n h i nde red by s ome ki nd o f b a r
rie r. I f it is a ble to pe net ra te this b a rrie r (b ) it pe rsists
wh e re th e c onditi ons o f life a re f a v ora ble If i t d o es .
.

c onti nue a nd the su rrou ndi ng c onditi ons rem a i n p ra c

476 DI S T R I B UTI O N OF LAND ANI M AL S
tic a lly th e s a me for a l o ng pe ri od o f time the desce nd a nts
,

cl osely resemble th e origina l pi onee r a ncest ors But if

, .

the c ond iti ons g ra d u a ll y ch a nge a nd this is ge ne ra ll y

,

the c a s e (c ) the successful species g ra d u a lly ch a nges

,

a ls o a n d so c onti nues to fi n d th e te rrit o ry f a v ora ble .

I t h as f reque ntly h a ppe ned i n f orme r ge ol ogic a l a ges

th a t a few i ndivid u a ls o f a species h a ve bee nb ornwhi ch
d i ff e red s omewh a t f rom thei r p a re nts a nd this d i ff e r
,

e nce h a s bee n su ffi cie nt to e na ble th em to pe net ra te to

s ome exte nt i nt o new regi ons S till fu rthe r m odified
.

desce nd a nts i n succeedi ng ge ne ra ti ons h a ve c o rre

sp on di ngl y exte nded th e ra nge o f th e species a n d by ,

thi s g ra d u a l p rocess wide a re as h a ve bee n occupied .

S m a ll g ro ups o f i ndividu a ls withi nthese l a rge a re a s ma y

bec ome is ol a ted f rom th ei r fell ows a nd a mid sligh tly
,

di ff e ri ng su rrou ndi ngs they ma y u nde rg o g ra du a l ch a nges

wh i ch ultim a tely t ra nsf orm them i nt o a disti nct type .

In this wa y on e widely dist ribu ted species bec omes

b roke nup int o a numbe r o f new ones e a ch o f wh ich is
’

sep a ra ted by s ome s ort of b a rrie r f ro m the o the rs .

A N I M A LS O F O CEA N I C I S LA N DS Th e l a ws o f geo
.
-

g ra phic a l dist ributi on a re a dmi ra bly illust ra ted by the

hist ory o f m a ny is ol a ted v ol ca nic isl a nd s th a t m ost
ce rt a i nly h a ve neve r bee n u ni ted wi th a c onti ne nt a nd ,

a cc ordi n gly h a ve bee n c ol onized by i ndividu a ls c a rried

by wi nd or w a te r from s ome d ist a nt l a nd m a ss The .

G a l a p a g os G ro up situ a ted m ore th a nfive h u nd red miles

,

west o f Pe ru is i nh a bited by a nim a ls s ome o f which

, ,

a re ide n tic a l with th ose o f th e m a i nl a nd while o th e rs

,

exhibi t v a ri ou s d eg rees o f m odific a ti on Th e l a nd tor .

t oises liz a rd s a nd s na kes a re a l l disti nct spe cies a nd the

, ,

s a me is t rue o f m ore th a nh a l f o f th e i nsects a nd s na ils .

Whe re rel a ted f orms a re k no w n th ese a re i nv a ri a bly

S ou th Ame ri ca n The s a me ge ne ra l f a cts a ppe a r i n the
.

c a se o f th e Az o res th o ugh he re th e a nim a ls resemble

,

th o se o f E u rope ni ne hu nd red miles dis ta nt Th e spe cies .

o f th e Be rmud a s a re a ki n to th o se o f Ame ric a while ,

 Z O OL O GI CAL R E AL M S 477

the H a w a ii a nG ro up i nthis reg a rd be a rs a nu nmist a k a ble

resembl a n ce to J a p a n .

In a ll o f these i nst a n ces a nd i no the rs th a t might be ,

me nti oned th e evide nce u nmist a k a bly le a ds to the con

,

elusi on th a t while s o me i mm i g ra n ts f ro m th e m a i nl a nd
ma y re a ch a new te rrit ory a nd pe rish owi ng to u nf a v or
a ble c on diti ons othe rs bei ng bette r fitted h a ve pe rsisted
, , , .

Th e new su ou di gs
rr n n — s o il clim a te veget a ti on a nd , , ,

e nemies — a re di ff e re nt from th o se o f th e h ome l a nd ,

h oweve r a nd in these is ol a ted dist ric ts the i ndividu a ls

,

O f th e species g ra du a lly u n de rgo m odific a ti ons a s th e

ce ntu ries p a ss u ntil th ey di ff e r m a rkedly f rom the
origi n al an cest ors .

ZOOL OGI CA L R EA L M S Th e c ont ine nts a nd isl a nds of

.

th e e a rth f rom th e froze n north a nd s ou th to the

,

bu rni ng t ropics a re t od a y i nh a bited by m ore th a n a

,

milli on di ffe re nt species of a nim a ls which i n p a st t imes ,

h a ve c rossed b a rrie rs a nd bec ome a d a pted to thei r sur

rou n d ings And i n much th e s a me wa y th a t th e e a rth
.

is divided i nt o l a rge p hysic a lly di ffe re nt regi ons S O the ,

an im a ls h a ve roughly bee n i ncluded i n five g re a t re a lms

on th e b a sis o f th ei r dist ributi on Th ese a re th e Hol .

a rctic Ne ot ropic a l or S o uth Ame ric a n E th i opi a n Ori

, , ,

ent a l a nd Aust ra li a nre a lms Th e c onditi ons p eculi a r to

,
.

e a ch O f these l a rge divisi ons se rve a s a b a rrier to th e

gre a te r numbe r O f species of a na dj a ce nt re a lm a lth ough ,

the re a re a few (excep t i n th e c a se of th e Aust ra li a n

re a lm ) th a t i n te rmi ngle a l ong th e b orde r li ne a nd re nde r
the b ou nd a ry s omewh a t v a gue a nd i ndisti nct .

H OLAR CTI C R EA L M Th e H ol a rctic re a lm c omp rises

.
-

a pp roxim a tel
y a ll O f th e l a n d su rf a ce n or th o f th e T ropic

of C a n ce r a nd a cc ordi ngly i ncludes ne a rly th e wh ole of

,

N orth Ame ric a a l l of E u rope a nd Asi a north o f th e

,
‘

Him a l a y a s N O oth e r regi on of th e gl obe is S O highly

.

dive rsified with t ra cts O f pe rpetu a l ice imm e nse t reeless ,

dist rict s dese rt s sw a mps a nd a n a lm ost endless v a riety

, , ,

of s oil c on diti ons .

478 DI S T R I B UTI O N OF LAND AN I MAL S
Ina ncie nt times l a nd b ridge u nited Al a sk a a nd As i a
a
,

an d a pp a re ntly the re wa s a simil a r c on necti on with

E u rop e by wa y O f G ree n l a nd so th a t a nim a ls i n b oth ,

h emisphe res we re f ree to mi ngle Th is a cc ou nts for the .

f a ct th a t b o th the Ol d a nd the N ew W orld a re th e

h ome O f wildc a ts f oxes we a sels be a rs elk dee r wild, , , , , ,

oxe n be a ve rs squi rrels a n

, d h a res
,
Am ong the a nim a ls
, .

ch a ra cte ristic O f this re a lm i n the Weste rn Hemisph e re

a re th e musk o x p rong h orn a ntel op e ra cc oon m usk
,
-
, , ,

ra t a n
,
d p ra i rie dog wh ile th e E a ste rnHemisph e re is the
,

h ome of the two h umped c a mel musk dee r ch a m oi s a nd

-
, , ,

y a k . Th e di ff e re nces betwee n these two sub re a lms a re -

,

h oweve r no g re a te r th a n th ose sep a ra ti ng Al a sk a a nd

,

C a lif o rni a ; a nd th ey a re f a r less th a nth ose disti nguish i ng

th e H ol a rctic f rom a ny o th e r re a lm .

T H E N E OTRO P I CA L R EA L M Th e N e o t ropic a l or S outh .

Ame ric a nre a lm i ncludes S outh Ame ric a Ce nt ra l Ame ric a , ,

an d th e West I ndies Like th e H ol a rctic it is a n e nor

.
,

m o us t ra ct ch a ra cte rized by m ou nt a i n ch a i ns v a st ex
, ,

p a nses o f g ra ssy pl a ins a nd fully h a lf of i ts a re a is c ove red

,

by de nse a nd luxu ri a nt f orests N O oth e r re a lm is so rich .

inspecies a nd ove r h a lf of the k now nf a milies o f a nim a ls

,

a re rep rese n ted with in its b orde rs Am ong the ch a r .

a cteri sti c m a mm a ls a re th e disti n ct g roup of b roa d nosed -

m onkeys th e v a mpi re b a ts th e gui ne a pig a nd seve ra l re

, ,
-

l a ted species sl o ths a rm a dill o s th e t a pi r a nd seve ra l wild

, , , ,

d ogs a nd tige r c a ts Th e bi rds a re rep rese nted by ove r.

f ou r h u nd red species o f h ummi ng bi rd s m a ny p a rro ts , ,

t ou ca ns t a na ge rs th e ost rich like rhe a s a nd nume rous

, ,
-
,

o th e r species I nM exic o th e b ou n
. d a ries o f this rea hna re
not pe rfec tly dis ti nct si nce seve ra l H ol a rctic sp ecies , ,

such a s th e h a re th e grou nd squi rrel ce rta i n mice th e

, , ,

p um a dee r a nd sku nk a re dist ribu ted a s fa r s outh a s th e

, , ,

I sthmu s of P a na m a a nd eve nf a rth e r Onthe o the r h a nd .

,

ce rt a i n species o f m onkeys th e j a gu a r a rm a dill o t a pi r , , , ,

pecc a ry (a pig like c re a tu re ) a nd seve ra l o th e r a nim a ls

-
,

exte nd f rom th e s o uth i nt o M exic o o r bey ond B roa dl y .

 E THI O P IAN R E AL M 479

pe a king h oweve r the S o uth Ame ric a nre a lm is cle a rly

' !

defi ned a nd is one O f the m ost i n

, ,

te resti ng i nth e w orld .

ET H I O P I A N R EA L M Th is re a lm i ncl udes the wh ole of

.

Af ric a s outh of the S a h a ra D ese rt A ra bi a s o u th of th e ,

T ropic O f Ca nce r M a d a g a sc a r a nd seve ra l a dj a cent

isl a nds I t thus c orresp on
,

. ds i n p ositi on to th e S o uth
Ame ric a n re a lm a nd while m a ny o f th e fish es a nd l a nd
,

S hells O f th e two dist ricts a re cl o sely r el a ted th e ma m ,

m a ls a re ve ry disti nct Th e a pes a nd m onkeys a re of th e

.

na rrow nos ed type a nd ina ddi ti onto the ma nlike chim

'

- -
,

p a n z ee a n d g or ill a th e r e a re seve r a l species o f b a b o o ns

an d lemu rs Ope nor f orested c o u nt ry is the h ome of the
.

A fric a n eleph a nt gi ra ff e zeb ra s a nd o th e r wild a sses

, , , ,

an d ma n y a ntel opes wh ile i nregi ons wh e re m a rshy t ra cts

,

p rev a il th e rh i no ce ros a nd h ipp op ot a mus a re ch a ra cte r

i sti c a nim a ls The li on le op a rd a nd ost rich a re likewise
.
, ,

i nh a bit a nts of this re a lm th ough they exte nd bey ond its

li mits th ro ugh A ra bi a The S a h a ra D ese rt a cts a s a na l.

m ost pe rfect b a rrie r so th a t th e Afric a na nim a ls a l ong the

,

M edite rra ne a nS ea a re ve ry disti nct from th ose elsewh e re

onth e c on t i ne nt .

T H E OR I E N TA L R EA L M emb ra ces th a t p a rt O f Asi a s outh

O f th e Him a l a y a M o u n t a i ns J a v a B orne o th e Philip , , ,

pi nes a nd m a ny o the r isl a nds betwee nI ndi a a nd Au stra

,

li a. Th e a nim a ls o f this regi ona re cle a rly rel a ted to th ose

of Af ric a yet the re a re m a ny ch a ra cte ristic species th a t
,

a re f o u n d i n no o the r regi on Th e ma nlike a pe s a re .

-

rep rese n ted by th e ora ng u t a n a nd th e g ibb ons Th e re -

.

a re a ls o th e I n di a n eleph a nt th e tiger seve ra l species , ,

o f be a rs a n d dee r th e pe a c o cks a nd th e j u ngle f owl f rom

, ,

which our d omestic a ted f owl h as desce nded On the .

o the r h a n d the re a re ve ry few species of h orses a sses , ,

an d zeb ra s no rh i n ,
o ce ro s an d sc a rcely a ny a ntel opes
,
.

T H E A U S TR ALI A N R EA L M c omp rises Au st ra li a T a s ,

m a ni a N ew Gui ne a a nd seve ra l of th e a d j a ce nt isl a nds

, ,

o f sm a lle r size The gre a te r p a rt of this te rrit ory is

.
“

a rid s ome dist ricts bei n

,
g a dese rt with bu t sc a nty vegs
4 80 DI S T R I B UTI O N OF LAND ANI MAL S
ta ti on . Acc ord ingly it is no t f a v ora ble fo r a rich a nd
,

v a ried d evel opme nt O f species bu t the ones th a t do ,

exist a re m ore distinctive th a nth o se o f a ny o the r re a lm .

D eep w a te r a cts a s a b a rrie r a nd for milli ons O f ye a rs ,

its a nim a ls h a ve ev olved with ou t e nte ring i nto c ompeti

ti on with m ore p owe rful a n d o re speci a lized types mA .

ve ry few h ogs th e di ng o or na tive dog a nd seve ra l species

, ,

o f b a ts ve ry n e a rly c omplete the list O f h igh e r m a mm a ls .

On th e oth e r h a nd the re a re m a ny u nique species o f

,

p rimitive m a mm a ls such a s the k a ng a roo s w omb a ts a nd , ,

nume rous o the r rel a ted species th a t likewise a re a ki n to

the op osSum Th is re a lm is a ls o the h ome o f the re
.

m a rk a ble egg l a yi ng duck bill a nd spi ny a nte a te r a s well

- -
,

a s of n ume ro us species of bi rds such a s the b rush tu rkey , ,

p a rr o ts a nd pige o ns th a t o ccu r i nno othe r re a lm

,
.

R EA LM S U B V S O N S
DI I I — I n m uch the s a me m a n
. ne r
th a t th e U nited S t a tes is c omp o sed o f s ta tes c ou nties , ,

an d t o w nsh ips e a ch o f th e z o Ol ogi ca l re a lms is sub

,

divided i nt o lesse r dist ricts e a ch with a ch a ra c te ristic ,

a ssembl a ge o f a n im a l f orm s The H ol a rc tic o f N orth

.

Ame ri ca for ex a mple is d ivided i nt o no t less th a neigh t

, ,

ge og ra ph ic a l regi ons (see ma p fa ci ng p a ge wi th

f a i rly well defi ned a nd ch a ra c te ristic co nditi o ns depe nde nt
up on th e na tu re o f th e clim a te s oil a nd veget a ti o n , ,
.

Ea ch regi ona ls o po ssesses a disti nctive g roup o f a n im a l

species wh ich a re spe ci a lly a d a pted fo r feedi ng u p o n a
p a rticul a r ki nd o f veget a ti on for ex a mple or f o r livi ng , ,

a mid su rro u n di ngs o f a ve ry defi nit e ch a ra ct e r a l th o ugh ,

a sm a lle r n umbe r o f less speci a lized types ma y roa m f ro m

on e regi o n to th e n ext .

As a rule th e dist ributi on o f a give n species o f a ni

,

ma l is no t a s exte nsive a s th e regi o n i t occupies I t .

ma y be c onfi ned fo r ex a mple to a f orested a re a a

, , ,

dese rt a m a rsh o r a defi nite typ e o f s o il N o two sp ecies

n
.
, ,

o ccupy ex a c tly th e s a me ra n ge h oweve r so th e co t rol ] , ,

in f ct s bvi usly di f
f e r with th e species con ce rn ed
g a o r o o .

S p ea kl y b roa dly th ese f a ct ors a re suit a ble a n

,
d a v a il a ble
 M IG R ATI O N S OF AN I M AL S 4 81

f o od ,
S helte r from e nemies ,
a nd fa v ora ble clim a tic con
di ti ons .

M I GR ATI O N S O F A N I M A L S — In th e discussi on of th e
dist ributi on of a nim a ls it h a s bee n a ssumed th a t e a ch
species is c onfi ned to a f a i rly defi nite a re a ; bu t while ,

i n ge ne ra l th is is t rue sh a d a nd smelt mig ra te f rom

,

th e sea to l a y thei r eggs i n fresh w a te r ; ce rt a i n species

o f eels le a ve f resh w a te r to dep osit thei r eggs i n
th e sea ; m a ny o the r Species O f fishes mig ra te f rom de ep
w a te r i nt o S h a ll ow w a te r a t th e b reedi ng se a s on The .

rei n dee r musk ox bis on ce rt a i n species of h a res ra ts

,
-
, , , ,

an d m a ny o the r a nim a ls ma y j ou rney widely f rom on e

regi on to a n o the r chiefly i n se a rch o f f ood .

Th e bi rds h oweve r a ff ord the m ost st riki ng ex a mples

, ,

o f exte n ded mig ra ti ons wh ose impelling c a use rem a ins

a n un settled questi on P ra ctic a lly a ll th e i nsect e a ti ng
.
-

species of tempe ra te a nd c old regi ons spe nd thei r

wi nte rs i n w a rme r clim a tes the sh ore bi rds such a s , ,

the s a ndpipe rs pl ove rs a nd ph a l a ropes t ra veli ng a t

, ,

le a st one th ous a nd miles Th e g olde n pl ove r nesti ng

.
,

i n th e neighb orh o od o f th e A rctic C i rcle i n N orth

Ame ric a flies s ou thw a rd to N ov a S c o ti a a nd th e nce by
, ,

a di rect fligh t o f twe n ty fi v e hu nd red miles re a ches S ou th

-

Ame ric a wh e re it wi nte rs i n A rge nti na I t is a fu rth e r .

rem a rk a ble f a ct th a t i n c ompleti n g its rou nd t rip of

seve n th ous a nd miles it retu rns north by wa y of Tex a s .

A few Al a sk a n sh ore bi rds wi nte r i n th e H a w a ii a n

Isl a nds two th ous a nd miles dist a nt ; a S ibe ri a n sw a ll ow
wi nte rs i n M exic o or Bu rm a h ; th e E u rope a n sw a ll ow
migra tes to Afric a ; a nd the re a re h osts of o the r S pecies
wh ose j o u rneyings a re a s rem a rk a ble a nd a s difficult to
c omp rehe nd .
 C HAPT ER XXX I X

DI S T R I B UTI O N OF LIF E I N T HE O CE AN

P H Y S I CA L C H AR A T C ER S — T wo th i rds o f th e su rf a ce
-

of th e e a rth is c o ve red by oce a n s Th rough out fou r fi fth s

.
-

o f th ei r exte n t th e w a te r is ove r a mile i n d epth m ore ,

th a n two miles o ve r two th ird s o f th ei r a re a wh ile one

-
,

fif tee nth is ove r th ree miles d eep Th e g re a test dep th .

eve r s ou nded ,
feet is l oc a ted e a st of the Ph ilippine
,

Isl a nds .

S ea w a te r a s we k now is s a lty c ont a i ni ng a b ou t 27

, , ,

g r a ms o f c omm o n s a lt t o th e lite r I t a ls o
. c o n t a i n s in ,

much sm a lle r qu a ntities sa lts of m a g nesium c a lcium

, , ,

p o t a s sium a nd t ra ces of a c onside ra ble numbe r o f o th e r

subst a nces s ome o f wh ich a re o f p rime imp ort a nce i n
,

th e grow th o f m a ny a nim a ls a nd pl a nts Oxyge n nit ro .

,

g e n a n d c a r b o n d i o xide g a ses a r e a l so diss o lved i n i t .

In th e n eighb orh ood o f l a rge l and m a sses th e sea b ot

tom u su a lly sl opes ra th e r gra d u a lly to a p o i n t wh e re the
w a te r bec omes a pp roxim a tely 600 feet i n dep th He re .

th is c onti ne nt a l pl a te a u a s i t i s te rmed u nites wi th th e

, ,

c onti ne nt a l sl ope th a t f a lls a w a y with c omp a ra tive

a b rupt n ess i nt o th e a byss I n th ese deepe r oce a nic re
.

gi ons th e c onfigu ra ti on o f th e b o t to m v a ries widely I t .

ma y p rese nt th e a ppe a ra nce o f a ge ntly u ndul a ting pl a i n ,

h u nd red s o r th ou s a nds o f squ a re miles i n ex te nt with ,

h e re a nd th e re a t reme nd o us s a uce r like dep ress i onknow n

-

a s a deep . Or th e re ma y be rolli n g hills wi th wide i nte r

v en in g v a lleys o r,
n a rr o w g o rges betwee n m o u n t a in s
wh o se l o fty summi ts risi ng to th e su rfa ce form isl a nds ,

such a s th ose of the H a w a ii a n G roup .

482
 T E M P E R ATU R E OF T H E SEA 4 83

T EM P ER ATU RE — Th e he a ti ng e ffect of
OF T H E S EA .

the sunonoce a nw a te r is not pe rcep tible bel ow a depth

of a b o u t on e h und red feet a nd Obvi o usly is m ost m a rked
,

i n t rop ic a l regi ons O wi ng to th e ro t a ti on o f th e e a rth

.

an d th e rest ra i n i ng i nflue nce o f th e c onti ne nt a l l a nd

m a sses this h e a ted w a te r is m a d e to t ra vel i n gre a t
ci rcles i n b oth the northe rn a nd s o u th e rn h emisph e res .

In th e n orth A tl a ntic this cu rre nt i s na med th e G ulf

S t re a m whil e th e c orresp ondi ng one i n th e north P a cific
,

is th e J a p a n C u rr e nt Th e deepe r w a te rs o f th e oce a n
.
,

si nce these a re u na ff ected by the su n p rob a bly rem a i na t ,

a b o ut th e s a me tempe ra tu re f ro m ye a r to ye a r ; a n d it ,

ma y be a dded th e tempe ra tu re i ndeep w a te r is cl ose to

,

freezi ng a t the b o t t o m a nd g ra du a lly rises a s the su rf a ce

is a pp roa ch ed .

E FF E CT O F O CEA N C U RRE N T s — I t ma y p e rh a p s be .

u nnecess a ry to c a ll a tte nti onto th e f a ct th a t th ese o ce a nic

cu rre nts a re of th e high est imp ort a nce i nth e dist ributi on
of life i n th e se a N ot only do th ey c a rry th e fl oa ti ng
.

an im a ls a nd pl a nts for th o us a nds o f miles a nd th us

est a blish m a ny species i n h o mes th ey w o uld o th e rwise
neve r occupy bu t a ls o to a l a rge exte nt th ey gove rnthe
, , ,

mig ra ti on of fi shes a nd wh a les th a t p rey up on this f ood

an d a re thus O f g re a t imp ort a nce to th e fishi ng i ndust ry .

P R ES S UR E
. Th e a i r p resses up onou r b odies a nd oth e r
O bj ects wi th a weigh t o f a b ou t 1 5 p o u n ds to th e squ a re
inch a t sea level I n th e sea th e p ressu re i s d oubled a t
.

a depth of 35 feet a n d c onti nues to i nc re a se a t p ra ctic a lly

,

th e s a me ra te to th e g re a test dep th s Acc ordi ngly th e .

b ody o f a ma n if su nk to th e b o tt om o f th e deep O ff th e
,

Ph ilippi nes w o uld supp ort th e e norm o us weigh t of

'

a pp roxim a tely lb s S i nce a nim a ls a re k nown

.

to live a t a dep th of a t le a st th ree miles th e questi on

‘

na tu ra lly suggested is h ow is th is p ossible wh e n th e

,

p ressu re a m ou nts to c onside ra bly m ore th a n lbs .

p er squ a re inch I t is chiefly bec a u se th e b ody is l a rgely

.

c omp osed o f w a te r a nd w a te r is p ra ctic a lly i ncomp res

,
4 84 DI S T R I B UTI O N OF LI FE I N T HE O C E AN

sible A sp onge livi ng a l ong th e c oa st if su nk to a gre a t

.
, ,

depth w o uld Sh ow little dimi nuti oni n size owi ng to its

w a te ry c onsis te ncy i ts tissues a c c ordi ngly w o uld su ff e r
,

no c oll a pse a nd if f ood we re a v a il a ble it p rob a bly w o uld

,

live . At a l l eve nts seve ra l species o f widely rel a ted

,

an im a ls d redged f ro m dep th s ra ngi ng f ro m one to two

mil es h a ve bee nkep t a live for d a ys a nd eve nweeks .

O nth e o th e r h a nd i t f reque ntly h a ppe ns th a t c omp a ra

ti v ely l a rge a m o u n ts of g a ses a re diss olved i nth e tiss ues
o f deep sea a n im a ls Wh e n such i ndividu a ls a re ca p
.

tu red i nnets a n d d ra w n to th e su rf a ce this ga s exp a n ds

with th e reduced p ressu re a nd a s i t esc a pes from th e ,

tissues lite ra lly te a rs th e b ody i nto sh red s

Whi te ligh t is th e c ombi n
.

L I GH T — . ed p rod uc t of
V ib ra ti ons or w a ves i n th e eth e r o f di ff e re nt le ngth s .

Th ose o f rel a tively g re a t le ngth p roduce th e se nsa ti on

o f re d wh e n th ey f a ll up on th e re ti n a o f th e eye and ,

sh orte r V ib ra ti ons p roduce ora nge still sh orte r yell ow , ,

an d s o th rough gree n blu e to V i olet with th e sh ortes t

, ,

le ngth of a ll Wh e n th e su ns ra ys e nte r th e oce a n

.
’

th e d epth of thei r pe net ra ti on depe nd s up on th ei r

w a ve le ngth N O red re a ch es a g re a te r depth th a n300
.

feet N O g ree n p a sses th e 1 50 0 f oo t level a nd a t th e

.

d ep th o f a mile th e re i s no su nligh t wh a teve r F o u r .

fi f th s o f th e o ce a n fl oo r th e ref ore is i n a bs olute d a rk

, ,

ness so f a r a s th e su nis c once rned N eve rth eless th e re .

,

is a st rong p rob a bility th a t a feeble ligh t ma y be p rese nt ,

a s we sh a ll see p rese n tly .

Th ese physic a l fe a tu res s a li nity tempe ra tu re p ress , , ,

u re a n d ligh t a re ve ry evide ntly no t pe culi a r to th e

,

o ce a n bu t pl a y a n imp ort a n
,
t p a rt i n th e dist ributi ono f
l a nd a nd freshw a te r a nim a l s a s well Th ei r rel a tive i m .

p ort a nce h oweve r d iff e rs i nth e two c a ses A few o th e r

, ,
.

dist ribu ti o nf a ct ors will be no ted i ns ome o f th e succeed

i ng p a ra gra ph s .

D I S TR I B UT I O N O F S H ORE P LA N T S P ra c tic a lly a ll o f -

the a t ta ched se a weeds o r m a ri ne a lg ae g row a t dep ths less

 FL O ATING P LANT S 4 85

th a n 1 50 feet a nd a cc ordi ngly a re c oa st a l f orm s Th e

,
.

g ree n species th rive best i n yell ow ligh t a nd a s th is is ,

m ost ple ntiful i nsh a ll ow w a te r th ey fri nge th e immedi a te

sh ore li ne B row na nd red species fl ou rish i nblue ligh t
.
,

an d the ref ore a re i ndeepe r w a te r Wh ile ligh t thu s con.

t rols the dist ributi onof these pl a nt s i na give n l oc a lity ,

tempe ra tu re is a p rime f a ct or i ndete rmi n i ng th ei r di stri

bu ti ono ve r wide r a re a s C old w a te r species exte nd well
.

to th e so uthw a rd bu t st op sh ort o f tempe ra te w a te rs

wh ose ch a ra c te ristic f orms a re i ntu rnrepl a ced by w a rm
wa te r species i nth e t ropics .

S ea weeds a re not only c omm erci a lly v a lu a ble for th e

p o t as h a nd i o di ne th ey c o n t a i n bu t th ey a re
,
exte n sively
e a te nby m a ny sp ecies of c ra bs sna ils sea u rch i ns etc , , ,
.
,

wh o se dist ributi on is th u s dete rmi ned O f g re a te r i m .

p ort a nce is th e f a c t th a t th e dec omp ositi on of thei r

b odies t ogeth e r with th a t o f org a nic m a tte r swep t f rom
,

th e s oil supplies th e oce a nwith nit roge na nd nit roge nous

,

c omp o u nds up onwhich th e g rowth of a v a st numbe r of

o th e r pl a n ts a s well a s a nim a ls depe nds .

F L OATI N G P LA N TS S i nce th e disc ove ry o f th e mic ro

.

sc ope i t h a s bee n f o u n d th a t th e sup e rfici a l w a te rs of

th e oce a nsupp ort a we a lth of pl a nt life th a t i np oi nt of
numbe rs of i ndividu a ls a nd a ggreg a te bulk fa r o utstrips
th e sh ore f orms Th ese floa ti ng species a gree i n bei ng
.
-

u nicellul a r a nd i n c ont a i ni ng chl orophyll a nd o th e r p ig

ments th a t e na ble th em to utilize th e e ne rgy of su nligh t
a fte r th e f a shi on o f l a n d pl a nts He nce di rectly or i n .
,

di rectly a ll of th e a nim a l life i n th e sea depe nds up on

,

m a rine p l a nts for thei r p rim a ry f ood supply .

Am ong th e c omm on fl oa ti ng pl a nts a re th e di a t oms

( Fig . 1 39 ) th a t a re a t o n c e disti nguished by bei n g en
cl osed wi th a Sh ell o f fli nt exquisitely sculp tu red a nd
,

with sh a pes ra ngi ng f rom discs a nd ellipses to rect a ngles ,

which ma y f orm th re a ds whe re m a ny a re a tt a ched to

geth er These pl a nts a re ext ra ordi na rily a bu nd a nt i n
.

c old w a te r but i nthe t ropics a re l a rgely repl a ced by o ther

,
4 86 DI S T R I B UTI O N OF LI FE I N THE O C E AN

types a m ong wh ich a re m a ny th a t p ossess skelet ons of

lime In b oth regi ons the re a re a dditi ona l species
.
,

know na s p eri dinea ns wi th out mi ne ra l skelet ons but im

’

p orta nt a s a n a rticle i n the d iet o f m a ny fish es a nd

i nte resti ng i n th a t seve ra l species a re b ril li a ntly
ph osph orescent .

P LA N K T O N Al th ough fl oa t ing pl a nts norm a lly occu r

.

a t dep th s less th a n 200 feet thei r b odies a re h e a vie r ,

th a nw a te r a nd a t a ll times g re a t numbe rs a re g ra du a lly

,

FI G 1 3 9
. . D i a to ms , mi cro sco pi c pl a nts wi th fli nty sk el e tons .

settli ng to th e b o tt om He nce it is no t su rp risi ng to .

fin d th a t e a ch o f th e m a ny diffe re nt species o f a nim a ls

th a t di rec tly o r i ndi rectly live on this food supply is
fitted to live a t f a i rly d efi nite depths a nywh e re f rom
the su rf a ce to th e o ce a n fl oor All O f th ese swimmi ng .

an d fl o a ti ng org a nisms pl a nt a nd a n im a l a re c ollectively

, ,

k now n a s p la nkton In th e pl a nkt on the pl a nts ou t

.

numbe r th e a nim a ls but to wh a t ex te nt is no t ex ac tly

,

k nown In th e B a ltic sea it h a s bee n estim a ted th a t

.

th e pl a n ts m a ke up fif ty six p er ce nt a nd the a nim a ls -

forty f ou r p er ce nt o f th e to t a l a m o u nt I t h a s fu rth e r
-
.

bee n c a lcul a ted th a t th e v olume o f pl a nts d oubles i n

 U N I N C E L L U L AR OR GANI S M S 4 87

a little ove r th ree d a ys th e excess se rvi ng a s food for

,

an im a ls or se ttli ng to the b o tt om Whethe r these esti .

m a tes a re of ge ne ra l a pplic a ti on is u nk now n I t is cer .

t a i n h oweve r th a t th e numbe r of i ndividu a ls p er u nit

n
, ,

v olume of sea w a te r v a ries a cc ordi ng to se a s ona l a d

o th e r c on diti ons .

U N I CEL LU LAR ORGA N I S M S AN D B OTT O M D EP OS I T S .

S eve ra l th o us a nd Species o f one celled a nim a ls occu r i n -

the pl a nkt onof a ll th e oce a ns of the gl obe The l a rge r .

numbe r of these a re rel a ted to the a m oeb a but di ffe r in ,

th e p rese nce of a supp orti ng skelet on (Fig Th ose .

FI G 1 40
. .
— Mi croscopi c One
'

M a ri ne A nim a ls (Gl ob irgeri na )

-
Ce ll e d
with ch a mb ere d cells l a rgely resp o nsible for ch a lk dep osits
, .

with fli nty skelet ons the ra di ol a ri a ns c omp rise a t le a st

, ,

species livi ng f or the m ost p a rt i n deep t ropic a l

,

w a te rs of th e P a cifi c oce a n O the rs with S kelet ons of .

lime (the f ora mi nifera ) a re likewise t ropic al f orms of

wide r dist ribu ti on Th e numbe rs of b oth of these g roups
.

bec ome i nc redibly g re a t u nde r fa v ora ble c onditi ons a s ,

m a ny a s i ndividu a ls occu rri ng in eve ry lite r o f

w a ter a t or ne a r th e su rf a ce th rough out th ous a nds o f
S qu a re miles .

Th e b odies of these a nim a ls a re h e a vie r th a n w a t e r ,

an d wh ile ve ry m a ny a re bu oyed up by oi l d rops a n d

4 88 DI S T R I B UTI O N OF LI FE I N THE O CE AN

o th er fl ot a ti ondevices othe rs gra du a lly si nk a nd fi na lly

,

se ttle to the sea b o tt om whe re th ey f orm exte nsive de

p osits I t h a s bee n c a lcul a ted th a t in ce rt a i n dist ric ts
.

this sedime nt i nc re as es one i nch eve ry ten ye a rs a nd ,

the numbe r o f org a nisms included is bey ond c omp re

h en si on .I n Engl a nd a n d F ra n ce for ex a mple the re , ,

a re g re a t ch a lk dep o sits a pp ro xim a tely 6 00 feet i nth ick

,

ness a nd i n th is c o u nt ry simil a r a c cumul a ti ons of s ome

,

wh a t lesse r th ick ness exte nd th rough seve ra l st a tes This .

m a te ri a l i n l a rge m e a su re c onsists o f th e skelet ons o f

, ,

u nicellul a r org a nisms th a t numbe r f rom

. to
to th e cubic i n ch .

S O M E P E LA GI C M A N Y CE LLED A N I M A L S - — I n a dditi on
to th e f ree swimmin -
g or fl oa ti ng y o u ng of p ra ctic a lly
eve ry l a rge g roup o f th e a nim a l ki ngd om the re a re m a ny ,

a d ult a n im a ls of widely di ffe re nt f orm a nd rel a ti on

sh ip wh ich a ls o swim or fl oa t (Fig Th e m ost .

imp ort a nt o f th ese a re th e c opep ods sm a ll sh rimp ,

like org a nisms be a ri ng a sligh t resembl a nce to a m i ni a

tu re m a nd oli n These c re a tu res ra rely o ve r a f ou rth
.
,

o f a nin ch i nle ngth feed onpl a nts a nd i nm a ny regi ons

, ,

a re so a bu n d a nt th a t they a re p ra c tic a lly th e only f ood

of m a n y c omme rci a lly v a lu a ble fishes as well a s o f sev
e ra l species o f wh a les They a re a ls o p reyed up on by
.

sh oa ls o f j elly fi sh es v a ri o us w orms seve ra l species o f

-
, ,

f ree swimming s na il s a nd nume ro u s o the r c re a tu res re

-

l a ted to th e ve rteb ra tes M a ny o the r a nim a ls such a s

.
,

th e squids fishes l a rge a nd sm a ll sh rimps a nd c ra b like

, ,
-

an im a ls O f v a ri ous f orms a nd sizes usu a lly with c rims on

c ol ored b odies or a s t ra nsp a re nt a s gl a s s a ls o live up on ,

c opep od s or up on o th e r a nim a ls th a t feed up on th em .

A t th e s a me time i t Sh ould be no ted th a t ve ry m a ny

o th e r a n im a ls feed up on pl a nts but none o f th ese is so ,

widely dist ributed or o f such h igh ec onomic imp ort a nce

a s th e c opep od
—
.

OR GA N S O F F LOTAT I O N P rot opl a sm is s omewh a t

.

h e a vie r th a n sea w a te r a nd h e nce will si nk To ove r

,
.
 OR GAN S OF FL O TATI O N 4 89

c ome this tende ncy m a ny a nim a ls a re supplied with

defi nite l oc om o t or org a ns th a t e na ble them not only to
keep a fl oa t but to m ove ra pidly th rough th e w a te r The .

sh a rks m a cke rels a nd h e rri ngs a ll a ctive swimme rs f a ll

, , ,

i n this cl a ss M a ny species of sh rimp like a nim a ls cer

.
-
,

t a i ns na ils a nd w orms a nd the y ou ng or l a rv a l st a ges o f

,

m a ny a nim a ls a re likewise p rovided with fins p a ddles , ,

cili a etc th a t e na ble them to m a i nt a i n f a v ora ble p o

,
.
,

si ti ons .

FI G 1 4 1
. . M a ri ne C op ep o ds . A
, co ld w a ter form ;
-
B , tro pi ca l .

In nume ro u s the r a nim a ls with or with ou t l oc o

o ,

m ot or orga ns th e b ody is m a de ligh te r by the p rese nc e

,

o f oi l or f a ts In ce rt a i n fishes a i r bl a dde rs filled by

.
,

gulpi ng a i r a t the su rfa ce or by sec ret ory a ctivity of the

encl osi ng w a lls a re a ls o of se rvice i n keeping the b ody
a fl oa t . Then a g a i n the re a re fe a th e r like a ppe nd a ges -
,

b ristles pl a tes etc th a t wh ile th ey do not a ctu a lly p re

, ,
.
, ,

ve nt si nking neve rth eless p reve nt i t t a ki ng pl a ce ra p

,

idly i n th e c a se of l a rge numbe rs of m ostly sm a ll sized -

org a n isms J ust a s a le a d S h o t Si nks m ore ra pidly th a n

.

when be a te n i nt o a th i n sh eet so these st ructu res th a t ,

gre a tly i nc re a se the su rfa ce of th e b ody e na ble the org a n

4 90 DI S T R I B UTI O N OF LIF E I N T HE O CE AN

ism to keep a fl oa t with o ut th e excessive use o f l oc om ot or

org a n s .

A N I M A S OF T H E S EA F LOOR
L — O ur knowledge o f th e
an im a ls livi ng i n sh a ll ow w a te r is f a i rly c omplete bu t ,

o f life a t dep th s o f o ve r a m il e much rem a in s to be

le a rned S uch deep sea a nim a ls a s a re b rough t to th e
.
-

su rf a ce a re usu a lly a tt a ched to the b ott om or a re of slug

gish h a bits , a nd a re thu s th e ones th a t a re re a dily ca p
tu red by n ets d ra gged a l ong th e b ott om I f one we re .

to d ra g a n e t f ro m a b a ll oo n a l ong th e su rf a ce o f th e
e a rth it is obvi ous th a t bi rds m a ny i nsects a nd o the r
,

a ctive c re a tu res w o uld ra rely if eve r be ca ptu red N ev .

erth el ess a sufficie nt numbe r o f specime ns h a ve bee n

secu red to e na ble u s to sa y th a t eve ry o ne of the l a rge r
g roup s of a nim a ls h a s its deep sea rep rese nt a tives -
.

S p onges c ora ls w orms o f widely di ffe re nt a ppe a ra n

, ,
ce ,

s na ils cl a m s c ra bs sh rimps sea u rch ins sta rfi sh es a s

, , , , ,

well a s fish es a re p rese nt i na bu nd a nce M a ny o f these .

a re quite simil a r to species a l on g sh ore O th e rs dep a rt .

m ore or less widely i nst ructu re f rom th ei r sh a ll ow w a te r

rel a tives It h a s a ls o bee n p oi nted ou t th a t deep sea
.
-

an im a ls h a ve a l a rge a m ou nt of w a te r i n thei r t issues

which e na bles th em to with st a nd th e e norm ou s p ressu re
to which th ey a re subj ected As no g ree npl a nt s live i n
.

deep w a t e r th e org a nisms th a t ra i nd ow nf rom the lesse r

,

depth s se rve a s f ood for m a ny deep sea Species th a t i n -

t u rnma y be p reyed up onby th e oth e rs .

P H OS P H ORES CE N CE F ro m time immem ori a l ma nh as

.

k now n of th e ph osph oresce nce of th e sea a nd for a ,

c omp a ra tively l ong pe ri od it h a s bee n rec og nized th a t

it is p roduced by nume rous species o f org a nisms Neve r .

th el ess i t is on ly i nve ry rece nt times th a t v a ri ous deep

sea expediti on s h a ve give n us a f a i rly a ccu ra te noti on
o f its ge n e ra l fe a tu res We now k now th a t m a ny u ni
.

cellul a r a nim a ls a nd pl a nts p o ssess this peculi a rity of

emit ti ng fl a sh es like th ose o f th e fi refly a nd th a t s ome ,

o f the fishes a c on side ra ble numbe r of f ree swimmi ng

,
-
 S U MM A R Y 4 91

w ms a nd ce t i n species of sm a ll c ra b like a nim a ls

or ,
r a -

libe ra te from the su rf a ce of thei r b odies slime th a t gl ows

like a t orch Inm a ny fishes a nd seve ra l squ ids eye like
.
-

ph osph oresce nt org a ns a re imbedded i n the ski ni nv a ri

ou s p a rts of the b ody a n d the ph osph oresce nt or gl owi ng
m a te ri a l is b a cked up by a reflect or a nd the ligh t is
m a de to shi ne th rough a le ns a fte r the f a shi onof a bull s ’

eye l a nte rn In ce rt a i n l oc a li ties it is know n th a t

.

the org a nic m a tte r on th e oce a n fl oor c ont a i ns g re a t

numbe rs of b a cte ri a or sm a ll u nicellul a r pl a nts th a t
, ,

a re a ls o lumi n o us He nce it is believed by m a ny stude nts

.

o f the subj ect th a t th e o ce a n dep ths th ough dev oid of ,

su nligh t ma y neve rtheless be feebly illumi na ted by

,

ph osph orescent light Th is w ould p e rh a p s a cc ou nt for

.

the g re a t owl like eyes of seve ra l deep sea a n

-
i m a ls a l -
,

th ough i t is to be noted th a t m a ny sp ecies a re t ot a lly

bli nd .

Th e use of th ese ph osph orescent org a ns is s omewh a t

p roblem a tic a l Ins ome species they a re a lm o st ce rt a i nly
.

used to a tt ra ct org a nisms th a t serve a s f ood In o the r

m
.

c a ses they ay se rve to d rive Off e nemies or be used a s

signa ls to a tt ra c t th e opp osite sex .

S U M M AR Y — Pl a nt life i n th e o ce a n c omp rises a

.

sc a nty fri nge O f a lg ae a tt a ch ed i n the sh a ll ows a l ong

sh ore a nd widely dist ribu ted a nd exceedi ngly a bu nd a nt
,

u nicellul a r Species fl oa ti ng ne a r th e su rf a ce V a s t nu m .

be rs of i ndividu a ls of this l a st cl a ss a re c onti nu a lly set

tli ng to th e b ott om a nd a t a ll depth s a re fed up on by
,

V a ri ou s species O f a nim a ls This flOa ti ng p opul a ti on

.
,

or pl a n kt on th us exte nd s from th e su rfa ce to the b ot

'

tom th ough th e l a rge r p rop orti on is c onfi ned to w a te rs

,

less th a n a mile in dep th On th e o ce a n fl oor othe r .

an im a ls c ra wl ove r or th rough the mud or a re a nch ored

in i t .

D eep sea expl ora ti on a s a S cie nce i s less th a n fifty

-
, ,

ye a rs ol d a nd much reg a rdi ng th e dist ributi onof m a ri ne

org a n isms rem a i ns to be le a rned bef ore we ca n dete r
4 92 DI S T R I B UTI O N OF LIF E I N T HE O CE AN

mi ne th e f a ct ors c ont rolli ng i t In m a ny i nst a nces the

.

e ffect s of ligh t a nd tempe ra tu re i ntheir rel a ti onto dis

,

tri b u ti on a p p e a r to be f a i rly cle a r bu t th e eff ects of

,

sa li nity a nd p ressu re a re too impe rfectly u nde rst ood to

a ll ow of wide ge n e ra l iz a ti on
. s
 C HAPTER X L

T HE EVOLUTI O N OF M AN

T I M E R EQ U I RED FOR M A N S EV OLUTI O N

’
At the .

p rese nt time the re is bu t one species of ma n c omp rising ,

seve ra l disti nct ra ces which di ff e r from e a ch othe r a s

reg a rd s c omplexi on c ol or of h a i r a n
,
d eye s st a tu re sh a pe
, ,

O f he a d,
an d m a ny o th er st ructu ra l a s well a s me nt a l
peculi a rities I f a ll of th ese widely di fferi ng ra ces a re
.

th e result o f ev oluti on f rom a S i n gle a nce st ra l type

an d the re is no g oo d r e a s on f or believi ng the c ont ra ry
the nthe le ngth o f t ime requi red f or these ch a nges must
n
h a ve bee e norm o u s .

I t is a ls o re a sona bly ce rt a in th a t th e g re a t diffe re nces

existi ng a m ong m ode rn ra ces with reg a rd to th ei r re
li gi ou s a n d s oci a l cust oms thei r l a ws a nd th e v a ri ous
m
,

ph a ses of c o mu na l life a re th e result of g ra du a l ch a nges

exte n ding ove r a l ong p e ri od of ye a rs Th e w orld wide
.
-

dist ributi ono f ma na s well a s the gre a t size of his b ra i n ,

when c omp a red w ith th e h ighest a pes a ls o i ndic a tes ,

his g re a t a ntiquity On the b a sis of th ese f a cts i t is

.

a s sumed by s ome i n vestig a t ors th a t ma n or ra the r a n ,

a p e like c re a tu re with a n a dmixtu re o f h um a n ch a ra c

-

te rs fi rst a ppe a red on th e e a rth

,
ye a rs a go .

Oth e r stude nts of th e subj ect a re c onvinced th a t

00 0 ye a rs is ne a re r th e t ru th .

P LA CE OF OR I GI N Gra n ted th a t ma n fi rst a p pe a red

'

milli ons of ye a rs a go, i t now rem a i ns to dete rmi ne his

pl a ce of origi n For seve ra l re a s ons Asi a a ppe a rs to
.

fulfil th e requi reme nts m ost s a tisf a ct orily In the fi rst .

pl a ce th e re a re ve ry st rong i ndic a ti ons th a t h um a n

civiliz a ti onh a s e ndu red for a much l onge r pe ri od onth a t
c onti ne nt th a n elsewhe re S ec ondly it is th e h ome of
.
,

4 95
4 96 T HE E V O LUTI O N OF M AN

m a ny o f th e oldest d omestic a ted a nim a ls th e c omm on ,

fowl c a mel g oa ts c a ttle a nd s ome o f the b re eds o f the

, , , ,

h orse dog a nd ca t bei ng O f Asi a tic origi n This regi on

, , ,
.

a ls o is a dj a c e nt to J a v a th e h ome of the m ost a ncie nt

,

know n type o f ma nlike a nim a l Fu rth e rm ore this

-
.

c onti ne nt wa s th e ce nte r of m a ny h um a n mig ra ti ons

wh ich a re k now n to h a ve ra di a ted i nt o o the r regi ons .

Al s o ce rt a in fe a tu res o f m a ns p rese nt dist ribu ti on a re

’

best expl a i ned on this a ssumpti on Fi na lly th e a ncie nt .

,

clim a tic c onditi ons of ce nt ra l A si a we re such a s to f a v or

th e ra pid ev olu ti on of th e h um a n a ncest or .

EAR LI ER H I S T OR Y O F T H E H U M A N R A CE Th e e a rlie r .

s ta ges of hum a n ev olu ti on a re u nknow n N eve rth eless .

the re a re ma ny fe a tu res a ss oci a ted with the ge ol ogic a l

ch a nges of Asi a a few milli on ye a rs a go which whe n
, ,

studied i nc onnecti onwith th e st ructu re O i m ode rnma n

like a pes a nd th e f ossil rem a i ns O f e a rly ma n e na ble ,

the skilled i nvestig a t or to m a ke a sh rewd guess a s to

wh a t a ctu a lly occu rred In th e fi rst pl a ce th e c onti ne nt
.

wa s l a rgely c ove red with t ropic a l f orests i n h a bited it ,

is believed by a nima ls resembli ng i n a b roa d wa y th e

, , ,

m ode rn lemu r a m onkey like species With th e ele v a

,
-
.

ti ono f the l a nd a nd th e result a nt l owe red tempe ra tu re

an d in c re a sed d ry ness these f orests i n l a rge me a su re
,

dis a ppe a red S ome of th e lemu r a ncest ors pe rsisted i n

.

livi ng i nth e t rees a nd thei r m od e rndesce nd a nts a re th e

,

gibb ons ora ng u t a n ch imp a nzee a nd g orill a O th e rs i n

,
-
, .

th e st ruggle f or existe nce we re c ompelled to a b a nd on

the t rees a nd m a ke th ei r h ome i n ope n c o u nt ry The .

h i nd limbs bec a me a d a pted for w a lki ng o n th e g rou nd .

Th e h a nd s we re rele a sed f rom climbi ng a nd se rved m ore

p a rticul a rly for g a th e ri ng f ood a nd g ra spi ng we a p ons .

Th e b ra i n d evel oped i n c onnecti on wi th th e i nc re a sed

f a cility o f h a nd m oveme nt a nd with th e p owe r o f speech ,

th e t ra its which ch a ra cte rize th e hum a n species bec a me

es ta blish ed Th is his tory a ppe a rs highly f a nciful p er
.

h a ps bu t th a t i t c ont a i ns m o re th a n a g ra i n o f t ru th
,
 T HE J AVA M AN —APE 4 97

will bec om e evide nt a fte r th e ex a mi na ti on of ce rt a in

f ossil rem a i ns wh ich a ctu a lly h a ve bee n disc ove red
(Fig .

T H E JA V A M AN AP E — Betwee n the ye a rs 1 891 a n

-
d
1 894 th e s omewh a t sc a tte red a nd i n c omplete f ossil re
m a i ns of a ma nlike c re a t u re the J a v a ma na p e (Fithe
-
,
-

ca n th rop u s ) we re disc ove red i n J a v a a n

,
d a roused a ,

FI G 1 4 2
.
— R econstru cti onof P rehistori c Ma n Ja v a Ma n
. Le f t ,
Ap e P i theca nthropu s erectu s ; cente r N ea nd ert h a l ma n a nd ri gh t ,
, , , ,

C ro M a gnonma n m od ell e d o nt h e res tore d skulls by Profess or J H

-
. .

M cGregor . T h e s econd a nd third hea ds a re m o delled on nea rly

complete sk u lls O f th e fi rst h ea d on
. ly th e upp er p a rt of the
cra nium a nd two teeth w ere fou nd Ph oto gra ph e d f rom origina ls in
.

America nMu seum of Na tu ra l His tory N ew York ,

.

high deg ree O f i nte rest a s well a s c ont rove rsy i n b oth
p opul a r a nd scie ntific ci rcles From th e p ositi onof the
.

ge ol ogic a l f orm a ti on a nd th e na tu re of th e a cc omp a ny

,

in g f o ssils i t is believed th a t th e a ge of this species is

,

not less th a n ye a rs It is a ls o re a s ona bly cer

.

t a i n th a t th e f orm a nd st ructu re O f th e b ody a nd its ,

m ode of l oc om oti on we re esse nti a lly h um a n while th e ,

skull a nd th e size a nd sh a p e of the b ra i na re a s ch a ra c

teri sti c of the highe r a pes a s o f ma n The S ize of the .

b ra i nO f th e high es t a p es is six h u nd red cubic ce ntimete rs ,

4 98 T HE E V O LUTI O N OF M AN

or a b out fo u r h u nd red cubic ce ntimete rs less th a ninthe

l owest ra ces o f ma n while th e ma na p e h olds a ni nte r
,
-

medi a te p ositi on with a c a p a city o f s omewh e re be twee n

,

eigh t hu nd red a nd fif ty a nd ni ne h u nd red cubic c e nti

mete rs The J a v a ma na p e thus rep rese nts a st a ge
.
-

i n h um a nev oluti on whe re the b ody wa s h eld e rect a n d

th e h a nds we re no l onge r used for l oc om oti on; whe re
the teeth m ost cl osely resembled th ose i nma n a nd speech ,

a ppe a rs to h a ve bee n feebly d evel oped ; wh ile the size

of the b ra i ni n dic a tes a deg ree of i ntellige nce much a b ove

th a t of th e a pes th ou gh i nfe ri or to ma n (Fig .

P I LTD O W N M AN Th e disc ove ry o f th e rem a i ns o f

.
-

a ve ry p rimitive typ e o f hum a nbei n g i nPiltd ow n s outh ,

o f L on d on wa s a nnou nced i n 1 91 3 Owi ng to the fa ct

,
.

th a t seve ra l imp ort a nt p orti ons of th e skull a re missi ng ,

its origi na l f orm a nd th e ge ne ra l a ppe a ra nce of the mem

be rs O f the ra ce a re s omewh a t u nce rt a i n I t is evide nt .
,

h oweve r th a t the b ra i nc a se is typic a lly hum a n th ough

,

u nusu a lly thick while th e f oreh e a d is high a nd with o ut

,

th e g re a t ridges a b ove th e eyes which ch a ra cte rize the

J a v a ma na p e a nd th e N e a nde rth a l ma n Th e l owe r
-
.

ja w on th e othe r h a nd is m ost emph a tic a lly a p e like

, ,
-
.

I nf a ct i t wa s a t on e tim e a ssumed by ce rt a i nc ritics th a t

th e ja w a ctu a lly is th a t o f a nexti nct ch imp a nzee Th is .

O bj ecti on h a s bee n rem oved by the rece n t disc o ve ry o f

a sec on d skele ton with th e s a me ch a ra cte ristic fe a tu res .

S o fa r a s scie ntis ts a re a ble to j udge the Pil td ow n ,

ma n wa s of l ow st a tu re with a t reme nd ous l owe r ja w

,

an d ret re a ti ng chi n but with a b ra i n c a p a city a t le a st

,

a s g re a t a s th a t of th e m ore p rimit ive ra ces o f m ode rn

ma n Th e a ge a t wh ich h e lived h a s bee nv a ri ously es

.

tim a ted from to ye a rs a go a nd th e f a c t ,

th a t h e u nde rst ood th e use of fire a nd f a s hi oned we a p ons

o f th e ch a se i n d ic a tes th a t th e h um a ns to ck wa s cle a rly
defi ned a t th a t rem ote pe ri od .

OT H ER T R A CE S OF M A N S A N T I Q U ITY
’
Th a t the .

h um a n stem is o f gre a t a ge i s a ls o evide nced by th e

 T R A CE S OF M AN S AN TI ! UIT Y
’
4 99

disc ove ry of c a mp S ites i n Engl a nd with t ra ces of fi re

an d a n a bu nd a nce o f fli nt impleme nts N o skelet a l re .

m a i ns h a ve bee ndisc ove red with the p ossible excepti on

,

o f a l owe r ja w but these o the r evide n

,
ces a nd th e ge ol ogi
ca l f orm a ti on i n which they o ccu r cle a rly p o i n t to the
fa ct th a t Engl a nd wa s i nh a bited by s ome typ e of h um a n
bei ngs a t le a st ye a rs a go .

In 1 91 7 th e skelet on of a no th e r a n cie nt typ e of ma n

wa s disc ove red i na c a ve inR h odesi a S outh Af ric a The,
.

ch a ra cte r of th e ge ol ogic a l f orm a ti on gives no clu e to

its ex a ct a ge but i n respect to th e f a ce especi a lly it is
,

on e o f the m ost p rimitive th us fa r disc ove red The .

ridges a b ove the eyes resemble th o se of a na dult g orill a ,

an d t ogethe r with ce rt a i nfe a tu res of th e nose a nd m o uth

must h a ve give n this ra ce a m ost besti a l a ppe a ra nce .

C once rni ng its rel a ti onship s little ca n b e s a id u nti l

qu a lified scie ntist s re nde r a ve rdict Th e size of th e .

b ra i n a nd v a ri o us othe r ch a ra cte rs suggest th a t it occu

pies a p ositi on s omewhe re betwee n the Piltd own a nd
N e a nde rth a l ra ces And th e re is a t le a st a rem ote p ossi
.

b il i ty th a t it ma y be cl osely rel a ted to the Heidelbe rg

ra ce which up to th e p rese n t time is know nonly f rom a
l owe r ja w disc ove red i n Ge rm a ny i n 1 907 .

T H E N EA N DER T H A L M AN —This species migra ted

.

(seemi ngly from Asi a ) i nt o E u rop e not less th a n

ye a rs a go a nd bec a me widely dist ributed th ro ugh ou t th e
,

western p orti on of th e c onti ne nt F rom nume rous .

skele tons it is evide nt th a t th e b ody wa s S h ort (4 ft 1 0 i n . .

to 5 ft 3 . th ick set with p owe rful sh oulde rs sh ort

-
, ,

a rms a n
,
d stubby fi nge rs a n d thumb s with less f reed om
of m o veme n t th a n i n m ode rn ma n Th e re wa s a ls o a .

pe rm a nent a p e like be nd to th e k nee a nd a hu nch to

-

the sh oulde rs th at very evide ntly resulted i na na wkw a rd

.

shuffli ng g a i t Th e he a d wa s rel a tively e norm o us with ,

gre a t ove rh a ngi ng eyeb rows mpi ng f orehe a d l ong uppe r

.

, ,

lip a nd ret re a ti ng chi n a ll of which must h a ve give n

,

th is ra ce a m ost repulsive a ppe a ra nce .

500 T HE E V O LUTI O N OF M AN

Th e f a milies of this pe ople m a de th ei r h omes in c a ves

o r ro ck shelte rs whe re th ey f a sh i on ed slings a nd spe a rs
wi th h e a d s of s to ne Wi th th ese we a p ons they p rob a bly
.

defe nded themselves a nd it a ppe a rs th a t they success

fully hu nted th e w oolly m a mm o th a nd rhi noce ro s the ,

c a ve be a rs li ons a nd le op a rds a nd seve ra l o th e r a nim a ls

,

which si nce h a ve bec om e exti nct .

While it is u nq uesti ona bly t ru e th a t thi s ra ce resem

bled the a pes in ce rt a i nrespects it is a m o st sig nific a nt ,

f a ct th a t they c a refully bu ried thei r de a d left f ood a nd ,

we a p ons ne a r them a nd ve ry evide ntly believed i n i m

m orta li ty Th is rep rese nts a v a st st rid e f orw a rd a nd
.

pl a ces th e Ne a nde rth a l ma na lm o st a t th e s a me level a s

th e Au st ra li a n a nd T a sm a ni a n na tive .

M ODER N M AN — N 0 t l ong si nce it wa s believed th a t

.

th e J a v a ma na p e the Piltd ow n ma n a nd th e N e a n
-
, ,

de rth a l ma nrep rese nted st a ges i n th e g ra du a l ev oluti on

o f m ode rnma n N ow it is th o ugh t th a t th ey a re me rely
.

o ff sh oot s f rom th e m a i n h um a n stem (Fig an d .

th ough th ey th row a fl ood o f l igh t on th e p roblem o f

m a ns ev olu ti on th ey a re not to be reg a rded a s h is di rect
’
,

an cest ors Th ese p rimitive types h a ve c ompletely dis

.

a ppe a red wh e re a s th e m a i nb ra n
,
ch h as pe rsisted a nd not ,

on ly h a s it c onti nued but a s ma nove rc a me b a rrie rs a nd

,

bec a me widely d ist ributed th rough ou t th e gl obe the ,

S pecies bec a me di ffe re n ti a te d i nt o th e p rese nt da y ra ces -

o r sub species
-
.S ome o f th ese pe oples like th e Au stra ,

li a ns a nd the rece ntly ex ti nct T a sm a ni a n ra ces neve r ,

ev olved i ntellectu a lly much bey ond th e N e a nde rth a l

St a ge ; o th e rs bec a me much m ore highly devel oped .

CR O M A GN O N M AN — Am ong the highly devel oped

-

ra ces o f th e m ode rn h um a n species th e C ro M a gn on -

h old s a p romi ne nt p ositi on Ev olved i n Asi a to th e

.
,

best O f o u r k nowledge i t e nte red Eu rope by wa y of th e

,

M edi te rra ne a n c oa st no t less th a n ye a rs a go ,

an d sp re a d o ve r p ra c tic a lly the s a me te rrit ory a s th a t

held by th e N e a nde rth a l pe ople Inge ne ra l a ppe a ra nce
.
 C R O—M AGN O N MAN 501

this l a te r ra ce wa s u nusu a lly t a ll (6 ft 1 i n onth e a ve r

. .

a ge ),
with b roa d f a ces l ong a nd na rrow S kulls a nd l ong
,

limbs c a p a ble O f g re a t freed om of m oveme nt Th e b ra i n .

c a p a city su rp a ssed th a t of th e a ve ra ge ma nof to da y -

.

Pf fi lf M
'

0 new

FI G 1 43 . Gen
. ea l o gi ca l T ree , ill u s tra t i ng p rob a b le igi nof ma n
or

and th e a pe s D a ta l a rgel y f rom K ei th

. .

Th e Cro M a gnon p e ople were s kill ed i n th e use of

- ‘

t ools thei r we a p ons a nd orna me nts displ a ying a h igh de

,

gree of w orkm a nsh ip a nd a rtistic a bility T h ey a ls o d rew .

rem a rk a bly we l l a s th e p a i n
,
tings on th e w a lls of m a ny
c a ves be a r wit ness Their bu ri a l cu st oms i ndic a te a
.

belief i n life h e re a fte r Fu rth e rm ore it a pp ea rs to be

.
,

f a irly well est a blish ed th a t th ey c on

,
que red the N e a n
5 02 THE E V OLUTI O N OF M AN
derth a l pe ople a nd the re a re no i ndic a ti ons th a t i n tu rn
,

they fell bef ore a supe ri or foe N eve rtheless a b out .

ye a rs a go th e ra ce g ra d u a lly declined le a vi ng no ,

desce nd a nts with the p ossible excep ti on of a few pe ople

now livi ng in s ou th e rnF ra nce B ritt a ny a nd p ossibly in , ,

the Py re nees .

L ATER EU RO PEA N R A CES N e a r the cl o se o f th e C ro

.

M a g non pe ri od seve ra l ra ces of pe ople mig ra ted f rom .

Asi a i nt o E u rope a nd b eca me di ff erenti a ted i nt o th e

'

pe oples living the re to da y One st re a m c onstitut i ng the

-
.
,

so c a lled M edi te rra n

-
e a n type fl owed a l ong the s ou the rn ,

c oa s t exte nded northw a rd a l ong the A tl a ntic se a b oa rd

,

an d e nte red the B ritish Isles Their desce nd a nts a re .

rep rese n ted by m a ny G reeks It a li a ns S p a ni a rds P ortu , , ,

g u e se a,
nd E nglishme n o f sh or t st a tu re d a,
rk c o mplexi on , ,

b row n eyes a nd l ong he a ds Th e Alpi ne t ype mig ra ted .

by wa y of th e Bl a c k S ea a nd o ccupied C e nt ra l E u rope .

Th ey a re rep rese nted by m a ny G reeks Tu rks A rme ni a ns , , ,

S outh G e rm a ns N orth I t a li a n
,
s m a ny F re nch a nd S wiss ,

an d othe r inh a bit a nts o f th a t ge ne ra l regi on Inst a tu re .

—
these pe ople a re sh ort thick set with d a rk eyes a nd ski n
, ,

an d ro u nd h e a d s Th e N ordic type j o u rneyed by wa y o f

.

R ussi a i nt o N orthe rnE u rope InO lde n d a ys they we re .

th e Pe rsi a ns D ori a ns Th ra ci a ns G a uls G o th s V a nd a ls

, , , , , ,

F ra nks D a nes A ngles a nd N orseme n Th ei r m ode rn

, ,
.

desce nd a nts a re the Sc a nd ina vi a ns D a nes s ome R ussi a ns , , ,

N orth G e rm a ns a nd m a ny i nh a bit a nts O f th e B ritish

Isles This type is ch a ra cte rized by g re a t st a tu re f a ir
.
,

h a i r a nd blu e eyes Thus th e E u rope a ns to da y bel ong

.
-

to th ree disti n c t ra ci a l g roups th e M edite rra ne a n th e , ,

Alpi ne a nd th e N ordic
,
.

M AN I N A M ER I CA —Th e fi rst imp ort a nt f a c t which

.

a ppe a rs i nc on necti onwi th th e study o f ma nin Ame ri c a

is th a t a l l o f th e na tive t ribes bel ong to one typ e Th e .

E skim o a n d Aleut o f Al as k a th e v a ri ous I n di a n t ribes ,

o f b o th N orth a n d S outh Ame ric a th e Aztecs O f M exic o , ,

the M a y a s o f C e nt ra l Am e ric a a nd the I nc a s o f Pe ru ,

 S U B S P E CI E S
-
OF M O D E R N M AN 5 03

a re a ll f a i rly cl o sely rel a ted m odific a ti ons o f a si ngle

st re a m or b ra nch o f the h um a n ra ce This c onclusi on .

rests n o t on ly on physic a l me a su reme nts bu t on m a ny ,

s imil a rities rel a ti ng to t ools we a p ons buildi ngs cust oms , , , ,

l a ngu a ge a nd lege nds .

It a ppe a rs the ref ore p rob a ble th a t from time to time

a dve n tu ro us Spi rits i n se a rch of new h u nti ng g rou nds
, ,

eithe r a ctu a lly c rossed a l a nd b ridge u niti ng Asi a wit h

Al a sk a or t ra velled in c a no es Whe n th e fi rst mig ra .

ti on occu rred is u nknow n Th e Oldest h um a n rem a i ns ;

in N orth Ame ric a do no t a ppe a r to be olde r th a n rece nt

gl a ci a l times a nd a cc ordi ngly ma y not be ove r
,

ye a rs ol d In S outh Ame ric a the re is no c onclusive

.

evide nce o f ma ne a rlie r th a n ye a rs a go .

S U B S P E CIE S O F M ODER N M AN — F rom th e p ictu re

-

d ra w n i n the foreg oi ng p a ra gra ph s it is see n th a t the ,

h um a n ra ce eme rged p e rh a p s ye a rs a go ; th a t
it g ra du a lly ev olved i ndi ffere nt di recti ons ; a nd th a t a ll
of th e b ra n ches with one excep ti onh a v e bec ome exti nct .

Th is excep ti on rep resents th e S i ngle species o f m ode rn

ma n which l ong a go a ls o u nde rwe nt ev oluti ona ry
,

ch a nges resulti ng a cc ordi ng to one V iew i n th e p roduc

, ,

ti on of fo u r sub species s ometimes te rmed races

-
.

Th e fi rst of th ese is th e Au st ra li a n r ep rese nted by

th e ch oc ol a te c ol ored t a ll a nd l ong limbed na tives of
-
,
-

Aust ra li a a nd H i ndust a n Th e sec ond sub species is th e .

-

N eg roid with fl a tte ned nose a n d w oolly h a i r a nd bl a ck

or ch oc ol a te c ol ored skin Th e ra ce s o f M a d a g a sc a r a n
-
. d
A fric a s outh of th e S a h a ra bel ong to this g roup Th e .

th i rd or M ong oli a n sub species i s Sh ort thick set with -

,
-

obliqu e eyes c oa rse bl a ck h a ir a nd g olde n or reddish

,

b row n ski n Th is type is widely dist ributed ove r Asi a

.

an d P olynesi a a nd lik ewi se i ncludes th e na tive ra ces of

the N ew W orld Fi na lly the re rem a i ns the C a uc a si a n
.
, ,

with white or Sligh tly d a rk S ki n wh ose divisi oni nt o th e ,

M edite rra ne a n Alpi ne a nd N ordic h a s bee n me nti oned

,

i na p revi ous p a ra gra ph .

504 T HE E V OLUTI O N OF MAN
M A N S F UT U RE
’
C once rning th e futu re of the h uma n
.

ra ce n o on e ca n S pe a k with a u th ority Owi ng to h is .

supe ri or me nt al devel opme nt inve ntive inge nuity a nd

,

S pi rit o f coOp era ti on ma nh as bee na ble i np a s t tim es to

,

o ve rc ome b a rrie rs a n d mig ra te ove r the e a r th a n d live

a mid su rro u n di ngs th a t w o uld h a ve a nni h il a ted a less
i nte llige nt species B rute st re ngth a nd kee nness of
.

se nse a re no l onge r th e imp ort a nt f a ct ors they once we re

in the st ruggle for existe nce a n d p rob a bly th ey a re no t
a s highly devel oped a s f orme rly A h a i ry c ove ring for
.

p rotecti onis no l onge r a necessity a nd i t is m ore sp a rse

th a ni na ncie nt ma n Th e teeth a re sm a lle r a nd we a ke r
.
,

t h e size o f th e t oes with the excepti ono f the fi rst digit

, ,

is red uced a nd m a ny o th e r org a ns a re m ore ru di

,

me nt a ry Th e b ra ina ppe a rs to be i nc re a si ng i nsize a nd

.
,

S kill i n th e u se o f th e h a n d ma y likewise be d evel op ing

a s a resul t o f m ore pe rfect a d j ustme n ts of b ones a nd
m uscles but bey ond th is p oi nt no p rog ressive ch a nges
,

a re k n ow n to be t a ki n g p l a ce .

Wh ethe r ma n w ill ultim a tely bec ome ex ti nc t a nd a ,

new a nd pe rh a ps m ore h ighly devel oped species t a ke his

pl a ce is a much deb a ted a nd i ns oluble q uesti on E ve ry .

high est species i nth e p a s t h a s bee nrepl a ced by m ore

c ompete nt bette r a d a pted types a nd i t ma y ve ry well
, ,

h a ppe ni nth e a ges to c ome th a t a we a ke ned c onsti tu ti on ,

u nfa v ora ble mut a ti ons new p a ra s itic dise a ses or s ome
,

o th e r a ge ncies w i ll forever r emove ma nfrom th e f a ce of

the e a rth .
 C HAPTER XLI

H U MA N I NH E R I TAN CE

H U M A N T R A I T S — M a n like a ll o the r a nim a ls is

.
,

a c omp osite of m a n y ch a ra cte rs which depe nd on the

,

p rese nce of defi nite ge nes or f a ct ors i n the ge rm cells

which u nite a s the fi rst step i nh is being Th e m a nne r .

i nwhich the ge n es b ri ng a b ou t the devel opme nt of thei r

“

c orresp ondi ng ch a ra cte rs is h oweve r i n m a ny c a ses

, ,

m ore c omplic a ted i nma nth a ni na V i neg a r fly or a pl a nt .

It h a s a l re a dy bee n sh ow n i n previ ous ch a pte rs (X V I ,

X V I I ) th a t ce rt a i nch a ra cte rs depe nd f or thei r exp ressi on

onth e a cti on s of h orm ones Obvi ously i nth ese c a ses the
.

f a ct ors wh ich c ont rol th e devel opment o f th e h orm one

p roduci ng gl a nd s will a lso i ndi rectly c ont rol othe r m ore
obvi o us ch a ra cte rs It h a s rece ntly bee ncl a imed th a t a
.

l a rge numbe r of h um a nch a ra cte rs a re dete rmi ned i nthis

indi rect fa sh i on th ough this rem a i ns yet to be p roved
,
.

On th e o th e r h a nd a g re a t m a ny ch a ra cte rs a ppe a r to
n
v a ry m ore or less c o ti nu ously th rough a wide ra nge .

Ex a mples of this s ort of qu a ntit a tive v a ri a ti on h a ve

a l re a dy bee n ex a mi n ed i n pl a nts a nd a nim a ls (Ch a ps .

XXX I— XXXI I I ) a nd sh ow n to depe nd on wh a t a re

c omm only c a lled cumul a tive f a ct ors ” .

D I FF I CU LT IES OF S TU DY I N G H U M A N I N H ER I TA N CE .

Th e fa ct th a t m a ny ch a ra cte rs i n ma n a re c ont rolled

i ndi rectly th rough h orm one sec reti on a nd m a ny m ore
th rough cumul a tive f a ct ors m a kes th e study of m a ns i n
,
’

he rit a nce difficult These a re not h oweve r th e only di fli

.
, ,

cu l ti es for th e chief on
,
es a re c once rned with th e i mp os
si b ili ty of c on ducti ng c ont rolled expe rime nts onma n In .

the fi rst pl a ce no one lives l ong e nough to Obse rve much

,

50 5
506 HU M AN I NH E R ITAN CE
-

m ore th a n his o wn ge ne ra ti on He ma y to be su re .
, ,

o rdi n a rily obse rve h is o wnp a re n ts a nd his ownch ild re n ,

.

an d m o re ra rely h is g ra n dp a re nts a nd g ra ndchild re n At .

m o st h e ma y h ope to see a p a rt of f ou r or five ge ne ra ti ons ,

c omm only t o t a ling fewe r th a n a h u nd red i ndividu a ls .

M ore ove r h e will h a ve p ra c tic a lly no c ont rol o ve r the

v a ri ou s m arri a ges a nd this will re nde r h is me a ge r Ob
,

serv a ti on s o f still less v a lue H oweve r no twithst a ndi ng

.
,

a ll this the re a re s ome h um a nt ra its which a re su ffi cie n

,
tly
st riki ng a nd simple th a t it h a s bee np ossible to d ete rmi ne
wi th ce rt a i nty h ow they a re i nhe rited a nd to sh ow th a t ,

they beh a ve i nthe s a me wa y a s sim i l a r ch a ra cte rs i nth e

an ima ls c omm only used i n ge netic expe rime nts
‘

H ER ITA B LE C H AR A CTER S I N M AN In pl a n ts a nd
.

the l owe r a nim a ls p ra ctic a lly a ll ou r knowledge of he red

,

ity necess a rily c once rns physic a l t ra its Inma n h oweve r .

, ,

we a re c once rned with me nt a l t ra its a ls o I t is i ndeed .

p rob a ble th a t k nowledge O f the wa y i n which g ra des o f

i ntellige nce ma y be a ccu ra tely me a su red a nd O f the ma n
ner i nwh ich they a re i nh e rited is of g re a te r v a lue th a n
an y o the r s ort O f k n owledge wh ich we ma y eve r h o pe

to a tt a i n f or o n i t d epe n
,
d s the s ole p ossibility o f a ny
exte nsive imp roveme nt o f th e hum a n ra ce Physic a l .

t ra its a ppe a r h oweve r to be simple r I t will be desi ra ble

, ,
.
,

the ref ore to t re a t b riefly i nthis ch a pte r o f s ome physi

,
~

ca l t ra its wh ich a re eith e r kn ow n to O bey th e ord in a ry

l a ws of he redity or ma y re a s o na bly be supp osed to do so

o n th e b a sis o f th e i n c omplete d a t a a v a il a ble a nd th e n ,

to c on side r the i nh e rit a nce o f me nt a l t ra its i n th e next

ch a p te r .

P H Y S I CA L C H AR A CT ER S I t is a ni nte resti ng f a c t th a t
.

ma np ossesses ve ry few disti nctly m a rked physic a l t ra its .

Alm ost a ll o f th em v a ry th ro ugh a wide ra nge with out

an y sh a rp li ne o f sep a ra ti o n Thus the re
. a re bl o ndes
an d b ru nettes fa t a nd le a n pe ople st ra igh t h a i red a n
, ,
d -

cu rly h a i red t a ll a nd sh ort st rong a nd feeble a nd a l l

-
, , ,

gra d a ti ons betwee n C e rt a i n m ore o r less a b norm a l

.
 A PI G M E NT o r O R G
.

R O I D C O AT A N D P I GM E N

OF I RIS A BS ENT . 1 . Th
‘

ALB I N O ey e . fro r
R ed
u o n bs cu red bloo d vessel:

B PI M E N T O F C B C
. G
RO ID PR E S E N T .

a IR I S W IT HO UT T R U
.

G
PI M E N T 2 B LU E Du . . .

to a pu rpl e l a y er onb ac
of ey e .

6 IR I S
. W IT H TR I '

G
PI M E N TS .

a Li pochrome or yelloz
.

pi gmen t 3 GR E E N o r ca . .

eye Y ello w pigment

. 0

blu e b a c kgro u nd .

I) . M clu ni e or bla ck p i g
ment . 4 . HAZE L or gra ,

ey e . b ro wn p ig Di lu t e
ment a ro u nd pupil only .

5 BRO W N
. ey e . M el a ni
pigment ; v a rio us sh a de
fro m v a ri o us dilu tions .

6 . BL AC K eye . A
abu nd a nce of mela ni c pig
ment .
 T HE IN H E R I TAN CE O F Y —
E E COL O R 5 07

t ra its do S h ow a sh a rp disc onti nuity a nd a re c onseque ntly

m ore e a sily studied Alt ogeth e r a b o u t fi fty t ra its h a ve
.

bee nsu fli ci entl y studied to k now th a t th ey a re i nhe ri ted

an d i n s ome c a ses to k now defi nitely h ow Am ong nor .

ma l ch a ra cte rs ma y be me nti oned : c ol or of h a i r eyes , ,

an d ski n; f orm o f h a ir whethe r st ra igh t cu rly or ki nky ; , ,

st a tu re p rop orti on a nd weigh t of the b ody H a nds a nd

, , .

feet with two j oi nted fi nge rs or t oes or with six digits

-

i n pl a ce o f five web feet c ol or bli nd ness (i na bility to

, ,

disti nguish red from g ree n m o st c omm on) nigh t bli nd ,

ness (i na bility to see i n dim light ) h oemop h i l i a (a cer ,

t a i ntype Of pe rsis te nt bleedi ng ) a nd c reti nism (a dise a se ,

due to deficie ncy O f th e thy roid gl a nd h orm one ) a re

a m on g th e bette r k now n .

T H E I N H ER I TA N CE O F EY E C OL OR — The c ol or o f the -
.

eye (Fig 1 4 4 ) depe nds on pigme nt g ra nules dep osited

.

i n th e i ris All eyes h a ve s ome p igme nt on the b a ck

.

S ide o f the i ri s which pigme nt whe n p rese nt a l one

,

gives pu re blue eyes N onbl ue eyes h a ve i n a ddi

.
-

ti on to this pigme nt m ore or less b row n p igme nt i n

th e cells f ormi ng th e front side o f th e i ris Th e v a ry .

in g sh a des o f eye c ol or f ro m g ra y-
t o bl a ck dep en d o n
th e a m ou nt a nd dist ributi on of this sec ond pigme nt on
th e front I f a blue ey ed p e rs on m a rries a pu re d a rk
.
-

eyed one a ll th e child re n will h a ve d a rk eyes If these .

d a rk eyed hyb rid ch ild re n m a rry blue eyed m a tes one

- -

h a lf th ei r ch ild re n th e F ) will be d a rk eyed a nd 2

-

on e h a lf blu e eye d I f th e d a rk e yed hyb rid s m a rry

- -
’ -
.
!

o the r d a rk eyed hyb rids-

th e offsp ri ng of a blue
eyed by a d a rk eyed p a re nt ) th ei r child re n
-
th e F ) 2

—
will S h ow th ree d a rk eyed to one blue eyed a pp roxi -
,

m a tely This is cle a r a nd su fficie nt p roof to sh ow th a t

.

th is eye c ol or di ffe re nce depe nds on only one p a i r of

-

f a ct ors or genes I t beh a ves ex a ctly like c oa t c ol or i n

.
-

gui ne a pigs or fl owe r c ol or i npl a nts .

B RA CH Y DA CTY LY is a c onditi on o f th e h a nds i n which

they a ppe a r p rop orti ona lly V e ry b roa d i n c omp a ris on to
‘
508 H UM AN I NH E R I TAN CE
thei r le ngth (Fig . I t is d ue to the fa ct th a t the fin
ge rs h a ve only two j o ints i npl a ce o f the usu a l th ree A .

b ra chyd a ctyl ous individu a l ordi na rily t ra nsmit s the p e

culi a ri ty to a b o ut one h a l f o f h is ch ild re n whe nm a ted to
-
,

a n orm a l w o m a n H is a ff ected chil d re nt ra n

. smit it i nthe

FI G 1 4 5
. . B ra chyd a ctyl o us Ha nds .1 , sh ows th e p ro port i o n
a ll y

sh o rt b roa d h a nds a nd 2, a n X- ra y ph o to gra ph o f a norm a l a nd

b ra chyd a c tyl o us h a nd N o te th a t i n the l a tte r th e l as t two bo nes
.

o f e a ch fi n ger a re fu se d to ge th e r to fo rm a si n gle sh ort one

—
.

Pla te, Vererbu n gslehre Engl em a n

. n .

s a me wa y bu t th e norm al child re np rob a bly neve r t ra ns

mit it to th ei r desce nd a nts F rom th ese d a t a one ma y
.

c onclude th a t a si ngle fa ct or di ffe re nce is resp onsible a nd

th a t th e ge ne fo r b ra chyd a c tyly is d omi na nt ove r the
c orresp ond ing norm a l ge ne I nthe c a ses o f wh ich the re
.
 P O L Y DA C T YL Y AND S Y NDA CT YL Y 5 09

a re r ec ords it w ould a ppe a r th a t the b ra chyd a c tyl ous

i ndividu a ls we re hyb rid si nce they give two equ a l cl a sses
o f desce n d a nts wh e n m a ted i e b a ck c ro ssed to the ,
. .
,
-
,

norm a l recessive D oubtless i ndividu a ls ma y occu r wh o

.

a re pu re f or b ra chyd a ctyly an d w ould h a ve a l l thei r

child re n a ffected i e p roduce a u nif orm F
,
. .
, 1 .

P OLY DA CTY LY AN D S Y N DA CTY LY (Fig —In th e .

f orme r th e re is a next ra fi nge r or toe I t not only occu rs .

i nma nbut is a ls o well k now ninp oult ry -

Th e ge ne for .

this ch a ra cte r is d omi na nt ove r norm a l a nd i s t ra nsmitted

usu a lly to a b out h a lf the offsp ri ng bec a use th e a ffected ,

0
FI G 1 46
. . In h eri tance of S u pernum era ry D igi ts . Aff ecte d ind i
vi du a ls a re indi ca te d by d a rk symb ols a nd u na ff ecte d by ligh t o nes .

F i ndi ca te s e xt ra fi n gers a nd Z e xtra toe s Fem a le s . a re in di ca ted

by ! a nd m a l es by
Pla te, Vererbu ngs lehre . Engl em a nn .

p a re nt is ordi na rily a hyb rid It occ a si ona lly h a ppens

,

th a t a pp a re ntly norm a l recessive ) off sp ri n g t ra nsmit

the a b norm a lity to th eir ch ild re n This w ould i ndic a te .

th a t othe r fa ct ors or c onditi ons a re c once rned i nth e ex

p r ess i on o f this ch a ra cte r I n sy nd a ctyly th
. e fi nge rs or

t oes a re u nited side by side hence th e c omm on te rm

web f oot
-
” .
,

Th is c onditi oni s a ls o due to a si ngle d omi

na nt f a ct or diffe re nce Aff ected i ndividu a ls (usu a lly b e
.

in g hyb rid ) t ra n smit it to a pp ro xim a tely h a lf thei r

offsp ri n g a nd the norm a l child ren (being pu re reces

,

si v es ) do n o t t ra n smit it .

S K I N COL OR p rese nts a m ore c omplic a ted situ a ti ona nd

51 0 HU M AN I N H E R I TAN CE
o ne which is no t yet ce rt a i nly or fully u nde rst ood Th e .

f a ct a ppe a rs to be th a t a hyb rid betwee na light skinned

Ca uc a si a n a n d a d a rk ski n ned N eg ro is usu a lly a b ou t
midw a y betwee nthe p a re nts i nc ol or (Fig D a rk .

ski n a ppe a rs th us to be like the red c ol or i n the f o u r

,

o cl ock desc ribed i n C h a p te r XX V I I I on

’
ly p a rti a ll y ,

d omi na nt ove r light H oweve r wh e n the pi nk fo u r

.
,

o cl ocks a re selfed o r i n b red the F c onsists of 1 red

’

2 pi nk : 1 wh ite wh ile the medium c o l ored mul a t

,

t oes whe n i nb red do not seg reg a te cle a nly int o

FI G 1 4 7
. . In h eri ta nce of S ki nC ol or a nd Ha ir Form T he f a mily .

of a kin ky h a-
i re d A f ri c a n N e gro a nd a s t ra igh t-
h a ire d Engl ish
w om a n No te th e d omi na nce of the ki nky ov er s tra igh t h a i r a nd
.

th e v a ryi ng col or of the child rendue to the f a ct th a t the d a rk p a re nt

wa s no t pu re for a ll th e co l o r gene s .

Pl a te , Vererbu n gs le hre
. Engl em a nn .

3 cl a sses i n th e ra ti o In f a ct th ey p rod uce

.
,

seve ra l sh a des o f c ol or o f wh ich i t ra rely h a ppe ns

th a t a ny a re ei th e r a s ligh t or a s d a rk a s th e g ra ndp a re nts .

This result st rongly resembles th a t obt a i ned i n the c ross

o f th e red g ra i n ed wh e a t by th e white g ra i ned one
- -

desc ribed i n Ch a pte r XXX I Th e cu rre nt expl a na ti on

.

a s sumes th a t th e N eg ro a n d th e C a uc a si a n ra ces d iff er

 S TATU R E AND W E IGHT 51 1

by two ch ief p a i rs o f f a ct o rs or ge nes neithe r of which is

fully d omi na nt If th e re a de r will w ork ou t such a
.

c ross by me a ns of the Pu nnett squ a re h e will fi nd th a t

this will a cc ou nt for ni ne di ff e re nt ge no types or sh a des
of c ol or th a t migh t be expected i nthe sec on d ge ne ra ti on .

If th e d om i na nt c ol or f a ct ors p rese nt i n th e N eg ro be
rep rese n ted by A a nd B of wh ich A p roduces m ore c ol or
,

th a n B a nd th ei r recessive ge nes i n the Ca uc a si a n

,

by a a nd b th e nth e mul a tt o o f i nte rmedi a te c ol or w ould

‘

be Aa B b Ge no typ e s o f this c onstituti on whe n i nte r

.
,

b red w ould be expec ted to p roduce a n F ge ne ra ti on

, 2

c onsisti ng O f 1 AAB B (like th e Origi na l N eg ro p a re nt ) ,

2 AAB b (a sh a de lighte r ) 2 Ad BB (slightly lighte r still )

‘

, ,

1 AAbb (still ligh te r ) 4 Aa B b (i nte rmedi a te or mul a tt o )

, ,

1 a a BB 2 Aa b b 2 a a B b (o f decrea smg c ol or g ra des ) a nd

, , ,

1 a a bb (like origina l C a uc a si a n p a re n t ) ou t of sixtee n ,

.

Whe nit is remembe red th a t c ol or is not u nif orm i neither

p a re nt ra ce owi ng to m odifyi ng f a ct ors i t a ppe a rs p rob ,

able th a t this hyp o th esis is suffici e nt to expl a i n a l l th e

dive rsity of c ol or a c tu a lly f o u nd i n these i nte rra ci a l
hyb rids .

S TATU RE AN D WEI GH T a re ve ry evide ntly h e redit a ry

(Fig 1 4 8) but o wi ng to the gre a t i nflue nce of th e e nvi ron
.

me nt i t h a s not bee n p o ssible so f a r t o dete rmi ne with

,
o

s a tisfa ct ory a ccu ra cy th e numbe r of f a ct ors c once rned .

Ce rt a i n f a milies a nd ra ces a re u n usu a lly t a ll a s for ex ,

a mple th e n a tives of P a t a g on i a a nd S c otl a nd while ,

o the rs for ex a mple Wels h or E skim o s a re u nusu a lly

, , ,

sh ort S ince these ra ces m a i nt a i nthei r ch a ra cte ristic st a t

.

u re eve nwhe nlivi n

, g inothe r c o u nt ries it is O bvi o us th a t ,

the c a uses c ont rolli ng h eigh t a re chiefly h e redit a ry I n .

f a milies wh e re t a ll h a s m a ted with t a ll th e child re n a re

m ostly t a ll S ome h oweve r a re sh ort a s w o uld be ex
.
, , ,

p e cte d f ro m th e f a ct th a t b o th p a re nts w o uld ra rely be

pu re f or a ll th e f a ct ors for t a llness Hum a nfa milies a re .
,

h oweve r too sm a ll to e na ble one to dete rmi ne th e nu m

,

b er of f a ct ors c once rned Wh a t h a s bee ns a id for st a tu re

.
51 2 H UM AN I N HE R I T AN CE
seems to h old for weigh t a l so I t is eve n m ore difficult
.

to a sce rt a i n th e f a ct ors g o ve rning weigh t si nce i t is so

g re a tly i nfl u enced by di et a nd exe rcise .

COLOR B LI N D N ES S se rves well to i llust ra te sex li nked

- -

inhe rit a nce i n ma n Th e re a d e r h a s a l re a d y le a rned

.

th a t the re a re m a ny sex l inked ch a ra cte rs i nthe vineg a r

-

fly wh ose ge nes or f a c t ors a re c a rried in the so c a lled

,
-

FI G .1 48 .
—
D w a rfs At th e righ t is a ma n o f n
. orma l s ta t re u .

The n ex t t wo fig res a re th o se o f dw a rfs wi th v ery sh o rt l e gs b u t

u
wi th ne a rly norm a l tr nk The nex t two figu re s a re of dw a rfs wh ose
u .

b odies a re p ro p ort i ona lly red ce d th rough o u t Th e g re to the

u . fi u
le f t is a dw a rf (cre ti n) wh o se u nderde veIO pment is due to the f a ilu re
o f a du ctl e ss gl a nd to p rodu ce the norm a l a m ou nt O f h orm o ne s .

Pl a te , Vererbu ngs lehre Engl em a n

. n .

sex ch m os omes
-
ro The typ e of c l bli nd ness i n o o r-

wh ich the a ff ected pe rso nis u na ble to disti nguish red a nd

g ree nis fa i rly c omm on(a b ou t one pe rs oni ntwe nty five ) -

a m on g men bu t ra re a m ong w ome n Whe n a c ol or .

bli nd ma nm a rries a norm a l w om a nhe t ra nsmits to h is

 C R E TIN I S M 51 3

d a ugh te rs the sex ch rom os ome c ont a i ning the recessive

-

f a ct or for c ol or bli nd ness but th e d a ugh te rs a re not

-
,

c ol or bli nd bec a use th ey h a ve a ls o received a sex ch rom o

- -

s ome wi th th e d omina nt norm a l f a ct or from th e m o the r .

S i nce the hum a n fem a le h a s two sex c h rom os o mes a ll -

eggs will be a like a nd a lw a ys h a ve a sex -ch rom os ome

’

B ut onthe othe r h a nd th e m a le h a s only one se x ch rom o -

s ome a nd the ref ore p roduces two kinds of sperms one

, , ,

kind with th e sex ch rom o s ome a nd th e o the r l a cki ng it

-
.

The s ons i n the f a mily j ust menti oned receive thei r

'

si ngle sex ch rom os ome f rom th e 1 r m o the r a nd a re con

-

seque ntly free f rom a ny t ra it of c ol or bli nd ness Sh oul d -

.

the norm a l a pp e a ring d a ugh te rs m a rry norm a l h usb a nds

-

a ll thei r d a ughte rs w o uld be a ble to dist in guish c ol ors but ,

h a lf of them w ould be hyb rid l ike thei r m o th e r H a lf .

‘
the s ons w o uld receive a c ol or bli nd sex ch rom os ome f
- -

an d h a lf a n orm a l on e Th e fi rst w ould of c o u rse be

.
, ,

c ol or bli nd like thei r g ra ndfa the r a nd the sec ond w ould

-
, ,

be st rictly norm a l .

CR ET I N I S M is a na b norm a l st a te of th e h um a n species
due i ns ome ea ses a t le a st to a deficie ncy O f the h orm one
, ,

p roduced i nth e thy ro id gl a nd i nth e neck The i ndivid

“

u a l a ffected is likely to be d wa rfed i n st a tu re a n d dis

torted i n fe a tu res a n d to h a ve a ve ry l ow g ra de i n tell i
g e nce . I t is know n th a t s ome forms a t le a st of , ,

c reti nism a re he rit a ble P resum a bly th e h e redit a ry

, .

f a cto rs di rectly c ont rol th e devel opme nt of th e thyroid

gl a nd a nd th rough it p roduce th e Obse rved e ff ects I t .

h a s rece ntly bee ncl a imed by a c ompete nt bi ol ogist th a t

the re is evide nce i ndic a ting th a t m a ny ra ci a l ch a ra cte rs
depend on h orm one a cti on I f this sh ould be sub sta n .

ti a ted by fu rth e r i n vestig a ti on these h e redit a ry t ra its ,

w ould al s o be c ont rolled in th e s ame i ndi rec t f a shi on a s

c retinism .
 C HAPT ER X LII

I N H E R I TAN CE O F M E N TAL T R AI T S
(B Y L E W IS M TE R M AN )
.

M EN TA L As W E LL AS P H Y S I CA L T RA I T S ARE H ERITAB LE
— Th a t me nt a l a s well a s physic a l t ra its a re he rit a ble
.

is i ndic a ted by a ll th e a v a il a ble evide nce This i s wh a t .

we sh ould expec t c onside ri ng th a t ment a l fu ncti ons

,

a re depe n de nt up on physic a l st ructu res na mely th ose , ,

o f th e ce n t ra l ne rv ous system .

H oweve r we k now much less a b out me nt a l th a na b out

,

physic a l i nhe ri t a nce This is p a rtly due to th e f a c t th a t

.

t ra i ts of mi nd c a nno t be id e ntified a nd me a su red with

an ythi ng like th e a ccu ra cy wh ich is p ossible i n the c a s e
o f heigh t weight ski nc ol or etc
, ,
N eve rth eless c onside r
,
.

a ble p rog ress h a s bee n m ade i n the m e a su reme n t O f in

telli gen ce a n d o f a few o f th e m ore speci al ized m e nt a l
a bilities .

A M O U N T AN D N ATU RE O F VA RI A B I LI TY I N I N TELLI
GEN C E — M e n
. t a l tests h a ve sh ow n th a t u nselected i n
di v i du a ls o f a give nra ce diffe r e no rm ously i nth ei r i n tel
l i gen ce ,
an d th a t th ese d iffe re nces a re c onti nu ous .

Fi gu re 1 49 wh ich gives th e d i st ributi o no f i ntellige nce “

” ,

qu otie nts o f ni ne h u nd red a nd five u nselected C a li

forni a sch ool child re n illust ra tes b oth o f these f a cts
,
.

The i ntellige nce qu otie nt is a ni ndex o f b right ness The .

qu otie nt 1 00 rep rese nts a ve ra ge b righ tness The high es t .

g ra de feeble mi nded h a ve a qu o tie nt O f 6 5 or 70 the

-
,

me rely dull a q u otie nt o f 80 or 85 Child re n b righte r .

th a nth e a ve ra ge tes t a b ove 1 0 0 a b ou t one i ntwo h un

,

dred fi fty testi ng a s high a s 1 40 .

51 4
 VA R IA B ILI T Y I N IN T E LLIG E N CE 15

I t will be noted th a t a m ong these 905 chil dre n th e

intellige nce qu otie nts c ove r th e ra nge f rom bel ow 60 to
a b ove 1 4 0 th a t is f rom f eeble mi n
, ,
ded ness to a ve ry high -

deg ree of s upe ri ority Th e b rightest ch ild in th e g roup

.

is a s fa r a b ove the a ve ra ge a s the dullest of th e g roup is

bel ow I t will a ls o be noted th a t a s we go f rom the
.

a ve ra ge to w a rd eithe r ex t reme th e n umbe r of c a ses de

c re a ses g ra du a lly The re a re fa r m ore c a ses j ust a b ove
.

53 65— —
66 7 5 7 6—
85 86- 95 96 1 05— 1 06- 4 1 5 1 1 6—1 15 —
1 1 6 1 35 —
1 36 1 4 5
. 33 % . 55 %

FI G . 1 49

(R ep roduced from Te rm a n Th e M ea su rement of ,

I nte lli gence ) .

th e b orde rli ne o f feeble mi nded ness th a n i n the s a me

-

ra n ge j ust bel ow it a nd m ore c a ses o f m ode ra te th a n O f

,

ext reme b righ t ness The re is no sh a rp dividi ng li ne

.

betwee n a ny two gra des O f me nt a l a bility The e nti re .

ra n ge of h u ma n i ntellige nce is from p ro f ou nd idi ocy ,

which is rep resented by a n i ntellige nce qu o tient of less

th a n 1 0 to a degree of supe ri ority r ep rese nted by a n
,

i ntellige nce qu o tie nt of 1 80 or 1 90 Th e g re a test ex .

t remes of i ntelligence we re not f ou nd i n th is g roup of

90 5 child re n .
51 6 I N H E R ITAN CE OF M E NTAL T R AIT S

VARI AB I LI TY I N OT H ER “ A s n — Tests O f music a l

a bility sh o w simi l a r ra n ges o f diff e re nces for such t ra i ts
a s disc rimi n a ti ono f p itch se n se O f rhythm tona l mem
, ,

ory a n
,
d a pp reci a ti onO f c ons on an ce ; a nd th a t th e v a ri a
ti on i n these t ra its a ls o is c onti nu ous I t is p rob a ble .

th a t if we c ould me as u re such t ra its a s conscie nti o us ness ,

will p owe r s a nity or s oci a l a d a p t a bili ty w e w ould fi nd

, ,

th e ra nge o f i ndividu a l d iff e re nces a s g re a t a s for i ntel

li gen ce a nd m usic a l a bility a nd th a t th e diff e re nces f ou n
, d
w o uld be c ontinu ous Wi th me nt a l t ra its it is ra rely
.

a questi on o f simple a bse n ce or p rese nce bu t ra the r o f ,

p rese nce i ng re a te r or less deg ree The i nfe rence is th a t .

such t ra its do not depe nd up on si ngle f a ct ors i n the

ch rom os omes bu t up on seve ra l f a c tors I ntellect l i ke
,
.
,

h eight is pe rh a p s i nre a lity a c omp osite of seve ra l m ore

,

eleme nta l t raits Epilepsy a n

. d p o ssibly a few o th e r
f orms O f me nt a l defect ma y be excepti ons to this r ule .

STAT I S T I CA L M ET H ODS U S ED M EA S U RE M E N T OF
I N TH E

R ES E M B L A N CE — S i nce th e study o f me nt a l i nhe rit a nce

.

i nv olves a c onside ra ti on of degrees of resembl a nce b e

twee np a re nts a nd offsp ri ng ra th e r th a nth e me re no ti ng
,

o f p rese n ce or a bse nce o f p a rticul a r t ra its ce rt a i nst a tis ,

tic a l devices h a ve bee n f ound necess a ry Among th ese .

is th e c oeffi cie nt o f c orrel a ti on which is a nume ric a l i n

,

dex exp ressi ng th e a m ou nt of a g reeme nt betwee n two »

sets o f f a cts .

For ex a mple , le t u s su p p ose t h a t i t is desi red to exp ress

th e rel a ti onship existing betwee n i ntellige nce o f f a the rs
an d intellige nce o f s ons a n d th a t th e i ntellige n
,
ce of a
th o us a nd f a th e r sonp a i rs h a s bee n me a su red by m e nt a l
-

tes ts I f th e b righ test f a the r we re f o u nd to h a ve the

.

b righ test son th e nex t b righ test f a the r th e next b righ t

,
- -

est son a nd so on d own to th e d ullest the c orrel a ti on

, ,

c oefficie nt w o uld be 1 wh ich w ould rep rese n

,
t c o mple te
a g reeme n t betwee nth e two sets o f fa cts I f th e b righ test .

f a the r we re fou nd to h a ve th e d ulles t son the next ,

b righ test f a th e r th e nex t dullest son etc th e c orrel a

-
,
.
,
 R E S E M B L AN CE I N I NT E LLIG E N CE 51 7

ti on w ould be 1 wh ich w ould rep res ent c omplete

,

dis a g reeme nt A c orrel a ti on o f 0 w ould me a n no m ore

.

resembl a n ce th a n we sh ould get if we p a i red Off a ny

f a th e r with a ny one else s son a t ra nd om C orrel a ti ons
’
.

betwee n 0 a nd 1 w ould the ref ore rep rese nt a ll deg rees

of p ositive resembl a n ce f rom ve ry sligh t to p ra ctic a lly
p e rfect We a re now re a dy to ex a mi ne the c orrel a ti on
.

c oefficients of resembl a nce wh ich h a ve a ctu a lly bee n

,

f ou nd for rel a ted i ndividu a ls i n the c a se o f i ntelligence .

FA M I LY R E S E M B LA N CE N I N T
I E LL I G N CE
E — Th e co
efficie nt of resembl a nc e betwee n p a re nts a nd child re n i n
i ntellige nce seems to be i nth e neighb orh ood O f 50 wh ich
i s a b ou t h a lf wa y betwee npe rfec t r esembl a nce a nd no
.
,

r esembl a n ce a t a ll I t is a st riki ng f a ct th a t th e p a re nt
.

child resembl a nce i nm a ny physic a l t ra its a ls o tu rns o u t

to be ra the r u nif ormly i nth e neighb orh oo d of 50 This . .

w ould suggest th a t me nt a l a nd physic a l t ra its a re i n

h eri ted to much th e s a me deg ree .

The c orrel a ti ons betwee n si bs child re n of th e

s a me p a re nts ) h a ve a ls o bee nfou nd to a pp ro xim a te x50 ,

b o th for physic a l a nd me nt a l t ra its This h olds for

spelli ng a nd a rithmetic a l a bility a s m
.

e a su red by st a nd a rd
tests a nd f or a bili ty Ino the r sch ool subj ects a s me a su red
,

by sch ool m a rks R a ti ngs of 2000 sch ool child re n by

.

th ei r te a ch e rs on such t ra its a s i ntellige nce viv a city , ,

c onscie nti o usness p opul a rity tempe r self c onsci ousnes s

, , ,
-

an d a ssertiveness g a ve c orrel a ti ons betwee nsibs a pp rox i

m a ti ng 50 f or a ll of these t ra its
. .

In G e rm a ny sch ool m a rks e a rned b y sev era l hu n d red ‘

child ren were c omp a red wi th the sch ool m a rks e a rned
by thei r p a rents a nd g ra ndp a re nts Correl a ti on coeffi .

ci en ts we re no t c omputed but th e results sh ow ni nT a ble

,

1 a re sig nific a nt .Th e m a rks we re give n on a sc a le o f

five 1 bei ng h igh est a nd 5 l owest
, .
51 8 I NH E R ITAN CE OF M E NTAL T R AI T S

T A B LE 1 .
(Fro m Pete rs)

A v era ge M a r k s 0 f
Avera g e M a r ks
o f C hild re n
One P a re nt Ot h er Pa re nt

T H E LAw F I LI A L R EGRES S I O N
OF 1 it
.
— F rom T a ble
will be see n th a t supe ri or p a re nts te nd to h a ve ch ild ren
wh o a re a ls o supe ri or but less supe ri or th a n themselves ;
,

an d th a t i nfe ri or p a re nts te nd to h a ve ch ild re n wh o a re

i nfe ri or bu t less so th a nthemselves Th is illust ra tes th e
”
.
,

l a w o f fili a l reg ressi on wh ich wa s fi rst f ormul a ted

“
,

by F ra ncis G a lt on I t is me rely a st a teme nt of the f a ct

.

th a t the Off sp ri ng of p a re nts wh o a re excepti ona l i n a

p a rticul a r t ra it te nd to be less excep ti ona l i n th a t t ra it
th a n their p a re nts The l a w h olds for c omplex t ra i ts
.

like heigh t or i ntellige nce but not for simple t ra its li ke

,

eye pigme nt c ol or bli nd ness etc

-
,
-
, .

M E N TA L R ES E M BLA N CE O F T W I N S B o th inphysic a l .

an d me nta l t ra its a much high e r c o efficie nt of resem

‘

—
bl a nce h as bee n f ou nd for twi np a i rs th a n for ordi na ry .

sibs na mely 7 0 to 90 f or twi ns a s c omp a red wi th 50

, . . .

f or sibs Twi ns h oweve r a re believed to be o f two ki n

.
, ,
ds ,

th ose origi na ti ng f rom two eggs sep a ra tel y fe rtilized a nd

 E NVI R O N M E N T N OT S OL E FA C T O R 51 9

th ose p roduced by th e divisi on of a S i ngle fe rtilized egg .

It is p rob a ble th a t th e l a tte r m ode o f origi n a cc ou nts

for a ll c a ses o f so c a lled ide ntic a l
-
twi ns th a t is twi ns , ,

wh o a re a lm ost ex a ct duplic a tes O f e a ch o the r Ide ntic a l

‘

twi ns a re a lw a ys of th e s a me sex while f ra te rna l twi ns

,

ma y be ei the r o f the s a me or opp osite sex I t is believed .

th a t it is the p resence o f ide ntic a l twi ns i n th e g roups

wh ich h a ve been me a su red a nd tested th a t is l a rgely re
sp onsible for th e g re a te r resembl a nce of twi np a i rs a s -

c omp a red with ordi na ry sib p a i rs H oweve r th e ques

-
.
,

ti onis still i ndispute l a rgely owi ng to th e imp ossibility

,

o f ou r k n owi ng to a ce rt a inty wh i ch twi ns a re O f si ngle

egg or of two egg origi n
-
.

F A M I LY R ES E M B L A N CE N OT W H O LLY A CCO U N TED FOR

BY E N VIRON M EN T — One is na tu ra lly tempted to ex
pl a i n th e ment a l r esembl a nce O f p a re nt a nd ch ild a nd ,

especi a lly th a t of sib a nd sib or twi n a nd twi n a s due

, ,

to simil a rity o f e n vi ronme nt a nd t ra i ni ng The re a re .

seve ra l f a cts h oweve r wh ich re nde r this the ory u nte n

, ,

a ble.
(1 ) Th e resembl a nce i n me nt a l t ra its is a b ou t a s
g re a t a s i nphysic a l t ra its a nd it is k now nth a t the l a tte r
,

a re n o t dete rmi n ed chiefly by e nvi ronme nt ( )

2 The .

resembl a n ce of twi ns i n me nt a l tests wh ich h a ve novel

c onte nt is a s g re a t a s i nte sts wh ose c onte nt is m ore sub
ject to the i nfluence O f t ra ini ng (3 ) Olde r twi ns h a ve.

no h ighe r c o efficie nt of me nt a l resembl a nce th a ny ou nge r

twi ns If th e resembl a nce we re due ch i efly to e nviron
.

me nt the n i t sh o uld bec ome g re a te r th e l onge r th a t en

,

v i ron ment ope ra tes . I t d oes not ( )

4 G a lt on f o u
. nd
twi ns wh o h a d been subj ected to ve ry diff e re nt e nvi ron
ment wh o we re neve rtheless ex t remely simil a r (5 ) I t .

is a m a tte r of c omm on obse rv a ti on th a t sibs wh o diff er

g re a tly from e a ch o the r i ne a rly childh oo d ma y c onti nue
to di ffe r no twithst a n
,
di ng the simil a r e nvi ronme nt they
e nj oy th ro ugh ou t th ei r p e ri od of pl a sticity (6 ) G a lt on .

f o u nd th a t th e a d opted sons of p opes we re fa r less l i kely

to bec ome e mi n e nt th a n the re a l s ons of g re a t menwh o
520 I NH E R I TAN C E OF M E N T AL T R AIT S

ra nked wi th p opes i ni ntellige nce (7 ) W ood s study of .

’

h e redi ty i nth e roy a l f a m ilies o f E u rope sh ows th a t s ons

wh o i n he ri t th rones a nd f ortu nes a re bu t little m ore
lik ely to be numbe red by p oste rity a m ong the g re a t th a n
a re thei r y o u n ge r b ro the rs .

T H E I N H ER I TA N CE O F G E N I U S — The ge n ius is not

.

sh a rply d i ff e re nti a ted f rom o th e r h um a n bei ngs bu t is ,

me rely a pe rs ono f ve ry supe ri or a bility i none or m ore

lines All deg rees o f me nt a l a bility te nd to be t ra ns
.

mi tted a nd ge nius is no excep ti on to th e rule G a lt on

, .

fo u nd th a t 97 7 meno f ge nius wh om h e studied ge nius

bei ng defi ned a s th a t deg ree o f a bility p ossessed by the
m ost i ntellectu a lly emi ne nt pe rs on in a p opul a ti on o f
h a d a t o t a l o f 53 5 f a th e rs b roth e rs s on,
s g ra nd , ,

f a the rs g ra nds ons u ncles a nd neph ews a s emi ne nt a s

, ,

th emselves I t h a s bee n c a lcul a ted th a t th is is a b ou t

.

1 3 5 times a s m a n y emi ne nt rel a tives a s w ould be f o und

for 97 7 a ve ra ge men O f 1 030 B ritish men a n
. d w ome n
o f ge n ius studied by E llis 4 0 p er ce nt h a d eithe r a f a the r
,

or m o the r O f ve ry supe ri or a bility An exte nsive study .

of seve ra l h u n d red gifted ch ild re n o f Ca lif orni a e a ch O f ,

wh om equ a lled or exceeded i n i ntellige nce the b rightest

o u t o f 200 chi ld re n t a ke n a t ra n d om S h owed th a t th ese ,

a ls o h a d seve ra l times a s m a n y rel a tives of supe ri or

a bility a s do child re n O f a ve ra ge in tellige nce .

C onvi nci ng evide nce h a s a ls o bee n f o u n d for th e h e ri

ta bi l i ty o f a r tistic a n d music a l a bili ty In3 0 f a milies i n .

wh ich b oth p a re nts we re a rtistic G a lt on fou nd th a t 64,

p er ce nt o f t h e child re n S h owed speci a l t a le nt i n a r t Th is .

wa s th ree t imes th e p rop o rti onf ou n d i nf a milies i nwhich

neith e r p a re nt wa s a r tistic M usic a l a bili ty especi a lly is
” ,
.
,

p rone to ru n i n f a milies Th e B a ch f a mily for ex

“
.
,

a mple ,
p roduced i n eigh t ge ne ra ti ons no less th a n 57
i ndividu a ls o f ve ry supe ri or music a l a bility 20 of wh om ,

bec a me emi ne nt .

I N H ERI TA N CE OF F EEBLE M I N DEDN Es s -

I nvestig a .

ti ons by G odd a rd D a ve np ort a nd o the rs h a ve sh ow nth a t

,
 T HE KALLI KAK FA M IL Y 5 21

a m a j ori ty o f the c as es of me nt a l deficie nc y a re t ra ce a ble

to he redity p o ssibly a s l a rge a p rop orti ona s two thi rds
,
-

or th ree f o u rth s
-
Th e rem a i nde r a re due to a v a riety o f
.

c a uses i nclud i ng i nj u ry a t bi rth dise a ses or a ccide nts

, ,

du ring inf a ncy a nd syphilis or a lc oh olism of p a re nts

,
.

S oci a l s tudies h a ve bee nm a de of m a ny feeble mi nded -

or o the rwise dege n e ra te st ra i ns g ood ex a mples of which ,

a re the Ka ll ik a ks th e N a m F a mily ,
th e H ill F olk th e , ,

Pin ey s th e J ukes a n
,
d th e Ishm a elites R ep rese nt a tives .

of such f a milies a re to be f o u n d i na lm ost a ny neighb or

h ood a nd eve rywhe re th ey m a ke up a l a rge p rop orti on
,

of th e ne e r do wells p a up e rs c rimi na ls a nd sex

’
- -
, , ,

offe n de rs .

S ome i nvestig a t ors not a bly Godd a rd a nd D a ve n

,
p ort ,

a re i n cli ned to believe th a t me nt a l deficiency is due to a

single f a ct or or ge ne which beh a ves a s a M e ndeli a n re
cessi v e .This w o uld me a n th a t defec tive m a ted with
defective w ould give only defectives i n the F D efe o 1 .

tive m a ted with norm a l w o uld give a n F c omp osed 1

e nti rely of norm a l but hyb rid i ndividu a ls I f th e l a tte r

sh ould m a te w
.

ith o the rs wh o we re hyb rid like th emselves ,

the resul t w ould be a nF yieldi ng th e f a mili a r 2 r a ti o .

I t is by no me a ns ce rt a i n h oweve r th a t this i nterp ret a , ,

ti on of the d a t a i s c orrect Th e g ra ded ch a ra cte r of .

me nt a l defect w ould suggest th a t p rob a bly m ore th a n

on e f a ct or is i nv olved .

T H E K A LLI K A K F A M I LY Th is f a mily is of speci a l

.

i nte rest by re a s on of th e f a ct th a t it c ont a i ned two

b ra nch es a norm a l a nd a d efective B oth b ra nches
,
.

t ra ce b a ck to a me nt a lly norm a l y ou ng s oldie r o f th e

R ev oluti ona ry W a r D u ri ng th e wa r this s oldie r m a ted
.

temp ora rily with a feeble mi nded w om a n givi ng rise to -

,

a feeble mi n -
ded st ra i n wh ich exists to th is da y F rom .

th is u ni on4 80 i ndividu a ls h a ve bee nt ra ced 1 4 3 of wh om ,

a re k n ow n to h a ve bee n feeble mi n ded a nd m a ny othe rs -

c rimi na l imm ora l or a lc oh olic Only 4 6 a re know n to

, ,
.

h a ve bee nnorm a l After th e wa r th is s a me s oldie r ma r

.
5 22 IN H E R I TAN CE OF M E NTAL T R AIT S

r ied a w om a n o f g ood h e redity a nd f o u nded a noth e r

f a mi l y O f wh o m 4 96 i ndividu a l s h a ve bee nt ra c ed Pra c
n
.

tic a lly eve ry membe r o f th is b ra nch wa s norm a l e a rly ,

a ll o f th em bei n g p rospe ro us f a rme rs or successful l a wye rs ,

j udges or h olde rs o f imp ort a nt public O ffi ce N o t eve n

,
.

a we l l c on
-
t rolled expe rime nt c o uld h a ve sh ow nm ore con
vi ncingly th e imp ort a nt p a rt pl a yed by h e redity i n the
c a us a ti on o f me nt a l deficie ncy .

I N H ER ITA N CE O F OT H ER FOR M S OF M E N TA L D E FE CT .

B o th ep i lepsy a nd i ns a nity a re a ls o k now nto be h e redi

t a ry th o ugh th e l a ws g ove rni ng th ei r t ra nsmissi on a re
,

no t fully u nde rs tood I t seems to be f a i rly well est a b

.
-

l i sh ed th a t epilepsy is d ue to a si ngle recessive M e n deli a n

f a ct o r Th e re a re m a ny f orms o f i ns a nity a nd i t is p os
.

sible th a t not a l l o f th ese a re t ra nsmitted a cc ord i ng to th e

sa me l a ws I t h a s bee n f o u nd th a t epilepsy i ns a nity
.
, ,

an d feeble mi nded ness f reque ntly a ppe a r i n th e sa me

-

f a milies O fte none ki nd o f defect a ppe a ri ng wh e na no th e r

,

w o uld be expected a nd it is p ossible th a t th ese v a ri ou s

,

t ypes of defec t ma y be rel a ted to e a ch o th e r i ns ome a s

ye t u nk now nm a nne r Th e p redisp o siti onto a l c oh olism
.

is a ls o to s o me exte nt h e redit a ry a nd O f te n a p p e a rs i n
f a m ilies m a rked by o th e r types o f defect Crim inal ity .
,

a s such is p rob a bly no t h eredi ta ry b u t a c onside ra ble

‘

, ,

p ro p orti on of c rimi na ls a re feeble mi nded a nd feeble -

,

mi nded ness a s we h a ve see n is h e redit a ry

, ,

Figu res 1 50 1 —2 a re typ ic a l f a mily hist ories O f epil epsy

-

an d in s a nity .

E N DO W M E N T OF T H E D I FF ERE N T S OC I A L AN D O CCUPA
T I O N AL C LA S S E S .
—Wh a t
bee ns a id i n th e p reced i ng
ha s
p a ra gra ph s w o uld suggest th a t i n ge ne ra l membe rs of , ,

th e so ci a lly successful cl a sses a nd h igh e r o ccup a ti o na l

gro ups a re onth e a ve ra ge bette r e nd owed me nt a lly th a n
th ose o f i nfe ri or s oci a l st a tus Th a t such is th e c a se is .

i ndic a ted by th e i ntellige nce o f th ei r O ff sp ri ng M e nta l .

tests h a ve b ro ugh t ou t th e f a ct th a t th e ch il dre nO f suc

cessf ul l a wye rs d oc to rs mi niste rs or c ollege p ro fessors
, , ,
 E ND O W M E N T OF D I FFE R E N T CLA S S E S 5 23

h a ve a n a ve ra ge i ntellige nce qu o tie nt of 1 1 5 or h igh e r ,

an d th a t ch ild re n Of t o t a lly u nskilled l a b ore rs h a ve a n

. om
© ThcN1 StateH e for Bo ys
© Childrcn
ls Industrial Hom r

NewBrun swick

FI G 1 50
. .
— The
P ro du ct o f a Feebl e- M i n de d M a n (wh o h a s a n
epil e p ti c b ro t h er) a nd his epil e p t i c wi e (wh o se f a t h er wa s insa n
f e

an d u ncl e eebl e mi n
f d ed ) ; th e fi rst child di e d i n i nf a ncy , th e next
-

two w ere f eeble - mi n de d a nd die d y o u ng, th e ne xt is a n ep il ep t ic

‘

a t t h e N ew J ers e y S t a t e V ill a ge ; the n e x t is f e ebl e mi n de d , h a s a

-

crimi n a l re cor d a nd is i nth e S t a te Ho m e f or B oys ; th e l a s t is fe ebl e

mi nde d a nd is i nth e C hil drens Industri a l Hom e Si x i nt his a mily
’
. f
h a ve b eenor a re w a rds of th e s ta te A , a l co h oli c ; C, crimina listi c ;
.

D , dea ; E , e pil e p ti c ; F, eebl e mi nde d ; I i n

f f -
s a ne, N , norma l , S V
in 3155? m e a ns a n i nm a te o f a S t a t e V i ll a ge f or E pil e p ti cs .

D a v enp ort, Heredi ty i nR ela ti onto E ugen

’

i cs .

aw s n
oou

FI G . 1 51 -
.An Al co h oli c
M a n of Goo d Fa mily b u t p rob a bly
si mpl ex i n m en norm a l w om a n 2 norm a l child ren
t a li t y h a s by
a

an d by a f e ebl e mi nd e d w o m a n 2 norm a ls a nd 1 fee bl e mi nd e d

- -
.

He h a s h a d 4 oth er children by f eebl e mi nd e d w om en a ll f eeble

-
,

minde d .

D a v enp ort, Heredi ty i nR ela ti onto Eu geni cs .

5 24 I NH E R ITAN CE OF M E NTAL T R AIT S
a ve ra ge qu otie nt o f 85 o r l ow er I t sh ould be emph a
.

sized h o weve r th a t th e figu res refe r to a ve ra g es In

, , .

b o th c a s es th e ra nge O f qu o tie nts is wide a nd e a ch di s

tri b u ti o ncu rve g re a tly o ve rl a ps th e o th e r .

Al th ough feeble minded child re n a re occ a si ona l ly

-

f o und i n s oci a lly successful f a mi li es they a re seve ra l ,

tlIIl eS a s f reque nt inth e f a milies o f l ow occup a ti on al a nd

s oci a l st a tus ; a nd a l th o ugh gifted ch ild re na re s ometim es
f o u nd i nth e f a milies o f u nskilled l a b ore rs they a re m a ny ,

times a s f reque nt i n f a milies bel ongi ng to th e p rofes

si on a l cl a sses .

O f seve ra l h u nd red C a lif orni a child re nwh o we re f ou n d

to h a ve a ni n tell ige nce qu o tie nt of 1 4 0 or h igh e r a nd a l l ,

Fi g 1 52.
—Inherita nce insa nity
Of From t h e ce ntra l ma t ing O f
.

two n orm a l pe rso ns t h e re a re d eriv ed 8 child re n 3 insa ne

,
B u t th ere .

is the h eredi ta ry te ndency in the germ pl a s m of both p a rents Mott

,
.
,

1 90 5 .

Of wh o m p rob a bly ra nked with i n th e top h a l f of th e top

1 p er ce nt O f th e ge n e ra l sch ool p opul a ti on a p p roxi ,

m a tely 50 p er ce nt h a d f a th e rs wh o bel onged to one or

an o th e r O f th e p ro fessi on s M ore th a n35 p er ce nt we re
.

f ro m th e semi p ro fessi ona l or me rc a ntile cl a sses s ome

-
,

thi ng ove r 1 0 p er ce nt f rom th e S killed l a b or cl a sses a nd ,

less th a n 5 p er ce nt f ro m th e u nskilled a nd Semi ski lled -

o ccup a ti on a l cl a sses c o mbi n ed G a lt on Ellis a nd oth e rs

.
, ,

h a ve b ro ugh t o u t simil a r f a cts reg a rdi ng th e s oci a l origi n

o f me no f ge n ius .

S u ch f a c ts a re O f g re a t signific a n ce Th ey me a n th a t
.

we must usu a lly l ook to th e high e r s oci a l cl a sses to p ro

 D E CR E A S E OF S UP E R I OR S T O CK S 525

duce th e i ntellectu a l le a de rs wh o will a dv a nce scie nce ,

a rt g ove rn
,
me nt educ a ti on a nd s oci a l welf a re ge ne ra lly
, ,

Ou r civiliz a ti on o f a th o us a nd or ten th o us a n d ye a rs
h ence w ill depe nd l a rgely up on th e rel a tive fecu ndity o f
ou r l ow gr a de a n d h igh g ra de st o cks .

T H E D E CR EA S I N G F ER T I LI TY O F S UP ER I OR ST O CK S .

U ntil rece ntly th e re h a d not bee n a t le a st for h u nd reds

,

of ye a rs a n , y m a r ked te n de n cy i nm ost civilized c ou nt ries

f or one cl a ss to rep ro duce m ore ra pidly th a n a no th e r .

For ce ntu ries th e a vera ge me nt a l e nd owme nt of th e

E u rope a n a nd Ame ric a n p e oples h a d h eld i ts own o r ,

p ossibly evena dv a nced sligh tly a s a result of th e g re a te r

m ort a lity a m ong th e u ndesi ra bles Ab out h a lf a ce ntu ry
.

a go h oweve r a n ill b o di n
, ,
-
g ch a nge beg a n to t a ke pl a ce .

Th e i ntellec tu a lly supe ri or f a milies a re no l onge r rep ro

du ci ng a s ra pidly a s f orme rly a nd th ei r ra te of rep rodu c
,

ti onh a s f a lle nfa r bel ow th a t of th e s oci a lly i nc omp etent .

Wh i le th e fecu ndity O f feeble mi nded f a milies c onti nues

-

u ndimi nish ed th a t of u nive rsity g ra du a tes a nd o th ers O f

,

c orresp ondi ng intellectu a l supe ri ority h a s i ntwo or th ree

ge ne ra ti ons d ec re a sed by s ometh i ng li ke one h a lf The -
.

te ndency t ow a rd a di ff e renti a l bi rth ra te is m a ki ng a l a rm

i ng p rogress i n a l l civilized c ou nt ries a lth o ugh less i n ,

It a ly Ge rm a ny a nd J a p a n th a n i n Fra nce Engl a nd or

, , , ,

Ame ric a Th e situ a ti on is only i n p a rt relieved by the

.

s omewh a t l ow er m ort a lity a m ong th e bette r cl a sses -

.

I f th e di ffe re nti a l bi r th ra te sh o uld c onti nue a nd es ,

p ec i a ll y if its di ffe r e nti a l ch a ra cte r sh o uld bec o me m o re

an d m ore m a rked it w o uld be bu t a few h u nd reds or

,

th ous a nds of ye a rs u ntil th e su rvivi ng stOcks w ould be

th o se descended ch iefly from th e d regs of ou r p rese nt
da y p opul a ti on If this sh ould o ccu r no a m o u nt o f
.
,

educ a ti ona l e ff ort w ould p reve nt th e dec a y O f th e le a di ng

m ode rn na ti ons Th e re is re a s on to believe th a t such
fluen
.

a n in ce wa s l a rgely a cc ou nt a ble for th e w a ni ng of

e a rlie r civiliz a ti ons not a bly i n th e c a se of G reece a nd
,

R ome .
526 IN H E R I TAN C E OF M E NTAL T R A IT S
TH E I M P ORTA N CE OF E U GE N I CS — The re is no evi
.

de nce th a t th e d ec re a sed bi rth ra te i n the supe ri or

cl a s ses is to a n y g re a t ex te n t d ue to bi ol ogic a l c a uses I t .

is p rob a bly due e nti rely to the v olu nt a ry a c ts of individ

u a l s i nthe p o stp o n eme nt of m a rri a ge a nd th e p reve nti on
o f c on cepti on Th e re is the p ossib i lity the refore th a t
.
, ,

the S itu a ti on ma y be s a ved by ed uc a ti on i n euge nics .

Th e diffi culty is th a t a l th ough i t is rel a tively e a sy to

te a ch pe ople th e sig nific a nce o f th e issues i nv olved it ,

is ve ry h a rd to pe rsu a de them to c a re e nough a b o u t the

futu re of civil iz a ti onto m a ke a ny s a c rifices i nits beh a l f .

I t is evide nt a t a ny ra te th a t ma n f a ces few

, , ,

if a ny p roblems of g re a te r imp ort a nce for h is futu re

,

welf a r e th a n th o se rel a ti ng to the c ont rol of h e redity .

By th e st ric t a pplic a ti on O f euge nic p ri nciples it w o uld

be p ossible to red uce g re a tly th e numbe r o f defectives ,

to in c re a se the p roducti on o f ge nius a nd to imp rove ,

the a ve ra ge level of m oral ity a nd i ntellige nce .

T H E E FF E CT O F U N DES IR A B LE l M M I GR AT I O N — The .

me nt a l tests give n to ne a rly two milli on o f ou r s oldie rs

d u ri ng th e rece nt w a r discl osed l a rge di ff e re nces i n th e
i ntellectu a l a bili ty of v a ri ous ra c i a l g roups E vidence .

to th e s a me eff ect is a ls o a v a il a ble f ro m n ume rous ex

p eri menta l studies Th e f oll owi ng c onclusi ons seem
.

w a rra nted .

I P ro b a bly n
. o t m o re th a n1 0 or 1 5 p er ce n t of Ame ri
ca n N e gro es eq u a l or exceed i n i n tellige nce th e a ve ra ge
White The i nte llige nce o f th e a ve ra ge N eg ro is v as tly
.

i nfe ri o r to th a t O f th e a ve ra ge white a nd th e mul a tt o

,

oc cupies a p o siti o n a b o u t mid wa y betwee n -

.

2 The intellige n
. ce o f th e a ve ra g e Ame ric a n I ndi a nis
a pp reci a bly but n o t gr e a tly supe ri o r to th a t o f th e a ve r

a ge N eg ro O u r M exic a n p opul a ti on (I n
. di a n hyb rid s
for th e m o st p a rt ) wh ich h a s rece ntly inc re a sed with
g re a t ra pidity b a rely su rp a sses th e N eg ro i n a ve ra ge
,

e nd owme nt .

3 The immig ra n
. ts wh o h a ve bee n c omi ng to us f rom
 E UG E NI C S A ND M A R R IAG E 52 7

the ext reme s outhe rn a nd s outhe a ste rn p a rts o f Eu rope

(S o uth It a li a ns P
,
o rtuguese G reeks a nd S l a vs ) a re i n
, ,

ge ne ra l disti nctly i nfe ri or to th ose wh o c ome to u s f rom

N orthe rn Ce nt ra l a nd Weste rn E u rope Th e i nfl ux o f '

, ,
.

th e forme r h a s bee nso g re a t a nd their ra te O f rep roduc

,

ti on is so excessive a s to give u se to a se ri ous me na ce

, .

4 Th e few ment a l test studies wh ich h a ve b ee nm a de

.

o f Ch in ese a nd J a p a nese i ndic a te th a t these ra ces a re

a pp rox im a t ely the equ a ls of E u ro pe a n s i nment a l a bility .

5 N O na ti on ca n a ff ord to ove rl oo k the d a nge r th a t

.

th e a ve ra ge qu a lity of its ge rm pl a sm ma y g ra du a lly

dete ri ora te a s a resul t of u nrestricted immig ra ti on .

E U GE N I CS AN D M ARR I A GE — The l a ws o f me nt a l a nd
physic a l i nhe rit a nce a re not yet well e nough u nde rst ood
to w a rra nt l a yi n g d ow n a det a iled p rog ra m for secu ri ng
eugenic m a t ings S ome o f th e f a cts which h a ve bee n
.

set f orth h a ve a nObvi ous be a ri ng on the p roblem One .

wh o h a s reg a rd f or th e qu a lity o f h is desce nd a nts sh ould

o f c ou rse a v o id m a ti n g i nt o a ra ci a l or f a mily st o ck
wh ich is i nfe ri or to his own eve nif it is not ch a ra cte rized
,

by out righ t defect The resul ts ma y be s a tisf a ct ory but

.
,

th e ch a nces a re onthe othe r side The u nwisd om of ma t

.

in g with o n e w h o se f a mily is se ri o usly t a i n ted with

feeble mi nded ness i ns a nity epilepsy p a upe rism or a l
-
, , , ,

coh ol i sm ,
is too Obvi ous to dem a nd emph a sis The .

d a nge r of such m a ti ngs is g re a tly i nc re a sed wh e n b o th

p a rties to th e m a rri a ge c ome of defective st ock .

Th e desi ra bility of a d opti ng leg a l me a su res to p reve nt

the feeble mi nded f rom rep roduci ng is self evide nt M a ny
- -
.

st a tes h a ve l a ws desig ned to a cc omplish this but unfor ,

tu n a tel y th ey a re ra rely e n f orced a g a i nst the higher

g ra des of defectives S te riliz a ti onof the u nfit h a s g a i ne d
.

lit tle he a dw a y H oweve r reducti on of th e numbe r of

.
,

defectives is only a sm a ll p a rt of the p roblem of euge nics .

E limi na ti onof a ll the feeble mi nded w o uld n -

o t? ra ise th e

a ve ra ge level o f i n tellige nce i n th e ge ne ra l p op ul a ti on

to m ore th a n a b a rely n o tice a ble ex te n t a nd i t is the
,
5 28 IN H E R I TAN CE _OF M E NTAL T R AI T S

imp roveme nt o f o u r e nti re p opul a ti on th a t c ou nts i nthe

l ong ru n S till m ore imp ort a nt is the a d op ti on of eu
.

ge nic me a su res which will inc re a se th e numbe r o f i ntel

.

lectu a l a nd m ora l ge niuses .

T H E I N F LU E N CE O F E N VI R O N M E N T AN D EDU CAT I O N
UP O N M E N TA L P ER F OR M A N CE F rom th e p reced ing dis .

cu ssi o nit migh t be i n fe rred th a t one s me nt a l t ra its a re ’

dete rmined a lm o st e nti rely by h e redi t a ry f a ct ors a nd ,

th a t th e i nfl ue nces of e nvi ronme nt a nd f orm a l t ra i ni ng

6 o o c!
!
a shows e ma nof '

seoe mi fie a bi l i ty shows a
'

ma nof m emi fic a bi li ty who i s a lso 0

.

Fel low of th e Royal Soeael y ; . en

ws five o ther ( hi ldxen n
6) sho d so o .

FI G . 1 53 . Inh eri ta nce

Abili ty (ch a rt co nd e nse d a nd i ncom l
of -

p le te ) i nt h re e m a rk e dly a bl e f a milie s (f ro m Ke l l i co tt a f te r Wh e th a m )

1 C h a rl e s D a rwi n; 2 his co usi n Fra n

, , cis Ga l to n f o u n d er of the
, ,

m o de rn e u ge ni c m o ve me nt .

Guy er B ei n g Well born

-
.
,

c ou nt fo r nex t to nothi ng S uch a c onclusi on is by no .

me a ns j ustified Alth ough it is t rue th a t we c a nnot

.

cu re c onge ni ta l feeble m inded ness d evel op a dull a rd i nto

-
,

a ge n ius or t ra nsform a pe rs on wh o is t o ne de a f i nt o a
,
-

M e ndelss oh n i t is p ossible by me a ns of educ a ti on to

,

ra ise e n orm o usly the ge n e ra l level o f a cc omplishme nt o f

a ne n ti re ra ce or p opul a ti ong roup O ne h a s only to com
”
.

p a re th e N eg ro es i n Ame ric a with th ei r c o usi ns i n “

A f ri ca to see wh a t a n imme nse di ff e re nce ed uc a ti o n ca n

m a ke O u r a ncest ors O f two th o us a nd ye a rs a go we re
.
 H E R E DIT Y SETS T HE LI M I T 5 29

p rob a bly a s well e nd owed me nt a lly a s Ou rselves but for ,

l a ck of ed uc a ti on th ey we re p ra ctic a lly b a rb a ri a ns O ur .

sch ools h a nd ove r to child re n i n ten or a d oze n ye a rs

much of th e k nowledge a nd skill wh ich it h a s t a ke n the
ra ce th o us a n ds of ye a rs to a cqui re .

H EREDI TY S ET S T H E L I M I TS I t is not a questi on of

.

he redity or envi ronme nt bu t o f he redity a nd e nvi ron

,

ment B o th i nflue nces a re op e ra tive i ndete rmi ni ng wh a t

.

m a nne r of pe rs on a ny give n i ndividu a l sh a ll bec ome .

Wh a t he redity d oes is to set limits to th e i nflue nce which

it is p ossible for educ a ti on to exe rt I t is to a g re a t ex
.

tent educ a ti onwhich decides wh a t th e a bsolu te a ch ieve

ment of a give n p opul a ti on g roup sh a ll b e — th a t is -
,

whethe r its i ntellec tu al life sh a ll i nge ne ra l be ona high

or l ow pl a n e On the o the r h a nd it is h e redity ch iefly
.
, , ,

which dete rmi nes th e re la ti ve a ch ieveme nt of a n i ndi

vidu a l a s c omp a red with o the r membe rs o f his p op ul a
ti on g roup . At le a st this tends to be th e c a se i n a
dem oc ra tic c ou nt ry whe re cl a ss p rivil eges h a ve bee n
a b olished a n d whe re i nequ a lities of o pp ortu nity h a ve
b eenreduced to a mi nimum T o st a te it c onc retely the
.
,

a ve ra ge En glishm a n of t o d a y is m ore civilized th a nh is

an cest ors Of two th ous a nd ye a rs a go ch iefly bec a use o f
, “

educ a ti on a nd envi ronment bu t Englishme n like F a ra

,

da y L ord K elvi n or Ll oyd Ge orge su rp a ss th e a ve ra ge

p rese n t da y English m
, ,

-
a n i n a chieveme n t ch iefly bec a use
of t hei r supe ri or e n d owme nt .

Ordi n a ry Obse rv a ti onsuggests th e p ossibility th a t edu

c a ti on ma y h a ve a gre a te r i nfl ue nce up on m ora l t ra its

th a n up on ge ne ra l i ntellige nce This ma y b e t rue but
.
,

it is p rob a ble th a t eve nh e re origina l e nd owment c ou nts

he a vily N O na ti onh a s eve r succeeded i n ref ormi ng i ts
.

e nti re crimina l cl a ss .

M ODI F I CATI O N S OF M EN TA L T R AI TS N OT TR A N S M I TTED .

-
I n e a rlie r ch a pte rs (XX V I I I a nd XXX I I I ) i t h a s
beensh ow nth a t physic a l t ra its a cqui red du ri ng th e life
tim e Of a nindividu a l do not a ffect the ge rm cell th a t is , ,
530 IN H E R ITAN C E OF M E NT AL T R AIT S

do no t bec ome h e redit a ry Th is a ppe a rs to be equ a lly

.

t rue O f me nt a l t ra its The f a c t th a t ou r p a re nts h a ve

.

S p o ke nE n glish or studied L a ti nor ref ra i ned f rom d oing

evil d oes not m a ke it a ny e a sie r for u s to a cqui re th ese
h a bits Th e ch a nces th a t a p a re nt of a give ndeg ree o f
.

music a l a bility will t ra nsmi t th a t a bility to h is Off sp ring

a re n o t i n the le a st a ffected by the a m o un t of music a l
t ra i ni ng h e himself h a s received The re is only one wa y
.

in which we ca n be nefit f rom th e less on s o u r p a re nts

h a ve le a rned na mely by f oll owi ng th ei r ex a mple a nd
, ,

le a rning th e s a me less ons for ou rselves Educ a ti onmust

.

begi na ll ove r a g a i nwith eve ry ge ne ra ti on; which me a ns

th a t th e sch ool is a nd will a lw a ys be the m ost imp ort a nt
,

o f a ll our s oci a l i nstituti ons .

 C HAPTER XLIII

M AN S PLA CE I N N AT UR E
’

M AN TH E APEx OF T H E A N I M A L S ER I E S If th e p res
.

en ta ti on o f th e f a c ts a n d p ri nciples o f bi ol ogy give n i n

this b ook h a s a cc omplished its pu rp ose i t sh ould be cle a r
,

i n th e mi nd o f th e re a de r th a t no sh a rp li ne sep a ra tes
ma n from o th e r livi ng c re a tu res Th e w orld O f na tu re
.

p rese nts a v a st a rra y of f orms v a ryi ng i nc omple xity but

, ,

a l l h a vi ng a c omm onb a sis of p rot opl a sm a nd a ll sh ow

in g a high d eg ree O f a dj ustme n t to thei r envi ronme nt .

M a n is with o u t d o ub t th e m ost h ighly speci a lized of

them a ll a nd yet his met a bo lic p rocesses a re essenti a lly
,

like th ei rs M a n occupies th e n a defi nite p ositi on i n

.
, ,

na tu re Th e f a ct th a t h is p ositi onis a t th e pe a k of the

.

sc a le of deve lopme nt sh ould not ca u se ma n to l ose sight

o f th e fu n d a me nt a l f a c t th a t he is a p a rt of the orde r of
na tu re a nd is subj ect to th e s a me l a ws b y which the .

a ctivities of o th e r livi n g th i ngs a re c ont rolled .

M A N S S I M I L AR I TY TO T H E H I GH ER A N I M A L S — F rom
’
.

wh a t h a s bee n s a id i n p revio us secti ons o f this b ook it

sh ould be cle a r th a t i nphysic a l c onstitu ti onma n di ffe rs
on ly sligh tly f rom m a ny of th e highe r a nim a ls His .

pl a nof b odily st ructu re is th e s a me a s th a t of m a mm a ls

ge nera lly E xcep t for a few det a ils h is ph ysic a l st ructu re
.
,

is st riki ngly simil a r to th a t O f the h igh est m a mm a li a n

g roups th e m onkeys a nd a p es Th e fu nd a me nt a l b a sis
, .

of n e v ous functi oni ng i n ma n is

r — so f a r a s we ca n
—
le a rn p recisely simil a r to th a t of o the r m a mm a ls a nd ,

i nf a ct of a ll except th e m o st l owly a nim a ls

,
I nma na s .

i no the r a nim a ls simple reflexes c o mplex reflex a cti ons

, ,

the p rop erti es of mem ory a nd a ss oci a ti on f orm th e b a sis

,

53 1
532 M AN S ’
P LA C E IN NATU R E
Of the i ntellectu a l life I n ma n a s in othe r a n
. im a ls ,

a g ree a ble a n d dis a g ree a ble em oti o n s a re a roused u nd er

p a r ticul a r c on diti ons a nd a re a cc omp a nied by ch a ra c
teri sti c b odily ch a n ges Ina ll these fu nd a me nt a l m a tte rs
.

the re i s no esc aping th e c onclusi on th a t ma n is a t one

with the wh ole g ro up of m a mm a ls to wh ich h e bel ongs .

T H E F A CT O F P R O GR E S S N o twiths ta ndi ng the p o ints

.

O f simil a rity em p h a sized i n the l a st p a ra g ra ph on e wh o ,

studies hum a nbe ha vi or i nc omp a ris onwith th e beh a vi or

o f high e r a n im a ls c a nno t f a il to be st ruck with a fe a tu re
o f di ff e re nce which is a bs olu tely fu nd a me nt a l a nd which ,

i n f a ct sets ma n i n a c a teg ory by himself

,
I n a w ord.
,

this d iff e re nce lies — a s exp ress ed i n the h e a d o f this

p a ra g ra ph — i n th e f a c t o f h um a n p rog ress which con

tra sts with th e a bse nce of p rog ress a m on g th e oth er
an im a ls Wh a t is me a nt by p ro gress i n this c onnecti on
.

is simply this : h um a nbei ngs f rom ge ne ra ti onto ge ne ra

,

ti on beh a ve diffe re ntly f rom th e ge ne ra ti ons which p re

ceded the m they do new thi ngs a nd thi nk new th ough ts
,

.
,

A nim a ls on th e othe r h a nd , ge ne ra ti on by ge ne ra ti on
, ,

beh a ve i n a m a nne r wh ich d oes not esse nti a lly ch a nge ,

thei r re a cti ons to th e e nvi ronme nt a re on the wh ole the

s a me now a s they we re h u nd reds of ge ne ra ti ons a go .

This d iffe re nce i nbeh a vi or is b oth re a l a nd fu nd a me nt a l ,

an d to th e exte nt to wh ich i t ca n be expl a ined m a ns

’ !

p ositi oni nna tu re i nc omp a ris onwi th th e o th e r a nim a ls ’

ca nbe m a d e cle a r .

T H E H U M A N R A CE U N CH A N GED I f we we re to fi nd
.

p rese nt ge ne ra ti ons of a nim a ls beh a vi ng d i ff e re ntly f rom

p a s t ge ne ra ti ons we sh ould h a ve no h esi ta ti on i n a s
,

se rti ng such diffe re nce i n beh a vi or to be du e to b odily

a lte ra ti on s Th e ch a nges i n hu ma nbeh a vi o r f ro m gen
.

c ra ti on to ge ne ra ti on a re not thu s a cc ou nted fo r The .

evide nce is c o nvi nci ng th a t ma ni nhis p rese nt physic a l ,

i ntellectu a l a nd em oti ona l ch a ra cte rs h a s existed onth e

,

e a rth si nce th e begi nni ng of re corded his tory a nd i na ll ,

li kelih ood for a pe ri od a t le a st two or th ree t i mes a s l ong

 E X PLANATI O N OF P R OG RE S S 533

p rec edi ng th e fi rst m a ki ng of pe rm a ne nt rec ords We .

know th a t ve ry e a rly i n hist oric time men lived wh o

we re i n a ll p rob a bility fully equ a l to m ode rn ma n not
‘

on ly i n physic a l p owe r but i n i ntellectu a l a bility a nd

,

em o ti ona l ch a ra cte ristics a s well Th is le a ds to the con

.

cl u si on th a t th e p rog ress o f hum a n ity i n hist oric a nd

l a te p rehist oric t ime h a s not bee ndue to ra ci a l devel op
ment bu t ra th er to th e utiliz a ti onof qu a lities which we re
'

p ossessed by p rimi tive ma nequ a lly with m ode rnma n .

EX P LA N ATI O N O F P R OGR ES S —The expl a na ti on O f

.

p rogress is to be s ough t i n s ome qu a lities which ma n

’

p ossesses a nd which oth e r a nim a ls do not si nce it is cle a r ,

th a t if the o the r a nim a ls we re i np ossesssi onof the s a me

or of simila r qu a lities they w o uld Sh ow th e Sa me
a bility . Hum a n a nd a nim a l ch a ra cte ristics h a ve bee n
cl a ssified a s physica l i ntellectu a l a n
,
d em oti on
,
al In .

these th ree r e a lms a re to be s ough t th e S peci a l p rope rties

wh ich h a ve f ormed the b a sis of h um a n p rogress .

P H Y S I CA L C H ARA CTER I S TI CS N o twiths ta nding m a ns ’

.

m a rked physic a l simil a rity to the o the r m a mm a ls his ,

b od y p rese nt s two speci a l fe a tu res which u nd oubtedly

h a ve c ont ributed to his a chievements Th e fi rst of these .

i s the e rect p ostu re I t is ch a ra cte ristic O f ma n th a t h e

.

a l on e a m ong a nim a ls st a nds definitely up ontwo feet a nd

pe rf orms th e m a j or p a r t of his l oc om oti onwith th e l owe r
limbs only Th e a dv a nt a ge th a t this c onfers up on him
.

is chiefly i nthe rele a se O f th e front limbs f rom l oc om ot or

duty a nd i nthus freei ng them for use a s org a ns of t o uch
an d for c a rryi ng on the g ra spi ng fu ncti on It is quite .

t rue th a t m a ny o the r ki ri ds of a nim a ls u se th e f ront

li m ,
b s ometimes to a l a rge exte nt a s a n org a n O f to uch
,

an d of g ra sp ing bu t only i n ma n a re these th e p rim a ry

,

fu ncti ons a nd l oc om oti on th e sec ond a ry I n a ll othe r

, .

m a mm a ls l oc om oti on is th e p ri ma ry fu ncti on O f th e
f ront limb o the r fu ncti ons bei ng only i ncide nt a l
,
.

Th e sec ond sig nific a nt physic a l d iffe re nce betwee nma n

an d o the r a nim a ls i s i n th e supe ri or flexibili ty of th e
534 M AN S’
P LA C E I N NATU R E
h a nd In no othe r a nim a l ca n th e th umb be m oved
.

a c ro ss th e p a lm o f th e h a n d a s ca n be d one f reely a nd
ac cu ra tely b y m a n This flexibility lies a t the b a s is of
.

m a nu a l dexte rity a nd whe n i t is rec a lled h ow much of

“

h is p rog ress ma n owes to ski llful use o f his h a nds th e ,

imp ort a nce of th is physic a l ch a ra c te r i s cle a r N ot .

on ly i nth e u se of mech a nic a l devices bu t i n th e use of ,

i nst rume nts for w ri ti ng a nd d ra wi ng is th is of p rime

sig nific a nce The re is no inte nti onh e re o f a sse rti ng th a t
.

these physic a l peculi a rities a re of themselves a sufficie nt

b a sis for h um a np rog ress which rests p rim a rily onch a r ,

a cters to b e desc ribed p rese n tly but a s c ont ribu tory ,

eleme nts they a re not to b e neglected .

I N TELLE CTUA L C H AR A CTER I S TI CS — Prob a bly ne a rly

eve ry one i f a sked to wh a t ma n owes h is supe ri ority to
,

an im a ls w ould reply i n p ossessi ng a bette r mi nd

, ,
.

And th e re is of c o u rse no d o ub t th a t th e a ve ra g e h um a n
mi nd is ve ry fa r supe ri or to th e a ve ra ge a nim a l min d ,

an d th e best h um a n mi nd i mmea su re a bl y supe ri or to

the best a nim a l mi nd Y e t sigh t mu st not be l ost o f the
f a c t th a t the re i s a m ong men— a s p rob a bly a m ong
.

an im ls
a — a n exceedi n gly wide ra nge i n me nt a l a bility ,

so th a t o n e is f a i rly s a fe i nm a ki n g th e a sse rti onth a t th e

m ost i ntellige nt i ndividu a ls a m o ng a nim a ls a re supe ri or
to th e le a st i n tellige n t h um a nbei ngs N o twithst a ndi ng .

th a t p ossibility th e re a re ce rt a i nki nds o f me nt a l p owe r

,

which a re p ossessed p rob a bly by eve ry norm a l h um a n

bei ng th e u ni ntellige nt a s well a s th e i ntellige nt th a t
, ,

a re n o t sh a red by a n y o f th e l o we r a nim a ls Wh e the r .

these speci a l ki nd s o f p owe r necess a rily indic a te me nt a l

supe ri ority is a deb a t a ble questi on C onside r for ex .
,

a mple th ree ve ry simple i n

,
st a nces : a pp reci a ti on O f the
p ossibilities o f fi re of cl othi ng a nd o f cultiv a ti on o f
, ,

th e so il
F IRE —Expl ora ti ons h a ve c ove red p ra ctic al ly the
.

e nti re h a bit a ble gl obe a nd no ra ce o f menh a s y et been

,

f o u nd which d oes no t k now a nd m a ke use of fi re The .

 ZCL OT H I N G 5 35 .

knowledge th a t fi re ma y be kep t goi ng by th rowi ng w ood

’
! '

on i t f rom tim e to time seems n ot to i n v olve a n ex

cep ti on a lly h igh orde r of me n t a lity I t i s with i n th e
.

sc op e of q u ite y ou ng ch ild re n; yet no c a se h a s eve r

bee n c redibly rec orded O f a ny l ow er a nim a l le a rni ng
h ow to keep up a fi re I t is quite t rue th a t i n a wild
’

st a te a nim a ls a re a fra id of fi re a nd th is fe a r ma y re a dily

,

a ct a s a d ete rre n t influ ence a g a i nst thei r le a rning to ma ke .

u se of fi re .Y et d ogs , which a re a m ong t h e most i ntel

_

l i gent o f a n im a ls h a ve no fe a r O f fire ; i n f a ct th e y
, ,

de rive g re a t e nj oyme nt from b a sking i nits w a rmth ; bu t

a dog wh ose i n tellige nce seems to b e a lm ost huma nwill
a ll o w a fi re to go ou t a n d itself to l ose th e e nj oyme nt o f
i t s w a rmth with ou t a n y e ffort t ow a rd m a i n ta i ni ng it .

Th e sig nific a nce of fi re i nh um a np rog ress is so obvi ous

a s to req i re on ly a w ord reg a rdi ng it I t i s one of th e .

f a ct ors which e na ble mento live outside th e t ropics The .

h ist oric a l f a ct th a t civiliz a ti on th rives a nd h a s th rive n

from th e b egInnIng i n clim a tes too c old for h um a n ,

h a bit a ti on with ou t a rtifici a l p rotecti on emph a si zes t h e ,

si gn i fi ca nce of fi re In view of th e fu rth e r f a ct th a t

.
-

met a l lu rgy a nd th e m ost imp ort a nt a pplic a ti ons O f p owe r

depend up on fi re th e signific a nce of this p a rticul a r
,

a bility o f m a n ki nd bec om es cle a rly m a nifest .

CLOT H I N G — . R el a ted t o th e a pp reci a ti on of fire i s

th e a pp reci a ti on of cl othi ng I t is a common th ing for
.
‘

an im a ls to seek p ro tecti on a g a inst th e i nclemencie s o f

clim a te by c ra wling i nt o sh elte red nooks bu t onl y ma n ,
‘ '

h a s di scov ered h ow to c a rry h is sh elter a rou nd with him

‘

i nt h e f orm of cl oth i ng J ust a s fi re ex te nds h is ra nge;

‘

by all owi ng hi m to live i n c old a s well a s i n w a rm cli

m a tes so cl othi ng i nc rea ses h is e ffective life by a ll owi ng
,

h im to be a ctive wh en th e we a the r i s so i n clement th a t

‘

with out it h e c ould rem a i na live only by cu rli ng himself

up i ns ome sh eltered sp o t a nd w a iting for condi ti onS to
'

m ode ra t e;
T H E CU LTIVATI ON OF T H E S OI L i n v olved i n its begi n
5 36 M AN ’
S P LA C E I N N AT UR E
nings no g re a te r e ff ort of me nta l ity th a nthe observ a ti on
th a t pl a nts sp ring f rom seeds a nd th a t by putti ng a num
ber o f seed s i nt o the g rou n d i n a ccessible l oc a ti ons one
will h a ve i n due t ime a c onside ra ble s tock of f ood co n
ve ul ent to h a nd N o a nima l so fa r a s w e know ca nor
.

d oes ti ll th e s oil The re is sc a rcely a ra ce of menoutside

.

th e p ol a r re gi on s th a t fa ils to p ra ctice a g ri cultu re inso me

form The p rim a ry a dv a nt a ge of a g ricul tu re rel a tes i t
.

self to th e a dv a n t a ges of fi re a nd cl oth ing in exte nding

m a ns ra nge Inth is p a rticul a r c a se the exte nsi on ta kes
’
.

th e di recti onof a ff ordi n g h im time for o th e r occup a ti ons

i nthe i n te rv a ls of food p roducti on P rogress is b orn of
-
.

sp a re time S O l ong a s eve ry w a ki ng m ome nt must be

.

dev ote d to th e fu nd a me nt a l a cti vi ties of s a tisfyi ng h un

ger sec u rin
,
g s a fety a nd pe rpetu a ting th e ra ce the re is
, ,

no O pp ortu nity for p rogress .

Alt h o ugh th e a bility to c a rry on a gricultu re is h e re

st a ted to be a st rictly h um a nch a ra cte ristic it must not ,

be forgotte n th a t a m ong ce rt a i n ki nds of a nts el a b ora te

sys tems of c rop ra isi ng a re m a int a i ned The re a s onthese
-
.

a re n o t a dmitt ed to be o f c omp a ra ble imp ort a n ce with

hum a n a gr icultu re is th a t th ey S h ow no S ig ns O f bei ng
othe r th a n c omplic a ted in sti ncts ; they h a ve no b a sis o f
ge nui ne i ntellige nce th a t is to sa y o f a ss oci a te d
, ,

mem ory a s is p roved by th e a bse nce O f a da pt a bility i n

,

th ei r pe rform a nce Crop ra isi ng a nts ca n be pl a ced i n

.
-

ci rcumst a nces wh e re the c a rryi ng onO f thei r re gul a r t as ks

is fu tile to a bsu rdity yet not th e sligh te st m odific a ti on
,

of th ei r routi n e beh a vi or is obse rv a ble ; th e e nti rely u se

les s m a noeuve rs a re pe rformed with the pe rsiste ncy wh ich
ch a ra c te rizes instincts a nd disti ngu ishes them f rom i h
tell i gen t a c tivities .

L A N GUAGE In a dditi on to th e th ree ex a mples O f

.

me nt a l p owe r desc ribed a bove with reg a rd to which one ,

migh t questi o nwh eth e r they req ui re h ighe r i ntellige nce

th a ndo s ome O f th e thi ngs regul a rly d one by i ntellige nt
an im a ls a nd ye t wh ose po ssess i on by ma n a nd not by
,
 L AN GUAG E 537

anim a ls n
e a bles him to p rogres s a l ong lines which they
c a nnot foll ow th e re is th e me nt a l p rope rty which f orms
,

the t op ic of thi s p a ra gr a ph inwh ich so fa r a s exp eri

ma nst a nds a p a rt from a ll oth er a nim a ls

,

ment reve a ls .

L a ngu a ge i s i n i ts esse nce no thi ng m ore th a n th e a d op

ti on by mutu a l c onse nt of a rbit ra ry sp oke n or w ritte n
symb ols to re p rese nt th ings a cti ons a nd ide a s These
, , .

symb ol s a re empl oyed a s a udit ory or visu a l stimuli

n
,

registe red i n the b ra i as mem ories a n d by th ep rocess,

of a ss oci a ti onc ombi n ed with oth e r mem ories of th e p a r

tic ul a r thi ngs a cti ons , or ide a s with wh ich they bel ong
, .

La ngu a ge mem ories thus bec ome p a rt of th e m a chine ry

o f a s so ci a ti on n a mely o f thi n
,
ki ng i n the hum a nmi n
, , d .

T o ill ust ra te : c onside r th e f oll owi n g seque nce of m a rks

bOok ” Looked a t s imply a s a sp o t fro
. wh ich light m
,

is reflected i nt o th e eye i n a ce rt a in m a nne r with th e ,

initi a ti on of a ce rt a i n visu a l stimulus th is seque nce of ,

m a rks h a s no me a ning wh a teve r But b ec a u se the

.

En glish spe a ki ng ra c es h a ve a greed th a t these m a rks

-

wh enpu t to geth e r i n th is way a re to st a nd i nour mi nds

for the a gg reg a ti on o f p ri nted p a ges wh ich we kn ow a s

a b ook the symb ol a ch iev es g re a t imp ort a n

,
ce I nma n .

th e th ough t p rocesses a re c a rried oni nte rms of l a ngu a ge .

We c a nnot th ink cle a rly a b o u t a nyt hi ng to which we

c a nno t a ssign w ords I t is th is f a ct which m a kes it so
.

difficult f or us to ev a lu a te p rop e rly the i ntel lectu a l a o

ti vi ti es o f a n im a ls wh ich be ca use they h a ve no w ords
,

with wh ich to th i nk — so fa r a s w e ca n j udge — h a ve

th ough ts th a t a re h a zy i ndeed i n c omp a ris on with ou r
own . C ont ra st the educ a ti on of a y ou ng wild a nim a l
with th a t of a child Th e f orme r le a rns ch iefly by ex
.

p e ri ence to s o me ex te nt by o bse rv a ti on B ut on ly thi ngs

which h a ppe nto i ts elf or withi nits ra nge of obse rv a ti on
.
,

ca n p o ss ibly c on t ribute to th e f orm a ti on of th ose a ss o ~

ci a ted mem ories wh ich m a ke up its i n tellec tu a l equip 4

me nt Th e ch ild on th e o th e r h a nd a lth ough b a si ng

.
, ,

much of i ts educ a ti on u p on expe rie nce a nd Obse rv a ti on,

538 M AN S ’
P LA C E I N N ATU R E
p a rticul a rly i n e a rly ye a rs receives in a dditi ona va st
,

a mo u n t o f i nst ructi on by me a ns o f l a ngu a ge The sum .

t o t a l o f the c hild s ed uc a ti o n g a i ned th us excee ds by

’

, ,

m a ny fold th e u tm ost th a t a ny o f th e l owe r a nim a ls ca n

a chieve . As wa s emph a sized i n Ch a pte r X V mem ory ,

an d a ss oci a ti onh a ve thei r p rime imp ort a n ce a s th e me a ns

whe reby th e org a nism is e na bled to p rofi t by exp eri
e nce L a ngu a ge exte nd s this a b ility so th a t i t i ncludes
.

th e expe rie n ce of o the rs b oth p rese nt a nd p a st I t must

,
.

be b orne in mi nd th a t i n th e a bse nce o f l a ngu a ge only

th ose expe rie nces wh ich h a ve h a ppe ned d i rec tly to th e
org a n ism or h a ve c ome withi n its immedi a te ra nge a re
of a n y v a l ue to i
. t B u t with l a ngu a e a d ded ags a fe a t
u re O f a ss oci a ted mem ory th e expe rie n,
ce of a g re a t m a ny
o the r i n dividu a ls bec omes a v a il a ble a s a guide to c onduct .

S tude n ts of hist ory believe th a t sp oke n l a ngu age c a m e

in t o use a m ong hum a n bei ngs e a rlie r th a n w ritte n .

S p oke n l a ngu a ge is na rrowe r i n its a pplic a ti on th a n

w ritte n bec a use of the impe rm a ne nce of th e thi ng which
,

gives rise to the stimul a ti oni nth e one c a se a s c ont ra sted ,

with its pe rm a ne nce i n th e o the r In th e m a i n sp oke n

.
,

l a ngu a ge c ove rs the p rese nt expe rie nce of o th e rs or thei r

'

expe rie nces i nth e rece nt p a st ; the l onge r a go th e exp eri

e nce th e gre a te r the li a bil ity to e rror i n its imp a rti ng .

W ritte nl a ngu a ge onth e o the r h a nd p e rmits the m a ki ng

, ,

O f pe rm a n e nt a nd a ccu ra te rec ords of p a st expe rie nces ,

an d so thei r utiliz a ti oni nide ntic a l f orm a t a ny t ime .

L A N GUA GE As A FA CTOR I N I N DI V I DUA L P R OGRES S .

Co n side red from the sta ndp oi nt o f th e i ndividu a l the re ,

is no d oubt th a t l a ngu a ge as a na i d to cle a r th inki ng h as

c ont ributed g re a tly to th e a dv a nceme nt o f m a nki nd N o .

pe rson with su ffi cie nt i ntellige nce to th i nk a t a l l , bu t '

a chieves much m ore th a n h e o th erwi se wo ul d th rough

his a bili ty to th ink i nte rms o f w ords F rom the st a nd .

p oi nt of the ra ce l a ngu a ge th e sp oke n w ord to a

'

, ,

g e a t exte n
r t a nd the w ritte n in inc a lcul a ble me a su re ,

a rded p rog ress n a ti onby n a ti ona n n ra ti on

h a s f o rw d ge e
 S IGNIFI CAN CE OF LANGUAG E 539

by gene ra ti on TO a pp reci a te th e a dv a nt a ges ma n enl

.

j o ys o ve r th e o th e r a nim a ls i n th is r eg a r d h e h a s only
to be a r i n mi n d th a t no m a tte r h ow much one of the
l owe r a nim a ls migh t le a rn o r h ow m a ny imp ort a nt in
s he migh t m
,

v en tion a ke his a chieveme n

,
t w o uld do his
f ell ow a nima ls a lm ost n o g oo d bec a use h is me a n s of
i mp a rti ng h is k nowledge to th em a re so li mi ted Th e
.

sp re a d of inf orm a ti on f rom ma n to ma n by w ord o f

m outh onth e othe r h a nd ca nbe a nd O fte nis ve ry eff ec
, ,

tive Th e limits to the useful ness of sp oke n l a ngu a ges

.
,

an d by th e s a me t okenthe supe ri ority of w ritte nrec ords ,

lie in th e impe rm a ne nce a nd l a ck of reli a bility of th e ‘

forme r due to th e f a llibility of hum a n mem ory con

, ,

tra sted with th e p e rm a n e nce a nd a ccu ra cy o f the l a tte r .

T H E S I GN I F I CA N CE OF LA N GUAGE T o R A CI A L P R OGRE S S
“

is tw of old Inthe fi rst pl a ce e a ch ge ne ra ti onis e na bled

.
,

to begi n subst a n ti a lly wh ere th e l a st left off On the .

wh ole th e imp ort a nt a ch ieveme nts of m a nkind a re re

,

c orded ge ne ra ti on by ge ne ra ti on; th us they bec ome

,

a v a il a bl e f or succeedi n g a ges a nd do not h a ve to be

redis co ve red In th e se c ond pl a ce a nd p e rh a p s even
.
,

m ore imp ort a ntly l a ngu a ge se rves to m a ke a v a il a ble

,

th e a chieveme nts of i ndividu a ls of supe ri or a tt a i nme nts .

Th e p oi nt h a s been emph a sized i n a p revi ous ch a pte r

( X L I I ) th a t a ce r t a i n pe rce n t a ge o f th e p o p ul a ti on is
defi ni tely supe ri or to th e ra nk a nd file i n i ntellectu a l
p owe r It is a f a c t of h ist ory which c a nno t be g a i ns a id
.

th a t on th e wh ole th e c ont ributi ons to k nowledge up on

which p rog ress depends a re m a de by th e h ighly i ntel
l i gent mi nority Th e a c tu a l p rog ress i tself h oweve r
.
, ,

depe nds on th e a d opti on by: th e ra nk a nd file o f th e

a chieveme n ts of th e i ntellige nt Only by me a ns o f
.

l a ngu a ge w ritten or s p oken i s this a d op ti on b rough t

, ,

to p a ss
Th e em
.

E M OT I O N A L C H AR A CTER I S TI CS .
-
o ti on s or ,

feelings rep rese nt the d riving f orces Of h um a n c onduct

,
.

In ma n, as i n the highe r a nim a ls c on duct Or beh a vi or

,
 40 M AN ’
S P LA C E I N NAT U R E
is m a de up o f ve ry c omplex a c tivities which resul t f ro m
an d rest up o n eq u a lly c omplex se n s ory a nd ne rv ous re
l a tion sh i p s . Among these a s h as a l re a dy bee n point ed
,

ou t ,
a re m a n y simple reflexes m a ny a ctivi ties which ,

th ro ugh f req ue nt repetiti on h a ve bec ome so h a bitu a l a s

to be to a ll inte nts a n d p u rp o ses reflexes a ce rt a i nnum ,

n n
be r th a t est up o a pu ely i tellec tu a l b a sis
r r — i nothe r
w o rd s depe nd up on a ss oci a ted mem ory — a nd fi na lly , ,

an o the r gre a t g ro up i ncludi ng p rob a bly ne a rly a l l our

v oli ti ona l a cts — th a t a re c a rried on a t th e i nst a nce of
some feeli ng S ince th e em oti ons u nde rlie v oli ti on so
.

l a rgely one is e ntitled to spe a k of th em a s the drivi ng

,

forces o f c onduct By wa y of illust ra ti on cu ri osity

.
, ,

a v a rice h a t red l ove ma y be me n

, , ,
ti oned a s such d rivi ng
forces Bi ol ogic a lly , they c ou nt a s f a ct ors i ni ndividu a l
.

or ra ci a l pe rpetu a ti on When one a ttempts to c omp a re

.

the em oti ona l ch a ra c te ristics i n men a nd i n a nim a ls ,

th e difficul ty is e nc ou nte red th a t the re is no me a ns of

j udgi ng wh a t em oti ons a re p rese nt in a nim a ls exce p t
th ro ugh th e study of the b odily m a n ifest a ti ons wh ich a c
c omp a ny them a nd th e a cti o ns wh ich result f rom them .

S o fa r a s this ki nd o f Obse rv a ti on g oes it indic a tes th a t

th e fu nd a me nt a l em oti ons th ose as s oci a ted i mm ed i a tely
,

an d definitely with in divid u a l or ra ci a l p rese rv a ti on a re ,

ve ry simil a r i n ma n a nd i n the highe r a nim a ls In .

a dditi onto these mene xh ibit ce rt a infeelin gs which so

fa r a s ca n be j udged — a re with o ut c o u nte rp a rt a m on

,

g
l owe r a nim a ls Among these a re feeli ngs a ss oci a ted
.

with high m ora l s ta nd a rds ; the se nse O f resp onsi

b i l i ty the tra i ts O f h onesty j ustice a nd ge ne rosity ih
, , ,

flu en ce hum a nc on duc t — an d the ref ore hum a np rogress

p ro fou ndly a nd a re c onseque ntly f a ct ors of p rime
,

imp orta nce .

T H E I N T ERA CT I O N O F T H E E M OTI O N S WI T H I N TELLI

GEN CE . I nthe a ctu a l w orki n g ou t o f me nt a l fu nc ti on
in g i t is cle a r th a t th e em o t i o ns m a y o n th e o n e h a nd
d irec t the i ntell ige nce a nd o n the o the r h a nd th e i ntel
'

,
 INT E R A CTI O N O F E M O TI O N S 541

l i gence ma y a ff ect the m a nifest a ti ons of feeli ng Th e

.

em o ti ona l re a cti ons of a s a v a ge to pe rs ona l a buse for

,

ex a mple a re likely to be ve ry di ffe re nt from th ose o f the

,

highly civilized ma n Likewise the th ough t p rocess itself

.
,

which lies a t the b a si s o f a ll i nte llectu a l p owe r is guided

,

to a ve ry l a rge exte n t by th e em o ti ons Evidently the

.
,

h ighest type of i ndividu a l is one wh o is e nd owed with

gre a t ment a l p owe r a n d i n wh om th e a ctu a l p ro cess of
m enta l fu nctioning is guided by well b a l a nced em oti ons
-
.
 C HAPT ER X LIV

HU M AN P R OG R E S S

P A S T P ROGRES S T H E R EA LI Z AT I O N O F I N H EREN T P OS S I
BI LI T I ES
. Inthe l a st ch a pte r emph a sis wa s pl a ced up on
th e f a c t th a t p rog ress withi n hist oric a l times h a s not
c onsisted — to a ny l a rge ex te nt a t le a st — i n th e de
,

v el op men t o f fu n d a me nt a lly bette r hum a n bei ngs but ,

i n a p a rti a l re a liz a ti on o f the p ossibilities i nhe re nt i n

m a nkind p rimi tive a s well a s m ode rn I t is Obvi ous th a t
,
.

if eve ry org a nism fully re a lizes its i nh e re nt p ossibilities

the re ca n be no fu rthe r p rog ress excep t by imp roving
th e ra ce Th is seems to be ge ne ra lly spe a ki ng th e S itu a
.
, ,

ti ona m ong a ll kinds O f a nim a ls excep t ma n G e ne ra ti on .

by ge ne ra ti on th ey a ppe a r to re a lize a pp rox i m a tely th ei r

,

i nhe re nt p ossibilities I t is pe rti ne nt the ref ore to i n

.
, ,

qui re wh a t a re th e h a ndic a ps which del a y m a ns p rog ress ’

an d th e me a ns by wh ich they a re to be ove rc ome .

I N H ERE N T P OS S I B I LIT IES AR E N OT Y ET F U LLY R EA LI Z ED

bec a use o f diffi culties imp osed by th e e nvi ronme nt I t .

is quite cle a r th a t if th e e nvi ronme nt we re ide a l i neve ry

respect eve ry i n dividu a l w ould ac tu a lly Sh ow a ll th a t h e
is c a p a ble o f By th e exp ressi on ide a l e nvi ronme nt is
.
, ,

me a nt one i nwh ich a ll the c ondi ti ons o f livi ng a nd a l l

th e se ns ory stimuli m a ki ng up th e b as is of beh a vi or a re
th ose best suited to th e welfa re o f th e org a nism a nd th e
ra ce.

A N I M A LS EX ER CI S E A L I M I TED CO N TROL OVER T H E E N

VI R O N M E N T . The study o f a nim a l beh a vi or sh ows th a t
an im a ls do c omp a ra tively little to b ri ng a b out c onditi ons
f a v ora ble to themselves They ma y m ove f rom a n ex
.

p osed sp o t to one th a t is sh elte red or f rom a regi on,

54 2
 M AN CO NT R OL S HI S E NVI R O N M E N T 5 43

wh e re f ood is Sc a rce to one whe re i t is a bu nd a nt or f rom ,

a p er il ous a re a to one o f secu rity ; a side f rom such a c

ti v i ti es a s these th ey a ccep t th e si tu a ti on th a t exists
,

an d get a l ong a s best they ca n D efi nite e ff orts to ch a nge

.

c ond i ti ons for th e bette r ca n h a rdly be s a id to exist

a m on g eve n th e m ost i ntellige nt of a nim a ls o the r th a n
ma n .

M AN LARGELY CO NTROLS HI S EN V I R O N M E N T — Th e .

fa ct th a t h um a n beings do not submit dumbly to u n

f a v ora ble c onditi ons but exe rt th emselves to b ri ng a b out
f a v ora ble c on diti ons lies a t th e b a sis of p rog ress C ivil .

i z ed ra ces su rp a ss th e u ncivilized In th e fi rst pl a c e

bec a use th ey a re h e redit a rily m ore i ntelligent a nd i nth e ,

secOn d pl a ce bec a u se th ey h a ve u n ce a si ngly imp rove d

thei r e nvi ronme nt ; s a v a ges a re on th e wh ole less in
tell i gen t and m ore Submissive a nd exe rcise themselv es
,

on ly to th e p oi nt of a ch ievi ng b odi ly c omf ort p e rs ona l ,

secu rity a nd ra ci a l pe rp etu a ti on In o the r w ords

,
.

s a va ges h a ve u tilize d t h ei r ch a ra cte ristic hum a n t ra its

,

little m ore th a nto Sc re for th emselves th e fu nda mental

bi ol og ic a l needs i n th e e a siest m a nne r p ossible AS
Soon a s thes e n eeds a re met th ey rest c ontent C ivilized
.

ma n on th e othe r h a nd h a s p ushed his c ont rol

, ,

ove r th e e nvi ronme nt f ar bey ond the s a tisfa cti on

of these immedi a te n eeds a nd c ompels i t to y i el d h i m
'

gre a te r a nd gre a te r we a l th o f expe rie nce onwh ich to b a se

b roa de r a nd eve r b road er m ent a l devel op ment ; a nd thus
a tt a i n s a pl a ne o f livi ng i nwh ich th e simple s a tisf a cti on
O f th e fu n d a me nt a l bi ol ogic a l requi reme nts o f f ood
’

secu rity a nd ra ci a l p e rpetu a ti onpl a ys a rel a tively mi nor

,

r61 e

PROGRES S — S lnce p rog ress depen

.

T H E DETA I LS OF . ds
up on th e c ont rol O f the e nvi ronme nt it is evide nt th a t
on e ca nle a rnh ow p rog res s h a s bee nm a de by n o ti n g th e
en vi ron ment a l h a ndic a p s which ma nh a s h a d to ove rc ome
an d by c onside ri ng h ow h e h a s gone a b o u t th e t a sk Of
‘

o v erc omin g them Als o

. s i
,
nce th e h u ma n ra ce i s V e ry
544 H UM AN P R OG R E S S
fa r f rom h a ving a ch ieved th e ultim a te Of p rog ress th is ,

s a me c onside ra ti on o f th e e nvi ronme nt a l h a ndic a ps

sh o uld p oi nt the wa y to fu rthe r p ro g ress .

T H E S AT I S FA CT I O N O F T H E P R I M AR Y N EEDS AN D T H E
E N R I CH M E N T O F L I F E — Th ro ugh o u t th e discussi on of
hum a n p rog ress th e fu nd a me nt a l f a ct Sh o uld not be
fo rg o tte n th a t th e p rima ry bi ol ogic a l requi reme nts o f
food secu rity a nd ra ci a l pe rpetu a ti onf orm the g rou nd
, ,

w ork O f h um a n as well a s o f a l l a ni m a l beh a vi or .

P rog ress c once rns itself fi rst with secu ring for the wh ole
ra c e sa tisf a cti ono f these fu n d a me nt a l needs ; the re a fte r ,

it ma y be dev oted to less immedi a te a lth ough exceed

in gly imp ort a n t dem a nds wh ich ma y be g rouped u nde r
th e ge ne ra l h e a d of enri chment of li fe I n c ont ra sti ng.

an im a l with h uma nbeh a vi or th e t ot a l a bse nce ina nima l s

,

of a n yth i ng th a t te nds t ow a rd e nrichme nt a nd th e i h ,

c re a sing p rese nce O f e nrich ing f a ct ors a s ma n becomes

m ore civilized a re th e outst a nding fe a tu res O f c omp a ri
,

son I nst rivi n

. g to imp rove h is env ironme nt ma ndev ot es
h imself fi rs t to th e s a tisfa c ti on of h is fu nd a me nt a l me
cessi ti es a n,
d th e re a fte r to th e e nrich me n t of h is life In .

the foll owi ng p a ra g ra ph s s ome of the m ost imp ort a nt of

,

the e nvi ronme nt a l h a ndic a ps a re me nti oned a nd the ,

w a ys i n wh ich m a nki nd st rives to ove rc ome them a re

c onside red .

T H E F OOD S UPP LY —E ve ry a nima l c on

. ce rns itself
fir st with secu ring food Th i s is a s t rue of h um a nbei ngs
.

a s of a n y o th e r ki n d o f a nim a l I f th e h um a n ra ce
.

is to su rvive foo d supplies m ust be a deq u a te for its

,

nou rishme nt The re a re no e ff orts di rec ted tow a rd co n

.

tro lli n g the e nvi ronme nt m ore fu nd a me nt a l th a n th ose

which c o nce rnth emselves with food res ou rces F rom the .

rude begi n nings O f p rogress i nwhich seeds we re sc a tte red

i nso il th a t h a d b ee nsligh tly l oose n ed with th e a i d o f a
sh a rp stick m a nki nd h a s see na dv a nceme nt to the p oi nt
,

whe re c o stly sch ools O f a g ricultu re expe rime nt st a ti ons

, ,

an d rese a rch instituti ons a re dev o ted wh olly to the p rob

 S E C UR I T Y OF LIFE 545

lem of p roduci ng c onse rving a nd dist ributi ng f ood

, , .

C ont rol O f the e nvi ronme nt i n thi s respect me a ns n ot

on ly th e p rod ucti on of su ffi cie nt f ood for th e feedi ng

of m a n ki nd but i ts p roducti on i n th e m ost efficie nt
,

m a nner p ossible si nce a s h a s a l re a dy bee np oi nted ou t

, , ,

it is only a s leisu re time is g a ined f rom th e s a tisf a cti on

O f th e fu n d a me nt al requi reme nts th a t ma n is e na bled
to dev ote h imself to the e n richme n t of life E ve n so .

element a ry a p a rt o f the p roblem a s secu ri ng for e very

hu ma n being the a m o u nt o f f ood requi red for h is a de
qu a te no u rishment is still fa r from s oluti on Compete nt .

a u th ori ties a ss er t th a t the re a re milli on s wh o live ou t

th ei r e nti re lives with ou t h a vi ng eve r beena dequ a tely
no ur ished S O l ong a s th is c onditi onp rev a ils the e nvi ron
.

me nt al h a ndic a p imp osed by f oo d sh ort a ge mus t be con

si dered a cute And eve n wh e n p rogress h a s re a ch ed
.

the p oint whe re eve ry h um a n bei ng a lw a ys h a s enough

to ea t th e re will still rem a i n the t a sk of p rod uci ng f ood
,

so efficie n tly th a t a ll m ankind wi l l be a ssu red of a de

qu a te leisu re which h e ma y dev ote to th e high e r ph a ses
o f livi n g .

S E CUR I TY OF LI FE — Th e sec ond of th e fu nd a me nta l

bi ol ogic a l requireme nts is th a t of i ndividu a l secu rity .

Ma nkind st rives to c ont rol th e e nvi ronme nt i n such a

wa y a s to lesse n h is li a bility to i nj u ry a nd de a th Th e .
‘

gre a test me na ce to h um a n secu ri ty h a s a lw a ys bee n

m a ns c om p etitive st ruggle with oth e r org a nism s
’
eithe r
with w ild a nim a ls or with th e org a nism sof dise a se or with ,

o th e r meneithe r i ni n dividu a l st ruggle or onth e wh oles a le

sc a le of wa r P rog ress tow a rd ove rc omi ng th e h a ndic a p of
.

n n
pe s o a l i secu ity is a chieved
r r — so fa r a s d a nge r from
wild a nim a ls or f rom dise a se p roduci ng org a nisms is con
-

n
c e ed
r — by a p o licy o f exte rmina ti on The fi rst c a re o f
.

the pi oneer is to dest roy such wild a nim a ls a s c onstitute

a nimmedi a te me n a ce Th e fi na l a i m O f medic a l scie nce is
.

to exte rmi n a te th e org a nisms of dise a se or to render ma n

'

immu ne to their a tt a cks A p recedi ng ch a pte r h a s em

.
5 46 H UM AN P R OGR E S S

p h a si z ed the d iffi culty of this st ruggle a nd th e g re a t

p rog ress th a t h a s bee n m a d e in s ome d i recti o ns N O .

on e ca nf a i l to re a l i ze h oweve r th a t dise a se still c on

, ,
sti
t u tes a se ri o us th re a t to pe rs on a l secu rity a n d th a t on
, ly
by c ont inued a nd p rol onged exe rti ons on th e p a rt of
medic a l i nvestig a t ors ca n we h op e to see this me na ce
red uced to a deg ree wh e re i t will ce a se to be a V e ry

p rese nt c o nce rn .

T H E D ES TR U CT I VE C O M P ET IT I O N O F M AN wI T H M AN
o n th e i n di vid u a l sca le h a s bee n reduced i n civilized ,

c ommu ni ties to th e a tt a cks up ons ociety by the crimi na l

'

cl a s ses S ecu ri ty a g a i nst th is me na ce O bv i o usly depe nds

.

on i n c re a si ngly efficie nt c ont rol O f c rimi na ls Th e v a st .

des tru cti ve n ess of wa r w i ll be e n ded whe n ma n re a lizes

th a t p rog re ss b oth i ndividu a l a nd ra ci a l depe nds up on
, ,

secu ri ng time a nd opp ortu nity for th ose a ctivities

whi ch e nri ch h is life a nd th a t to th e exte nt to wh ich he
,

dev o tes time a nd eff ort to secu ri ng himsel f a g a i nst th e a t

‘

t a cks of his fell owm a nor to fitti ng himself to a tt a ck them

i n tu rn h e is de n
,
yi ng h imself o pp ortu nity for devel op
me nt a l ong li nes th a t a re re a lly w orth wh ile Th e p ra c .

tic a l me a n s to b ri ng tOa ne nd th e d est ruc tive a c tivity of

ma n a g a i nst ma n must fi na lly be th e a d j ustme nt of
h um a nrel a tio nsh i ps ona b a sis no t o f selfish p rej udice,
bu t of u nselfish a ppli c a ti ono f scie ntific p ri nciples .

COOP ERAT IO N A B I O LOGI CA L N E CES S I T Y FOR P ROGRES S .

At th is p o i nt it w o uld be wel l for th e re a d e r to rec a l l th a t

p ro g ress a m ong a ll s o rts O f o rg a nisms h a s bee nrepe a tedly
sh ow n to depe nd on speci a liz a ti o n a nd h a rm oni o us co
ope ra ti on This is equ a lly t rue of the p a rts of a m a ns ’
.

b ody or o f the i nd ividu a ls o f a nim a l a nd pl a nt a ss o

ci a ti on M uch o f m a ns p rog ress i nth e p a st h a s rest ed
’
s .

o nthis s a me secu re f o u n d a ti o n Wh e nthis t ru th is once

.

fully re a lized des truc tive c ompetiti on wh e the r o f wa r

,

in g n a ti o ns indust
,
ri a l st ri fe o r i nd ividu
,
a l c o mb a t — w i ll
‘

be see nto be a t w o rst a h o rro r a nd a t bes t a silly w a s te

O f time an
. d will be repl a ced by c onst ruc tive e ff ort s
di rected to th e e nrichme nt o f life .
 L E S S E NI NG C HILD MOR TALIT Y 5 47

L ES S E N ED CH I LD M ORTA LI TY — Am ong a nim a ls gen

.

era l l y th e re is a di rec t rel a ti on betwee n th e n umbe r of

Offsp ri n g a nd th e h a z a rds of e a rly life Th ose species i n
.

which the ch a nce of a tt a ining m a tu rity is g ood p roduce

few y o u ng ; th ose i nwhich th e h a z a rds a re g re a t p roduce
l a rge — n
s ometimes e o m ous r — numbe rs With i ncrea s
.

in g civiliz a ti on hum a n f a milies te nd t o bec o me sm a lle r.

Th a t this f a c t c onstitutes a me na ce to the p e rpetu a ti on

o f the highly civilized ra ces c a n no t be de nied M a nmust
.

st rive to ove rc ome th is h a ndic a p a s h e st rives to o ve rc ome

o the r h a n dic a ps w hich i nte rfe re with h um a n p rog ress ,

an d a n evide nt li ne a l ong wh ich a ch ievement i n this

'

respec t is to be secu red is by p rese rvi n g to m a tu rity a l l

the y ou ng th a t a re fit ex a mples of thei r ra ce M uch h a s .

bee n d one i n this di recti on i n th e p a st few dec a des so ,

t ha t a t the p resent time inf a nts h a ve a much bette r

ch a nce o f livi ng to ma tu ri ty th a n did th ose b orn fifty
ye a rs a go M uch rem a i ns to be a chieved i nthis di recti on
.
,

b ut it is re a s ona ble to l ook f orw a rd to a time wh e neve ry

i nfa nt b ornin fa milies of g ood st ock will su rvive to ma
tu ri ty a n d thu s be a ble to be a r i ts p a rt i nthe p e rpetu a
ti onof th e ra ce .

EDU CATI O N . In the l a st ch a p te r p rog ress wa s st a ted

to be c omm on ly b a sed onthe a chieveme nts o f the highly
i ntell ige nt bu t to depe nd on the a d opti on of these
,

a chieveme n ts by m a nki nd i nge ne ra l I t is evide nt th a t

.

si nce this is th e c a se it i s only a s such a ch ieveme nts

bec ome k now n th a t p rog ress will be m a de Org a nized .

s ociety h a s a d opted th e pl a n of setti ng a side the ye a rs

of ch ildh ood a n d a d olesce nce a s a p e ri od o f f orm a l edu
c a ti on na mely , a s a p e ri od du ri ng which sh a ll be m a de
,

k now n to th e membe rs o f the c omi ng genera ti on th ose

fa cts of h um a n a c h ieveme nt up on wh ich th ei r own b e
h a vi or a nd such p rog ress a s they i ntu rnma y m a ke sh a ll
be est a blished Th a t p a rt of th e c ommu ni ty which de
.

v otes itself to the t a sk O f educ a ti on h a s to select f rom

the sum t ot a l of hum a n expe rie nce th ose a chieveme nts
. -
.
5 48 H U M AN P R OG R E S S

which me a n m ost to m a nki nd a nd the n to dr ive them ,

h ome insuch fa sh iona s to m a ke them a ctu a l c ont rolli ng

f a c to rs in beh a vi o r Th a t this t as k is impe rfec tly per
.

fo rm ed eve ry one will a dmit The re rem a i ns the n room

.
, ,

for g re a t p rog ress i nth is field .

I SO LAT IO N . I t is evide nt th a t si nce the a ch ieveme nts

o f th o se wh o h a ve th e a b ility to a cc omplish must be

p a ssed on to the ge ne ra l p opul a ti on if p rog ress is to be

m a de th e ex te nt to which membe rs of the p opul a ti on
,

a re is ol a ted h a s g re a t i nfl ue nce up onp rogress InC h a pte r .

XXX V I I I th e te nde ncy O f a nima ls to sp re a d th rough

o u t regi on s to which th ey a re a d a pted h a s bee np oi nted
o ut. M a nsh a res this te nde ncy with th e othe r a nima ls ;
i n f a c t h e g oes fa r bey ond a ll o the rs S i n
,
ce h is c ont rol ,

o ve r his e n vi ronme nt e na bles him to est a blis h h imself in

sit ua ti ons i nwh ich o the rwise h e c ould no t e ndu re Th e .

i nevit a ble eff ect o f m a ns mig ra ti ons is is ola ti on b oth

’
,

physic a l a nd i ntellectu a l The fi rst is bei ng ove rcome

.

by imp roved me a ns of t ra nsp ort a ti on the l a t te r by ,

p ri nting a nd the a llied a rts .

P RI N T I N G
.
— The re ca nbe no d o ubt th a t th e gre a tes t
si ngle a ch ieveme nt b y whi ch is ol a ti on h as bee n ove r
c ome is th e i nve nti on of me a ns for multiplyi ng rec ords
o na l a rge sc a le P ri nt ing with m ov a ble type forms th e
.

n
b a sis o f this a chieveme t a l th ough — as h a s bee n
p o inted o u t rece ntly — th e i nve nti on of p a p e r m a king
,

h a d p rob a bly a lm o st a s much to do wi th th e a c tu a l

devel opme nt o f the b ook m a ke r s a rt TO th e knowl
-
’
.

edge O f h ow to p rod uce rec ords h a d to be a dded the

k nowledge O f h ow to rep roduce them The re is p rob a bly .

no h um a ne nvironme nt which i s no t suscep tible o f bette r

me nt th ro ugh i nflue nces th a t ca nbe b ro ugh t to be a r by
me a ns o f th e p ri nted p a ge I t is a lm o st imp o ssible for
.

an y one no w to live i nsuch rem o te ness th a t his is ol a ti on

c a nnot i nl a rge me a s u re be relieved th rough the a ge ncy
o f the p ress

T ELEP H O N E T ELEGR AP H AN D R ADIO —

.

, , A gl a n ce a t .
 T R AN S P O R TATI O N 5 49

o ne s m orni ng p a pe r is suffi cie nt to i ndic a te wh a t these

’

a ge n cies a re d oi ng to a nnihil a te is ol a ti on Th e re one .

ma y fi nd wh a t the P remie r o f Engl a nd s a id but a few

mi nutes bef ore th e p rice of st o cks onthe P a r1 s or Be rli n
,

st ock exch a nges th e p rog ress o f a n expediti on a ttempt

,

in g to climb M t E ve
. r est a s well a s th e mul tif a ri o us
,

d oi ngs of h is own Co u nt ryme n All th is is p ossible b e .

c a use news ma y be fl a shed ove r th e w orld a t a lm ost the ,

S peed of ligh t by c a ble or ra di o

,
With r u ra l m a il de
.

live ry p a rcels p ost a nd teleph ones th e f a rme r is not only

, , ,

likely to be a s well i nf ormed a s h is city b ro th e rs bu t o fte n

bette r S o efli ci ent i nf a ct a re the me a ns of di ssemi na t
.
, ,

in g news th a t th e questi o nn o w is n o t one of m ore n ews

but of sifti ng it a nd ch oo si ng only th a t w orth k nowi ng .

TR A N S P ORTATI O N We h a ve the g re a test di ffi culty i n

.

a pp reci a ti n g th e physic a l is ol a ti on a nd c onseque nt utte r

p rovinci a lism of ou r a ncest ors of h a lf a d oze nge ne ra ti ons - -

ago . Wh ole f a milies lived ou t th ei r lives with out eve r

w a nde ri ng so much a s ten miles f rom h o me T o the .

m a j ority O f pe ople th e a dj o ini ng shi re wa s a s u nk nown

an d f oreig n a regi on a s a re th e e nds o f th e e a rth to us
to da y
-
Th e difficulties a nd h a z a rds o f t ra nsp ort a ti on
.

O pe ra ted as effective dete rre n ts to a ll but the a dve ntu rous

or th ose d rive n by n ecessity C ont ra st with th is the
.

p rese nt time i n wh ich a l a rge pe rce nt a ge o f th e p op ul a

ti onis a ble to t ra vel a b o u t M ech a nic a l t ra nsp ort h a s
.

imp roved a nd che a pe ned the me a ns of getti ng f rom pl a ce

to pl a ce u ntil sc a rcely a n y a dult membe r o f a civilized
c ommu nity but h a s s ome fi rst h a nd expe rie nce of dis -

tri cts o th e r th a nhis own T o th e f ree m o ti onO f pe rs ons

.

must be a dded th e t reme nd ous v olume a tt a i ned i n the

t ra nsp ort a ti onof g oods Th us one ma y to da y not only
.
-

g o to th e e nds O f th e e a rth on o cc a
,
si on but h e is d a ily i n ,

c ont a ct with a rticles b rough t f rom its f o u r c orne rs Any .

on e wh o re a ds this will h a ve a b o ut h is p e rs on a d o ze n

things f rom rem ote p a rts of th e w orld : sh oes from A rgen

tine h ides ; c oa t oi Aust ra li a n w ool wi th butt ons of ,
55 0 H UM AN P R OG R E S S

S o u th -
veget a ble iv ory ; a pe ncil m a de in G e rm a ny
sea
,

wi th a ne ra se r m a de f rom the sa p o f a B ra zili a nor S um a

t ra nrubbe r tree ; a S wiss w a tch Egyp ti a ncig a rettes or
, ,

wh a t no t .

A DEQ UATE P R ODU CT I O N AS civi liz a ti ona dv a n

. ces a n d
life bec omes riche r m ore a nd m ore m a te ri a l thi ngs a re
,

felt to be desi ra ble C ont ra st th e Sa tisf a cti on of a n

.

un tu to red s a v a ge i n the p ossessi on o f a few st rin gs of

be a d s a nd a sm a l l b a nd mi rror with th e ye a rning for
books pic tu res music a l i nst rume n
, ,
ts a nd the o the r
,

a ccess o ries O f li fe o nth e p a rt O f the civilized w orld The .

O bvi ous h a n dic a p to th e re a liz a ti on o f th e desi re for

these be n efits is th a t the re a re not e no ugh of them to go
ro un d By wa y O f illus tra ti on t a ke the m a tte r of indi
.
,

vidu a l t ra nsp o rt N o one will de ny th e enorm ous con

.

tri bu ti on a ut om obiles m a k e t ow a rd th e e n richm e n t of

life Any one ma y legitim a tely desi re to sh a re i n th is
.

e nj oyme nt Obvi ously this desi re c a nn

.
,
o t be fulfill ed

u ntil sufficie nt numbe rs of m o t or c a rs h a ve bee np rod uced ,

wi th sufficie nt supplies O f m ot or fuel so th a t th e re will ,

be e nough for a ll The i nc re a si ng desi res of m a nkind

.

c a ll fo r c onti nu a lly i nc re a sed p roduc ti on This d em a nd .

ca nbe met on ly by i nve nti ons a nd disc o ve ries especi a l ly ,

o f m a ch i n e ry which ta kes th e pl a ce o f h um a n h a nds .

Th e limit o f p roductivi ty by h a nd is Obvi ously ve ry fa r

bel ow p rese nt h um a n needs a nd desi res M ech a nic a l .

devices h a ve pl a yed a g re a t p a rt i nh um a na chieveme nt ,

an d must pl a y a n eve n g re a te r p a rt if m a n ki nd is to
c onti nue to p rog ress to th a t p oi nt a t which a l l l egit i m a te
hum a ndesi res a re s a tisfied .

H IGH I DEA LS — In p revi ous p a ra g ra ph s it h a s bee n

emph a sized th a t a n esse nti a l fe a tu re o f h um a n p rog ress
h a s bee ni ts p a ss a ge bey on d th e s a tisf a cti ono f th e fu nd a
me nta l bi ol ogic a l requi reme nts o f f o od se cu rity a nd , ,

ra ci a l pe rpetu a ti o n to th o se a ch ieveme n
,
ts which c onsti
tu te e n richme n t o f life An imp o rta nt eleme nt i n this
.

l a tte r gro up h a s bee n the se tti ng fort h O f high ide a ls of

 I MP UL S E TO CO M B AT 551

h um a nc onduct M enO f supe ri or i ntellige nce f rom e a rly

.
,

h ist oric a l times d ow n to the p rese nt h a v e st ressed the ,

enrichme nt of life th rough th e p ra ctice of th ose t ra its

wh ich we a ss oci a te wi th th e high est a nd m os t u nselfish
st a nd a rds H um a n p rog ress a l ong these li nes d e pe nds
.

up on th e e ffective ness with which high ide a ls ca n be

c a used to p rev a il a m ong men Th is is to be a tt a i ned.

by a c once rted e ffort on the p a rt o f th ose i ndividu a ls

wh o a re su fli ci ently intellige nt to pe rceive th e h i nd ra nce
offe red to a chieveme n t by l ow ide a ls a nd to d rive h om e ,

to thei r fell ow men the duty o f livi n g up to th e high est

!

st a nda rds I t is necess a ry the refore to disc ove r wh a t

.
, ,

ide a ls le a d to p rog ressive a ch ieveme nt a nd the nto i nclude

this knowledge i nth e educ a ti onof eve ry one but p a rti cu ,

l a rl y o f th ose desti ned by high a bility to be le a de rs o f

th ei r fell ow men
-
.

T H E I M PU LS E T o CO M B AT A f a ct of fu nd a me nt a l bi o
.

l ogic a l imp ort a nce a nd one which h a s bee n emph a sized

,
“

r epe a tedly i n p revi o us secti on s is th a t org a nisms c ome

,

i nt o c omp etiti on i n a ttempti ng to obt a i n f ood supplies ,

inm a ki ng th emselves secu re or i np e rpetu a ti ng th e ra ce

,
.

Th is c ompetiti onfrequently le a ds to di rec t c omb a t And .

j ust a s hu nge r is a p owe rful imp ulse to th e t a ki ng of f ood ,

an d its e ffective ness is rei nf orced by the e nj oyme nt

a ff orded by e a ti n g so the re is a n impulse to c omb a t
,

wh ich is a tte nded by feelings o f e nj oyment or s a tisf a cti on

wh e reby the c omb a t imp ulse is re inf orced It i s cle a r .

th a t th e th rill wh ich a tte nds st rife is a ve ry re a l b i o

l ogic a l phenome non a pplyi ng to ma na s well a s to o the r
,

an im a ls a nd mu st b e t a ke n i nt o a cc ou nt th e ref ore a s
, , ,

a f a ct or to be r eck on ed with i n h um a n p rog ress Th e .

rec og n iti onof th e fu nd a me nt a l ch a ra cter of the impulse

to c omb a t h a s led i n s ome i n st a nces to th e exp ressi on
o f th e V iew th a t st rife a m on g h um a n beings is not only
desi ra ble bu t bi ol ogic a lly necess a ry This a rgume nt h a s .

bee na dv a nced a s a j ustific a ti onfor wa r a lth ough p rob a ,

bly not m a ny will be f ou nd a t th e p rese nt time wh o a re

 H U M AN P R OG R E S S

p rep a red to m a i nt a i nthe a rgument inthis ext reme f orm ;

I t a ppe a rs to be s ou n d l ogic to rec o gnize th e exi s te nce
o f th e s trife impulse an d to seek me a ns whe reby it
,

ma y find exp ressi onwi th o u t h a rm ing m a nki nd .

S P ORT S . A suggesti onO fte nm a de a nd one th a t cer ,

fa in ly h a s much to rec omme nd it is th a t i n org a nized ,

c ompe ti tive sp o rts th e impulse to c omb a t ma y be s a tis

fi ed to the be n
,
efit ra the r th a nto the h a rm o f hum a nity .

The g re a t a nd a pp a re ntly inc re a si ng v ogue of c ompetitive

sp orts offe rs th e p o ssibili ty th a t ma n ma y th rough th is
ch a nnel a chieve a t le a st i np a rt such s a tisfa cti onof the
, ,

st ri fe impulse as he desi res with o u t such h i nd ra nce to h is

,

p rog ress a s the dest ructive c ompetiti onof wa r i nevit a bly

e nt a ils
.

CO M M ER C AI L S U CC S S
E — With ou t d oub t much of th e
lu re o f busi ness life lies i n th e Opp ortu nity it a ff ords for
O bt a in i ng the th rill of st rife T O c ompete successfully .

i n th e w orld o f c omme rce ; to pit one s a bilities a g a i n

’
st
the a bili ties o f o the r men a nd c ome off vict ori ous ; to
wield th e p owe r th a t a tte nds success ; for a l a rge f ra cti on
O f civilized h um a n ity these c ou nt a m ong th e sup reme
rew a rds life h a s to off e r Wh ile to th e a bst ra ct th i nke r
.

such ide a ls ma y seem to fa ll sh ort of the best O f which

m a nki nd is c a p a ble th e f a ct ca nnot be de nied th a t com
,

merci a l st rife i ncludes with its u nd oubted evils much

, ,

O f g ood ; while th e st rife o f wa r pe rh a ps n o t wh olly evil

,

i np a st times h as for m ode rn civiliz a ti onno redeemi ng

,

fe a tu re T o substitu te i nd ust ri a l c ompetiti onfor w a rf a re

.

me a ns the n to m a ke a na dv a nce O ne ma y re a s ona bly

, ,
.

l ook f orw a rd to a time wh e n a n a dequ a te substitute for

c omme rci a l st rife w ill be fo u nd wh e na fu rth e r a d v a nce ,

will be a chieved O f th e na tu re O f this substitute th e re

.

is o nly th is to sa y th a t i t must be fo u nded up on th e

na tu re O f ma na s a bi ol o gic a l e ntity a nd no t up on a ny ,

p hil o s o ph ic a l spe c ul a ti o n s up o n wh a t m a n migh t a tt a i n

we re h e di ff e re nt th a nhe is .

F UTU RE P R OGR ES S S O fa r a s we a re a ble to pe rceive

.
,
 R A C IAL B E TT E R M E N T 53

p rog ress i n th e futu re must be i n l a rge p a rt a l ong th e

s a me ge nera l li nes a s i n th e p a st S O l ong a s the re a re.

envi ronme nt a l h a ndic a ps th a t h a ve not bee n fully ove r

c om e so l ong a s a ny i ndividu a l is u na ble to re a lize his
,

i nhe re nt p ossibilities bec a use th e c onditi ons f or such

re a liz a ti onh a ve n o t bee n fulfilled the re is still room for ,

an d need O f e ffor t Achievement h a s g one so fa r th a t i n

.

im a gi na ti on a time ca nbe f oresee nwhe nma nwill h a ve

a tt a i ned to c omplete c ont rol o ve r h is envi ronme nt Only .

whe n th a t time h a s c ome will p rog ress i n th e di recti on

h e re outli ned c ome to a nend .

R A CI A L B E TTER M E N T Alth o ugh p rog ress by c ont rol

.

li ng th e e nvi ronme nt migh t c onceiv a bly thu s c ome to

a n en d th e prog ress o f h um a nity need not st op si nce
, ,

th ere c ontinues the eve r— p rese nt p ossibility of a ctu a l i m

p rovement of th e ra ce I t will be rec a lled th a t i n the
.

discussi onO f p rog ress thus fa r th e p o i nt h a s bee nst ressed

.

th a t the a chieveme nts wh ich m a nki nd h a s m a de a re not

to be a cc ou n ted for b y dem onst ra ble I mp roveme nt i n
hum a n qu a lity Wh e n m a nkind w ith his p rese nt a bili
ties h a s g on
.

e a s f a r a s h e ca n the re will still rem a i n

,

p ossibilities o f b reedi ng bette r men I t is t rue th a t to .

obt a i n i n dividu a ls supe ri or to th e best th a t h a ve yet

lived is a fe a t of gre a t difficulty th e oretic a lly a s well a s
,

p ra ctic a lly But to ra ise th e ge ne ra l a ve ra ge of the ra ce

.
,

physic a lly i ntellectu a lly a nd em oti ona lly is enti rely

, , ,

fe a sible requi ri ng not hi ng m ore th a n th e a pplic a ti on to

,

th e hum a n f a mily of well k now n p ri nciples o f ge netics .

The re a re to be su re g re a t diffi culties i n a pplyi ng these

n
, ,

p ri nciples eve to th e exte nt of ra isi ng the a ve ra ge by

,

cu tting d ow n th e numbe r of disti nctly i nfe ri or i ndivid

ua ls
. T o do wh a t is eve n m ore desi ra ble viz to b ri n g,
.
,

a b o ut a na ctu a l i n c re a se i n th e numbe r o f supe ri or i ndi

v i du a l s is a m a tte r O f g re a te r p ra ctic a l difficulty
, It ma y .

be th a t m a nkind will neve r be a ble to ove rc ome these

difficul ties e nti rely I t is quite su re th a t a t best thei r
.

s luti on ca n only be g ra du a l a nd
o — bec a use of the i n
,
554 H U M AN P R OG R E S S
f reque ncy o f h um an ge ne ra ti o ns wh ich ru n only th ree
,

to th e ce n tu ry — i t will o bvi o usly be a l ong time a fte r

a begi n ning is m a de bef ore st riki ng resul ts ca n be Ob
ta ined. I t i s cle a r h oweve r th a t the s oone r se ri o us
, ,

gene ra l a tte nti o n is p a id to ra ci a l be tte rme nt th rough

euge nics th e be tte r i t will be for m a nki nd b o th i n
,

ne a r a nd i n th e l ong dist a nt futu re .

FIN IS
 IN D EX

Absolu te a chi evement 529 , Algae 1 1 6 ; b a cteri a symbi oti c

,

Abs orbi ng O rga ns l ea v es a s 59 wi th 27 5

.

, , ,

Abs orp ti on 96 ; of f oo d m a teri a ls , , Ali m enta ry tra ct 92 (fi g ) ; m ove

,

38 m e nts o f 96
,

A ccess ori es di eta ry 50 , , Alpine typ e of ma n 502 ,

A ccess ory eye s tru cture s 1 4 5 (fi g ) ; ,

Al te rna ti ono f genera t i ons 3 27 ; i n
,

respira tory or ga ns 1 00 ; sex ,

f erns 33 0 ; i nhigh er pl a nts 3 42
, ,

or ga n s 298 , (fi g) ,
3 4 5 (fi g)
A ccomm od a tion 1 4 5 ,
Am eri ca , ma ni n, 502
A ceti c ferm enta ti on 264 , Ami no a cids , 4 6 , 7 7 , 7 8, 95 , 26 7 ;
A chi evem ent a bso lu te 529; rel a , ,
synth eses of proteins from , 4 7
t iv e 529 ,
Amm oni a , 266
A cid a mi no (see a mino a cids ) ;
,
Amm oni um ca rb ona te , 273
f a tty 26 7 ; l a cti c 264 ; ni trous
, , ,
Amcn b a , 2 1 (fi g ) , 25 (fi g ) , 1 1 9,
27 3 ; u ri c 261 ,
226 (fi g )
A cqui re d ch a ra cters 3 57 ,
Am oeb oid m o v em ent, 22, 1 1 9, 1 22
A ctivi ty nerv ous 7 6 ; refl ex 1 6 1 ;
, , ,
Amyl a se , 6 7 , 263 ‘

v oli tiona l 1 62 ,
Amyl o b a cter , 27 5
A cts v olunta ry 1 62
, ,
Amyl opl a s ts , 64
Ada p ta ti on 4 3 1 ; a djus tm ent i n , ,
Ana ero b e s , 26 7
an im a ls 4 39; convergent a m ong
,
Ana l o gy , 4 1 4 (fi g )
an i m a ls 4 3 1 ; conv ergent a m o ng
, A na ph a s e , 288 (fi g )
pl a nts 4 32 ; f ood secu ring 4 34 ;
,
-
, Ana to my , co mp a ra tiv e , 4 1 4
f or riv a lry 4 3 5 ; f or self de f ens e ,
-
,
A nemi a , 1 89
4 36 A nger, 1 6 6
Adj us tm ent a d a p ta ti ons i n a ni , A nim a l , b eh a vi or, 54 1 ; b ree di ng ,
m a ls 4 39; gra du a l nece ssi ty
, , 395 ; ki ngd o m , 4 1 1 ; p a ra si tes ,
f or 1 69; of l ea v es
, 1 1 4 (fi g ) ; ,
1 99, 20 8; w a s tes , 27 2
necessi ty for 1 0 9; o f orga ni sms , ,
A nim a ls , 1 1 0 ; a djus tm ent a d a p
6 ; pl a n t 4 39; of rep ro du cti on , ,
t a ti o n s in ,
4 3 9; ch emi ca l corr e
1 1 5 ; of roo ts 1 1 0 ; o f s te ms 1 1 2 , , l a tio ni n, 1 7 1 ; cold bl oo d ed , 7 9;
-

A djus tors 1 24 1 4 8; ch emi ca l 1 7 0 ;

, , , co l o r s o f , 4 4 0 ; co nv ergent a d a p
m ech a nis m of 1 4 8 , t a ti o n s in ,
4 3 1 ; dis tribu ti on o f
Adre na l gl a nd 1 7 5 ; h orm one of , ,
l a nd , 4 73 ; f ood re quirem ents of ,
1 75 1 1 0; migra ti ons o f, 4 81 ; of
A dre na l in 1 7 5 , o ce a n i c isl a nds , 4 76 ; o f th e s ea
A erob es 267 , fl oor, 4 90 ; reprodu ctio n a m ong ,
A gri cul tu re 53 6 292 ; s o m e p el a gi c m a n y celled ,
-

—
,

Ai r co mp o si ti on of 98; co nta mi
, ,
4 88; w a rm bl o o d ed , 7 9
na ti on 235 ; dry a nd m o is t 20 5 ;
, ,
A nnu a l produ cti on of f ood , 37 ;
pol lu te d 20 5 ,
v a lu e o f f oo d , 3 7
Album os es 268 ,
A nth eridi a , 328
Al coh ol 4 4 263 ; illi ci t ma nuf a c A nth ers , 33 7
A nti—
, ,

t ur e o f 263 b odies , 23 1
—
,

Al coh oli c f erm enta ti on 262 ,

A nti to xi n, 232
Al co h olism 52 1 ,
A nts , 1 95
5 55
 I ND E X
Anxi e ty 1 66 ,
B od y ce lls , 3 09; s urfa ce , 84
Appe ti te 1 3 7 , la y ers 3 02 ,

A pprecia tio n 1 86 , _
Bo il s 222
,

A rch egoni a 3 29 , B o ta n y 4 ,

A ris to tle 420 , Bo tto m de po si ts 487 ,

Artfi cia l s e lecti on 4 1 8 , Bo tul ism 21 7 ,

As cent O f sa p 5 7 ,
Bra chyd a ctyly 50 7 (fi g ) ,

Asexu a l R e pro du ctio n 3 1 6 ; , e sse n Bra in 1 1 0 1 5 8

, ,

ti s ls of, 3 22 ; in a l n ,
31 7; te Bra ndy j a ck a ss 263
, ,

su l ts O f , 295 B rea thing 1 3 3 ,

As h co ns ti tuents , 50 B ree ding a nim a l 397 ; co tton 393 ;

, , ,

As s o cia tion ,
1 86, 5 3 1 , 537 ; na ture pl a nt 4 3 90 , ,

of , 1 93 Bree ds 4 1 8; imp rov ed 403

, ,

Asso cia te d m emory 1 6 3 , B ri tish me n 520 ,

Asso rtm ent i nd e pe ndent , ,

3 63 ; B uddi ng 293 (fig ) ,

ra ndom 3 1 3 ,
B uds 1 1 7
,

Aus tra li a nra ce of ma n 503 ; ,

rea lm , B umbl e be e 1 94 ;
-
mi mi ck ed by ,

4 79 flies 4 44 (fi g )
,

Au to into xi ca tion 2 1 7
-
, B urb a n k Lu th er 391
,

A u to lysis 260 266 , ,

B urns 1 89,

Aw a re ness 1 65 1 86 , ,
B u tter fa t, 264
Az o to b a cter 27 5 ,

C a ctus pl a n ts , a da p ta ti ons a mong ,

B a ch fa mi ly 520 ,
4 33 , 4 66 (fi g )
B a ck cross 3 6 7 ; o f li nk ed ch a ra c
, C a l ci u m , 38
ters 3 6 9 ; o f i n
,
d epe nd ent ch a ra c C a li f ornia chil dr en, 524
te rs 36 7 ,
C a m e ls , e vo lu ti o no f , 4 1 0
B a c te ri a 2 1 9 (fi g ) ; d e n
,
i trifyi ng , C a ncer, 21 2
27 3 27 6 ; f ree livi n
,
g s oil 275 ; -
, Ca n e suga r, 96
ni tra te 27 4 ; b a c te ri a ni tri fy
, , C a rbo hydr a te s , 7 7 , 92 ; ferm en ta
in g 27 3 ; n
,
i tri te 27 3 ; sulphur , , t i o nof, 26 1
27 1 27 7 ; symbi o ti c 27 5 ; tube r
, , C a rb on, 38; cy cl e , 27 7
cul e 27 6 , C a rb o ndi o xid , 3 8, 4 0 , 81 , 98
B a c te ria l inf e ction 21 2 ; zym a se , , C a rnivo ro us pl a nts , 4 5 , 4 35
263 C a rrie rs O f dis ea se , 221
Ba rk , 56 C a ta lyz ers , 66
B a rri ers , 4 5 1 ; bio lo gi ca l ,
4 56 ; in C a tching co ld , 224
ta ngibl e ,
4 54, 4 75 ; ta n gibl e , C a u ca si a nra ce , 5 1 0
4 53 , 4 7 4 C a us e s O f dis ea s e , 204
Ba s i c m e ta b olism 74 ; eff ect onof , C e ll , co ndi tio n s f a vo ring a ctivi ty
a d re n a li n 1 7 5 1 7 7 , , of , 83 ; divisi on o f, 1 5 , 1 9, 283
B e e co l o n i e s 1 94 , (fi g ) ; grow th O f, 283 ; sa p , 6 1 ;
Bee r wor t m a l t suga r 26 2 ,
s tru ctur e o f, 20 ; surfa ce , 84 ;
Be h a vi or an im a l 54 2 ; h u m a n
, , , th eo ry , 1 5 ; typi ca l , 20 (fi g ) ;
w a ll , 20 (fig )
B en efi ci a l i nse cts , 236 C ells , 1 3 ; a re b a gs o f w a ter, 52 ;
Be tte rm e nt , ra ci a l , 553 bl oo d , 84 ; bo dy 30 (fig ) , 309 ;
B i o l o gy , 3 ; gen e ra l , 5 ; s a ni ta ry , 4 di s cov ery of, 1 3 ; germ , 297 ,
B i rds , mi gra ti o nO f , 1 6 2 30 9, 3 24 ; gua rd , 58; m us cl e ,
B irt h ra te , hum a n, 524 1 22 ; n umber of i n ma n, 84 ;
Bla n chin g, 4 1 x S l ZB o f
,
13
B l a s t l a , 302 (fi g )
u C ellul a s e , 6 7
B l ee d in g, 6 1 C e llul o se , 20, 44 , 265
B loo d , 88; cells , 84 , 229 C h a l k , 4 88
 I ND E X
C h a p a rra l , 4 6 8 (fi g ) C ol ors of a n
i ma l s ; 4 40 ; o f physi o
C h a ra cter expre ssio nof gen es , 381 l o gi ca l v a lue , 4 4 2 ; p ro tective,
C h a ra cters , a c qu ire d , 3 58; co m 4 4 1 (fi g ) ; w a rning , 44 3
bina tion, 359; h eri ta bl e, i nma n, C o mb a t , th e impulse to , 55 1
5 06 ; inde pe ndence O f , 3 59; C o mb i n a ti o n s of ch a ra cters , 359
physi ca l , of ma n, 506 ; physi ca l , C o mm ens a lism , 1 96 (fi g )
of o cea n ,
4 82 ; s econda ry s ex u a l , C o mm erci a l su ccess , 552
1 7 4 , 3 08 C ommu ni ti es , d ef ense s of a ga in st
C h a ra cteris ti cs , e m o tion al h u ma n, disea se , 234
53 9; intell ectu a l , 534 C omp a ra tiv e a n a to my , 4 1 4

C h e mi ca l a dj us ters , 1 7 0 C o mpe ti ti on, i ndus tri a l , 552

C h e mi ca l correl a ti on in a n i m a ls , C o mpl em enta ry genes , 3 86
1 7 1 ; i npl a n ts, 1 7 0 C onj uga ti on, of chr om os o m es , 26
C h emi ca l inju ry , 21 0 (fi g ) , 3 1 0 (fi g ) , 3 4 4 ; \ O f on e

C h emi ca l re gul a tio n, 1 69, 3 27 cell e d org a n is ms , 3 0 1

C h emi ca l s en s e, 1 39, 1 4 1 (fi g ) C onn ecti n g nerve- cell s , 1 55
C h em o tro pis m , 1 1 1 C ons ci o usn e ss , 1 6 5, 1 86

C h ewing , 96 , 1 33 C ons erv a tiono f en er gy , 7 8

C hild morta li ty , 54 7 C onta ct , rea cti onO f pl a n ts to , 1 1 3 ;

C hili s a l t peter , 272 rece p to r, 1 53 ; s en se s , 1 3 9
C hines e ra ce , 527 C onta mina ti on, o f a ir, 235 of

C hl a mydo m on a s , 22 (fi g ) w a ter a nd f oo d , 23 5
C hl oro phyll , 23 , 4 0 ; pigm en ts , 4 0 C ontra ctil e v a cu ol e , 26 , 1 03
C hro m a tin, 20 (fi g ) , 286 C onv er gen t a d a p ta t i on s , a mong
C hro m a to ph ore s , /40 an im a ls , 4 3 1 ; a m ong pl a nts, 432
C hr o m oso m e s , 287 , 3 35 53; conjuga C o Op era ti on , 5 6
4
t i on of , 3 4 4 ; con ta i n th e gen es , C ork , 56
3 53 ; hi s tory of , indi vidu C orrel a ti on, ch emi ca l , i n a n im a ls ,
al i ty of, In p a ir s , 291 ;
_ 1 7 1 ; ch emi ca l , i n pl a nts , 1 7 0 ;
i n m eta z o a , 1 4 9; i n pl a n ts , 1 1 7 ;
fl

mov emcnt of, 288; ra ndom a s

s ortment of , 3 1 1 ; red u ction o f, in p ro to z oa , 1 4 8; m e ch a n i sm of,
3 1 1 , 34 2 ; s e gre ga t i on of, 3 4 2 ; 1 18
S pli tti n g of , 288 C orn, 3 86 , 393
C il ia , 27 (fi g ) , 1 20 (fi g ) C orp s cl es , red, 88; whi te, 88
u
C ili a ry m oti on, 1 1 9 C orr ens , C , 3 5 1
.

C ir cul a tory sys tem of high er a n i C ort i , B , 1 4

.

m a ls , 87 (fi g ) C o tton, 3 93
C l a ss , 4 1 3 C o tyl e dons , 3 4 1
C l a ss e s , o ccup a ti on a l , 522 ; socia l , C ra bs , h ermi t, 1 96
522 Cretinism , 1 83 , 5 1 3
C l a ssifi ca ti on, 4 1 1 —
C ro M a gn onma n ,
4 9 7 (fi g)
C l os tridi u m , 275 Cross , 3 4 9
C l othing , 53 5 C ro ssi ng -
o v er, 3 7 4 (fi g )

C o a gu l a ting enzyms , 68 Crow nga lls , 2 1 2

-

C oa gul a ti onof bl oo d , 88 C rys ta lli ne l ens , 1 4 6

C oeffi cient o f corre l a t ion, 5 1 7 C ul tiv a t ion, eff ects of , 274
C o ld bl o o de d a ni ma ls , 7 9, 1 40
- C umul a t iv e gen es , 3 87

C olds , ca t ching, 224 ; co mm on , C u rd , 26 4

223 C u ttings , 1 1 7
C o llo ids , 4 5, 49, 52 y
C c , l e ca r bo n,
2 7 7 ; e ner gy ,
27 9;
C o l on i a l or ga nisms , 27 ni tro gen, 27 7 ; ph osph oru s , 279;
C ol oni es , b ee , 1 94 sulphur , 27 9
C ol or , percep ti onof , 1 4 6 C y ta s e, 265
C olor b lindne ss , 1 4 6 , 5 1 3 C y topl a sm , 20 (fi g )
 55 8 IND E X
D a ir y ca t tle , inh eri ta nce , 402 D o rm a ncy , O f see ds , 3 4 1

D a rw in ,
C h a r le s , 9, 420 Dro so phi la , 36 5 (fig ) ; d ete rmin a
D a rwins t h eo ry , fa c to rs o f , 4 2 1 ti on of se x i n, 3 66 ; he redi ty of ,
’

D a v e npo rt, C B , 520 366- 384 ; li n ka ge i n, 3 66

—
. .

D e a mi n a s es , 68 D ctle ss gl a nds , 1 7 1
u
D ea t h , 7 , 49, 24 9; a n a dv a nta ge , D ra ti ono f li fe , 24 9
u
255 ; imm e dia te ca u se s O f, 253 ; Dwa rfis hn ess , 1 7 3 , 5 1 2 (fi g )

Weisma ns th eory of, 252

’

D eca y , 25 9 Ea r , 1 43 (fi g )
De cidu o us fore s t , 4 60 (fig ) Ea r ly d ev el o pment o f a n i m a ls , 302
D eco mpo si ti on, 7 , 25 9; a ge nts o f , E a rly hi s tory , o f ge rm ce lls , 3 1 0
26 1 ; k inds O f, 260 ; o f fa ts a nd (fi g ) ; of th e h um a nr a ce , 4 96
O i ls , 266 ; of n i tro geno s w a ste ,
u Eco n o mi c impo rt a nce O f di sea s e,
26 6 204 ; o f pl a n t b re e d in g, 390
D e fe n se , of comm ni ti es , 234 ; a nd
u E du ca t i on, 528, 53 7 , 54 7
of o rga n isms , 228; o f yo ung , Eff e cto rs , 1 50 , 1 54 ,
4 37 E ff ects O f c u l tiv a ti o n ,
27 4 ; of

D ege n era tio n , 202

free zing , 250 ; of ocea ncurrents ,
De n i tri fying b a cte ri a , 273 , 27 6 4 83 ; o f p a ra si te o nth e h o s t, 203
D e pos i ts o f u n i cellul a r orga n i sms , E ggs , 1 63 , 298 (fi g ) , 3 25 , 329, 334 ;
27 , 4 87 m a tura tion o f, 3 1 3 ; pa renta l
D e sert, 4 6 7 ca re o f , 3 0 6 ; wi nter produ cti on
D e te rm in a te ev o lu tio n ,
4 29 o f , 40 1

D e termi n a t io nO f se x , 3 1 3 (fi g ) , 3 66 E mb ry o , 3 29, 33 9
De v el o pm en t of a n ima ls , 302, 4 1 6 Emb ry o lo gy , e vid en ce of ev ol u
D e V ri es , Hugo , 3 5 1 t i onf ro m , 4 1 6
D e x te ri ty, m a nu a l , 534 Emo ti o n s , 1 6 6 , 1 86 , 533 , 539
D i a be te s , 1 7 5 , 1 84 E m o t i on a l ch a ra cte ris ti cs , 53 9

D i a to ms , 3 , 44 , 4 85 (fi g ) E mu lsi on, 6 7
D i eta ry a cce sso ri es , 50 E nd os perm , 3 38; n u cl eus o f, 33 9
D iffe renti a l ge n es , 381 (fi g )
D i f e re n
f ti a t i o n ,
3 0 5 ; o f se x 3 27 En er gy , 1 7 , 3 7 , 3 9, 7 3 , 7 6 ; c h a n ges
D iff si o n
u , 5
2, 84 , 98 in d e co mpo si ti o n ,
262 ; co n

D iges ti o n ,
91 serv a tion O f, 7 8; cy cl e, 27 9;
D i o xid , ca rbo n, see ca rbo ndi o xid ra d i a n t , 3 9; rel e a se i n m e ta b
D iph th e ri a , 23 1 o lis m, 260 ; s o ur ce o f , 1 7

D ise ase s , o f pl a n ts , 204 ; d ue to h a rm En ri c hm e n t O f l ife , 54 4

ful orga nis ms , 2 1 6 ; due to m a l a d
jus tment, 1 79; a nd i nsects , 23 6
D ispe rsa l , m ea n s O f a nima l , 4 7 3
D ispo sa l O f o rga n i c w a s tes , 259
D iss o lved subs ta n ce s , as s timuli ,
1 3 5 ; m o ve m e n t o f i npl a n ts , 56
En v iro nm ent, 1 09; huma n, 5 1 9,
528; co n tro l o f , 54 3 ; id ea l , 54 2 ;
u nsui ta bl e , 204
En zyms , 66 , 7 4 , 7 7 , 96 , 260 ; cla ssi
fi ca ti o n o f, 6 7 ; co a gul a t in g , 68;
d ecompo si ti on, 26 1 ; ferm enta ~

Di s ta n ce , pe rce p t i o n o f , 1 4 6 ; re ti o n , 6 8, 26 3 ; m a n uf a c ture of,

ce p to rs , 1 3 6 , 1 42, 1 57 93
D is tribu ti on, geo gra phi ca l , 9, 4 1 7 ; E pil e psy , 522
l a ws a nd re sul t s , 4 7 5 ; O f a nima l s , E q u i librium s n e se , 1 37 , 1 39
4 73 ; o f life i nth e o cea n, 482 ; O f Ere ct po s t ure , 53 1
—
pl a nts , 44 9 4 7 1 ; of ra i nfa ll , 1 09 E thi o pi a nrea lm , 4 7 9
D ivisio n, ce ll , 1 9, 283 , 285 (fi g ) ; Euge n i cs , 526 , 527 , 554
redu ci n g , 3 1 0 (fi g ) , 342 ; o f E uro pe a nra ce s , l a te r, 5 02
l a b o r, 30 E vid e n ce s o f ev olu ti o n ,
4 08
D o dd er , 1 98 (fi g ) Ev olu ti o n, 9, 40 7 ; d e te rmina te ,
D o mi n an ce , 3 54 4 29; evid e n ce o f , 4 08; o f ca m e l,
 I ND E X 55 9

4 1 0 ; O f h o rs e , 4 09; o f ma n, 4 93 ; Foo ds , 1 7 , 3 7 , 58, 91 ; a nim a l ,

a h
res ul ts of , 4 3 1 ; t h eori e s of , 4 20 s orp tion of 97 ; a ni ma l cla sse s , ,

E x cre ting o rga ns , of a n ima ls , 1 03 o f 96 ; a n

,
im a l diges ti on o f 96 ; , ,

(fi g ) ; o f pl a nts , 6 1 an nu a l p ro du cti on of 3 7 ; a n ,

Ex cre ti on, 6 7 , 1 0 3 nu a l v a lu e o f 3 7 ; conta mi na ,

E xh a us ti onof th e s oil , 27 1 t i on of 23 4 ; for th e y oun

, g
Exi s te nce, s truggle f or, 4 23 pl a nt 3 3 9; pla nt 38; pl a nt
, , ,

Expe rience , 538; m a chinery for diges ti onOf 66 ; pl a nt kinds of , , ,

usi ng , 1 58 64 ; pl a n t s tora ge of 6 3 ; pl a n
, t , ,

E xtern a l s ens es , 1 36 m ovement of 65 ; p oisoni ng of , ,

Ey e , 1 44 ; a cce ssory s tru cture of , 21 6 ; u ti li z a t i onof 7 3 ,

.

1 4 5 (fi g ) ; col or of , 50 7 (fi g ) ; Foo d re qu i rem ents of a nim a ls 1 1 0 ; ,

iris of, 1 4 6 ; retina of , 1 4 5 supply hum a n 544 , ,

‘

Foo d s ecu ring a d a p ta t i on

-
s 4 34 ,

Fa mi ly , 4 1 3 Fora m i n if era 4 87 (fi g ) ,

Fa mi ly gro ups , 1 94 ; rese mbl a nce ,

Fores t coni fero us 4 6 1 (fi g ) ; de
, ,

ci du ous , 4 58 (fi g )
Fa s ci a tion, 20 7 Form , pe rce p tionof , 1 44
Fa t, 7 7 , 92 decompo si tion of , Form a t i onof im a ges , 1 44
262 Free z i ng , efl ects of , 250
’

Fa una , 4 52 Frigh t , 1 66
F1 genera tion, 35 1 Fronta ge , w a te r, 83
F2 genera ti on, 3 5 1 Fros t resis ta nce , 3 95
Feebl e m inde dne ss , 520
-
Functiona l m eta b ol ism , 7 6 , 1 22 ;
Feeli ngs , 1 66 , 1 86 w a s te i n 81 ,

Ferm enta t i on, 26 1 ; a ceti c, 264 ; a l Fu ngi 21 0 ; a nd di s ea s e

, ,
21 9, 224
coh oli c, 262 ; ca r bohydra te ,
26 1 ; Fu ngus p o ta to 2 1 1
, ,

enzyms 263 ,
Fuse l o ils 263 ,

Ferns 3 30 (fi g ) ,
Fusi on tripl e 33 9
, ,

Fert ility soil 1 09 27 1 ; of of , ,

Fu t u re m a ns 504
,
’
,

sup eri or hum a nstocks 525 ,

Fert ili z a t i on a mo ng a nima ls 299 , , Ga l a cto se, 265

(fi g ) ; a m ong pl a nts 338 ,
Ga ll fly , 207
Fert ili z ers 27 1 ,
Ga lls , 20 8 (fi g )
Fever 1 81 , Ga l ton, Fra ncis , 5 1 8, 5 1 9
Fibers m u s cl e 1 26 , ,
Ga m ete s , 297 , 3 24 ; puri ty o f , 352
Fire u se of by ma n 53 4
, ,
Ga m e ti c ra ti os , 383
Fi si on 26 (fi g ) 294
s
, ,
Ga m eto phy te , 3 30 ; f e m a l e , 3 3 6
Fi ttes t su rviv a l of th e 4 25
, , (fi g ) ; m a l e, 332 , 3 38 (fi g )
Fla gell a 22 (fi g ) 28 (fi g ) 1 20
, , ,
Ga nglion, 1 52, 1 54 (fi g )
Fl a gell a te m oti on 1 20 ,
Ga s e x ch a nge i n l ea ve s , 60 ; na tu

Fl a tw orms kid neys of 1 04 ; tra ns , ,

ra l , 44
p orta ti onsys tem O f 85 (fi g ) ,
Ga s es , diff usi onof , 98
Fl a v or 1 4 2 ,
Ga s tri c jui ce , 94
Fl ea s a nd pl a gu e 24 3 ,
Ga s trul a , 302 (fi g )
Fl exi bili ty Of h a nd 53 4 ,
Genera l B i ol o gy , 5
Fli e s ga ll 208; a nd dis ea s e 237 ;
, , ,
Ge n era ti o n,
a l terna t i on of , 330 ;
mimi cking w a sps 4 44 (fi g ) ,
sponta neous 293 ,

Fl oa t ing pl a n t s 4 85 ,
Gene s 3 53 ; ch a ra cter e xpressio n
,

Fl ora 4 52 ,
of , 3 81 ; co mpl em e n ta ry 386 ; ,

Fl ota ti on or ga ns of 488 , ,
cumul a tiv e 387 ; di fferentia l, ,

Fl ow ers a nd inse ct s 44 5 ,
3 81 ; i n ch rom oso m es 3 53 ; in ,

Fl uk es 200 ,
bibi tory 3 87 ; l eth a l 3 99; li nk ed
, , ,

Fly ga l l 20 7 ; vine ga r 3 6 5
, , ,
segrega tionof 3 70 ; mu ta tions of , ,
 IND E X
383 ; pa i red , 356 ; pos i tion o f, Here di ty , 8 3 50 M end el s ’
, , 424 ;
3 7 9; se gre ga t i o n o f, 3 54 ; sup la ws Of 35 1 ; pri nciple s of 3 78;
, ,

p le men ta ry , 3 86, 3 98 t hr o ugh c y to pla sm 357 ,

Ge n e ti cs , 4 1 9; a ppli ca ti o n o f to Heri ta bl e ch a r a cte rs inma n 506 ,

h uma npro gre ss , 553 Hermi t cra bs 1 96 (fig ) ,

Gen i us , inh eri ta nce o f , 520 Hete ro cys ts 3 20 ,

Ge n o type s , 3 62 ; n umber o f, 385 He te ro spo ry 3 3 2 (fi g ) ,

Ge n us , 4 1 3 Hi be rna t ion 99 24 9 , ,

Geo gra phi ca l dis trib u ti on, 4 1 7 Hi gh er a nim a ls ci rcula tory sys ,

Geo tro pism , 1 1 0 (fi g ) te m o f 87 ; kid n , eys O f 1 0 5 ,

Germ ce lls , 297 (fi g ) , 3 0 9, 324 ; High e r pla nts a nd ni tri fyi ng or

ea r ly hi s to ry of , 3 1 0 (fi g ) ; re ga nis ms , 27 4
ducing divisi on O f , 3 1 0 (fig ) , Hill Folk 521 ,

34 2 (fi g ) ; of p la n t an d a nim a l His to ry o f th e h orse ,

4 09; ea rli er,
co mp a re d , 3 4 2 of th e hum a nra ce ,
4 96
Ge rmina ti on a nd spring grow th , Hoe mophili a , 50 7
en zyms concerned in, 69 Ho la rcti c rea lm , 4 7 7
Gia nts , 1 7 3 Hom o l o gy , 4 1 4 (fi g )
Gi f te d chi ldren, 520 Honey-bee 1 94 ,

Gills , 1 00 Hoo d ed p a tterni nra ts , 398 (fig )

Gira ff e , evo lu t i onO f, 4 25 Hoo k , R o be rt 1 4 ,

Gla n ds , d u ctle ss , 1 7 1 ; h orm one Hoo kw orm , 203 , 226

ma nuf a cturing, 1 7 1 ; se creting, Horm one, 1 1 8 1 7 1 , 5 1 3 ; -m a nu ,

93 (fig ) fa ctur in g gl a n ds , 1 7 1 ; O f a d
Glu cose , 96 , 263 re n a l s , 1 75 ; o f inters ti ti a l gl a nds ,
God da rd , H H , 520 . . 1 7 4 ; of p a n crea s , 1 7 4 ; o f p i tu i

Go i tre , 1 7 3 ta ry bo dy , 1 73 ; rela te d to ra cia l

Go n i um , 28 (fi g ) ch a ra cte ris ti cs , 1 7 7 ; O f thy ro id

Gra f ts , 34 5 gla nd , 1 7 2, 1 83 (fig )

Gra nul o b a cter, 27 5 Horse , his tory of t h e , 4 09 (fi g ) ;
Gra s pi ng , 1 3 3 spee d i nh eri ta nce in, 4 02
Gra ss la n ds , 4 63 Hos t , 209 ; e ff ects o f p a ra si te on,
Gra vi ty as a s t imul us , 1 1 0 , 1 35 203 ; re l a ti on o f to p a ra si te,
Gree ks , 502 , 527 21 8
Gree npla n ts , 5 , 3 6 H uma nbeh a vi or, 53 2 ; inh eri ta nce,
Gro u ps , f a mily , 1 94
Grow th , 1 7 , 4 9, 3 05 ; cell , 281 H uma nra ce , ea r lier hi s tory of th e ,
Grow th m eta bo l ism , 7 5 ; w a s te in ,
4 96 ; tra ts , 50 5 i
80 H n er, 1 3 7 ; e o t
u g m ions co nnecte d
Gua n o , 27 2 wi h
t , 1 66
Gua r d cel ls , 58 Hyb rid 34 9; V i go r o f 393
, ,

Gui n ea pigs , coa t-co lor, 3 98 (fi g ) Hyb ridiz a ti on 392 ,

Gu tta ti on, 6 1 Hyd ra 29 (fi g ) 85 293

, , ,

Hyd rol ase s 6 7 ,

Ha bi t forma ti on 1 64 Hypo th esis Vo nBa yer s 43

’
, , ,

Ha dro ma se , 265 . Hys te ri a 1 86 ,

Ha ir huma n, color of, 50 7 ;

,

of, 50 7 Id ea ls , h u m a n, 550
Ha irs , roo t 54 (fig ) , 27 6
,
I m a ge form a ti on, 1 4 4
Ha nd fl exib ili ty of 53 4
, ,
Imm edia te ca us e s o f d ea th , 253
Hea ri ng 1 3 6 ; o rga no f 1 4 3
, ,
Immigr a ti o n, 526
Hea rt 87 ; muscle 1 25
, , Imm orta li ty o f the un i cellu la r or

Hem o gl o bi n, 88, 1 02 , 1 88 ga n is ms , 254

Herds , 1 94 I mm uni ty , 233
 I ND E X 61

I mmu n i ty rea ction, 230 K a llik a k f a mily 521 ,

Improv e d b ree ds , 4 0 3 Ka n r ed wh ea t, 3 91

Indep ende nce O f ch a ra cters , 3 59 K ephi r 265

,

Indi a nra ce , 502 , 5 26 Ki dneys 1 04 , 1 0 5 (fi g )

,

Indi a ncorn, b ree di ng of , 393 ; ori Ki ngd om , a nim a l , 4 1 1

ginof , 393 K oumi ss 265 ,

Indi v idu a li ty of th e chr om o so m es ,

289, 300 La bo r, di v isi onof, 24 , 3 1
Indus tri a l comp eti t i on, 552 La cta se , 265
I nf e cti on s , 20 7 , 21 8; low gra de, -
La cti c a cid , 264
223 ; o v erco mi n g Of, 229; p re La ctose , 264
v en ti onof, 228 La m a rck , J ea n ,
427
In fectiou s dis ea se , 1 80 La m a r ckism , 4 27
Influ ence of D a rwins w ork , 4 20 La nd a nim a ls , di s tribu ti onof, 4 73 ;
’

Ingredi ents , so il , 38 re spira to ry o r ga n s of, 1 00

I nh eri ta nce , h u m a n, 5 05 ;
o f ge n La ngu a ge , 5 3 6 ; S p ok en , 53 7 wri t
n
ius , 520 ; of m e ta l tra i ts , 5 1 4 t en , 53
8
In hibi tory genes , 387 La w of fili a l re gre ssi on , 5
1 7 ; of i n
In jury , ch emi ca l , 21 0 ; m ech a ni ca l ,
. dep end ent a ssoci a tion, 3 59; of
indep endent a ss ortm ent, 363 ; of
In k, 20 9 rea ss ortm en t , 3 5 1 ; of se gre ga
In sa n ity, 522 t i on ,
3 5 1 , 3 56
In s ectiv orous pl a nts , 435 La ws a nd resul ts of dis tribu ti on,
In sects , a nd di s ea se, 23 6 ; a nd —
4 7 5 ; o h eredi ty , 3 50 389; firs t
f
fl owers , 4 45 ; b enefi ci a l a nd la w, 3 56 ; s econd la w, 3 63 ; thir d
h a rmf ul , 236 ; tra ch ea l sys te m l a w, 3 6 4 ; f our th la w, 364 ; fi f th
of , 1 0 1 (fi g ) l a w, 3 7 4 ; S ixth l a w, 3 74
Ins tinct, 1 6 2 ; re p ro du ctive , 1 68 Lea f g a ll s , 208
Inta ngi bl e b a rriers , 4 5 4 , 4 7 5 Lea v e s , a djustm en t of , 1 1 4 (fi g ) ;
In te ll ectu a l ch a f a cteris ti cs , 534 a s a bs orbin g orga ns , 59 (fi g ) ; a s
Intelligence qu o ti en t, 5 1 5 orga n s of ga s exch a nge , 60
Intera ctionof orga nisms , 7 Le g m e s a nd b a cteri a , 275
u
Interb reeding , 3 97 Lengt h Of l if e , 25 1
In tern a l respir a t i o n,
98; s en s es , 1 3 6 Lens , crysta l line, 1 46 (fi g )
In terrel a ti on s of orga ni sms , 4 23 Let h a l gen es , 399

In ters ti ti a l h orm on e, 1 7 4 Leu w en h oek , A v a n

.
,
14
Intes tin a l j u i ce , 95 ; pu trefa cti on, L i c h en s, 1 96
21 7 Lif e , di stri bu ti on of m a rine, 4 80 ;
In ulin, 64 dur a ti on of, 249; enri chm ent
In verta se, 6 7 of , 54 4 ; s e cur i ty of , 54 5

Ir is O f th e ey e , 1 4 6 Ligh t a nd pl a nts , 39, 206 ; a s a

Irri ta bili ty , 1 1 7 , 1 3 5, 1 48 s t im ulus , 1 3 5 ; pen etra ti on of i n

I shm a eli te s , 521 sea w a ter, 4 84

I sol a ti on 4
, 5 9 Li mi ts of ph oto sy nth esis , 4 1
Ivy , 1 1 2 (fi g ) Li n k a ge , 364 ; in vi ne ga r fly , 3 66 ;
s ex , 3 7 4 ; s tren gt h of, 3 7 1
Ja ck a ss bra ndy 263 ,
Lip a s e s , 6 7 , 94 , 266
Ja p a nese ra ce 527 ,
Living caus e s o f di sea se , 207 ;
Ja v a ma na p e 4 97 (fi g )
-
,
m a chi ne, th e, 4 5
Je n ner, E dw a rd 233 ,
Lo co m o t i on, 22, 1 1 9
J oi nt m o tions 1 29 ,
Low er an im a ls , excretory or
J ui ce, ga s tri c 94 ; inte s tina l
, , 95 ga ns of, 1 04 ; w a ter ronta ge in,
f
p a ncrea ti c ,
94 85
Juk es fa mi ly , 5 21 L mi n
u o s ra ys , 39
u
5 62 I ND E X
Lungs , 1 02 ; e xt ent of s urf a ce , M e ta z oa ti o n in, 1 4 9;
, correla dis
1 02 e a se d u e to , 226 ; o t onin , 1 21
m i
Lymph , 88 (fig ) h ds s is
M et o , ta t ti ca l , 5 1 6
Lymph a ti c v e ssels , 89 (fig ) M e ca nra ce, 526
xi
M i ce y ell ow l etha l genes in 399
, , ,

M a chine the living 4 5 , ,

M i croorg a nisms a nd disea se 2 1 6 ; , ,

M a inte na nce O f li fe 6 ,
O f so il 272 ,

Ma la djus tm ents 1 79 , M igra ti ons of a n i m a ls 4 81 ; O f ,

M a la r ia 225 ; a nd m osq ui toe s 23 9

, ,
bir ds 4 81 ,

(fig) M il d ews 2 1 0 ,

M a la r ia l p a ra s i te s 225 (fig ) 239 , , M ilk s o uring of 26 4 ; s u ga r of 96

, , , ,

M a l nu tri ti on 1 88 ,
26 4
Ma l ta se 263 ,
M imi cry 44 4 ,

M a l t-su ga r 94 ,
M inera l sa l ts 3 8 ,

M a na nd a nt a comp a rison 1 95 M is tletoe 1 98 (fig )

M a n—
, , ,

a pe J a v a 4 97 M i tes a nd di sea se 244

,
—
M a n C ro M a gnon 500 (fi g ) ; e v ol a
,
,

,
M o d era tely compl e x tra nsporta
,

t i o nof 4 95 ; fu t ure o f 504 5 25 t i onsy s tems 86

,

5 5 4 ; i n Ameri ca 50 2 ; m od ern ,
, , ,

, M od ern ma n, 500 ; s u b spe cies of

,
— ,

500 ; N ea nd ert h a l 4 99 (fig ) ; Pil t , 503

d own 4 98; R h od es ia 4 99; sub
, , M o d ern sys tem of c a l ssifi ca tion ,

spe cie s oi m od ern 50 3 ; th e ,

41 2
high es t a ni m a l 53 1 ,
M o difi ca ti ons ,
3; Of m enta l
M a nu a l d ex teri ty 5 3 4 , tra ti s 529; Old a ge 252
, ,

M a nu fa ctu re of enz yms 93 ,

M o is ture 1 09 205 , ,

M a rri a ge 527 ,
M o lds 2 1 1 ,

M a teri a ls foo d 3 8; m a nuf a ct ure

, ,
M ongoli a nra ce 50 3 ,

O f 7 3 ; ra w 3 9; w a s te 80 M orpho lo gy 4
M a tu ra ti on of germ cells 3 1 1 —
, , , ,

31 3 , , M orta li ty ch ild 54 7 , ,

(fi g ) M o sq ui toes a nd di sea se 23 9 ,

M ea sles 225 ,
M o ss 3 28 (fi g )
,

Mech a ni ca l inj ury , 1 89, 20 7 , 21 0 ; M o ti o n a m oeb oid 22 1 1 9; b io

, , ,

re sul ts O f roo t s tru cture 55 , l o gi ca l impo rta nce o f 1 22 ; cil i ,

M ech a nism a d j us to r 1 4 8; O f ce ll
, , a ry 1 1 9; cla me s O f 1 33 ; fla ge l
, ,

di visi on 285 ; O f corre la tion 1 1 8;

, , l a te 1 20 ; i n m eta zo a 1 2 1 ; o f
, ,

of cro ssin g over 3 7 4 ; O f rea s , a lim e n ta ry t ra ct 96 ; o f ch ro m o ,

sortm ent 3 60 ; o f tro pisms 1 1 7

, , so m es 288; of j o ints 1 29 ; o f
, ,

M edi ca l s ci e nce a im o f 54 5 ; trea t , , m a teri a ls in pl a nts 56 6 5 ; o f , ,

m ent Of disea se 2 1 4 , p ro to pl a s m 1 4 ; re gu la ti on of , ,

M edi te rra nea nt ype o f ma n 502 , 1 23

Membra ne nu cl ea r 285 , , Mo to r nerv e-c ell 1 53 ,

Mem o ry 1 59 53 1 53 7 ; a sso ci a te d
, , , , M ul a tto 526 ,

1 63 , 5 40 M ul ti cellula r orga nisms , 24 ; d ea th

M e nd e l Gre gor 3 50 (fig )
, , in 255
,

M end el s la ws o f he redi ty 3 50
’

, M us cl e cells 1 22 1 26 ;
, h ea rt , , of ,

M enta l d e fects 522 ; tra i ts mo di , , 1 28; sm oo th 1 25 ; s tru cture of ,

fi ca tio n s Of 5 29 , sm oo th m us cle fibe r 1 28; form , ,

Mes s ma te ism 1 96 (fi g ) , 1 3 1 ; m a k e u p O f sm oo th 1 27 ; -
,

M e ta bo lis m 7 3 82 ; b a si c 7 4 1 7 5, , , , , se nse 1 38 ,

1 77 ;fu ncti ona l 7 6 1 22 ; grow th , , , M us cl es 1 22 ; a tta chments O f 1 29

, ,

7 5 ; pro te in 260 ; w a s te f ul 7 8 , , h ea rt 1 25 ; l ong 1 32 ; sk el eta l

, , ,

Meta llu rgy 53 5 , 1 25 ; sm oo th 1 25 ; s tren g th of , ,

Me ta m orph o sis 3 06 (fi g ) , 1 31
M eta ph a se 288 , M u shroo ms ,
21 0
 I ND E X 5 63

M u ta tions 3 50 ; of,
genes , 383 O rd er, 4 1 3
M y ch orrh i z a 276 ,
O rg a ni c subs ta nces , 1 8; w a s tes , dis
p os a l of, 259
N a egeli , C a rl , 1 5 Orga nisms , a dj us tm ent o f , 6 ; a s
N a m f a mily, 54 soci a tion of, 1 93 ; col o n i a l , 27 ;
N a m es , S ci entifi c, 4 1 3 d efens es of, 228; gro wt h Of , 283 ;
N a t u ra l , fert ili t y o f soi l , 27 1 ; ga s , intera ction of , 7 ; inte rrel a tions
44 ; o il , 27 ; S el e cti o n , t h eo y o ,
r f of , 4 23 ; m u l ti cell ul a r, 24 ; p ro
42 1 du ci ng d ecomp o si ti o n 26 1 ; p ro ,

N a t ure sso cia ti on Of orga nism

of a ,
du ci n g f erm e n t a t i o n,
2 6 2 ; pus?
1 93 ; of de a t h 24 9 ,
f ormi ng , 222 ; u ni cell ul a r a nd
N ea ndert h a l ma n 4 99 (fig ) ,
b o tto m dep osi ts, 4 87
N eg roes 526 ,
O rga n iz a tionof p ro topl a sm , 48
N egroi d ra ce, 503 O rga n s , 32 ; a ccess ory respi ra to ry ,

M ema tode w orms 208 ,

1 00 ; a cce ss ory sex, 298; ex

N eotrop i ca l rea l m 4 78 ,
creti n g, 6 1 , 1 04 ; of fl ota ti on,
N erv e cell s
-
connect i ng
, ,
488; of h e a ri n g, 1 4 3 ; l ea ves a s
m oto r 1 53 ; sensory 1 53
, ,
a bs orbi n g , 59; resp i ra tory , 99;
N erv o us a ct ivi ty 7 6 co ntrol , ,
im ru di m en ta ry , 4 1 4 ; sense, 1 1 0 ;
p a ired 1 85 ; orga ni z a tion
, ,
1 54 ; sex, 3 28

p ro stra tion 1 86 ; system, , ,

1 1 0, O ri enta l rea l m , 4 79
Origi no f S p eci es , D a rwi ns , 42 1 ’
1 48, 1 52
N e u ra s th eni a , 1 86 Orth o genesis , 4 29
N ew La m a rcki a ns ch oo l , 428 O rt o n, D r W A , 395. . .

N i tra te s , 38, 27 1 O s cill a to ri a , 3 1 7 (fi g )

N i trifyi n g b a cteria , 27 3 Osmo ti c p ressur e, 53
N i trify i ng or ga n i sms a n d high er Ov erp ro du cti o n, 4 22
pl a nts , 27 4 O v u l e, 336 , 3 4 0 (fi g )
N i tro gen, 4 5 ; cy cl e, 277 O xida s es , 68
N i tro genous w a s te , d eco mpo si t i on O xida t i on, 76 , 81 , 98
o f , 26 6

N o d ul es , root, 21 2 Pa i n, 1 37 ; e m oti ons co nnecte d

N on -se xu a l r p r du
e o ct i on,
116 wi th , 1 66
N or di c t yp e of ma n, 502 Pa ire d gen es , 356

N osto c, 3 1 8 (fi g ) P a l eo nto l o gy , evidence of e volu tion

N o u rishm ent of th e y o ung , 308 f rom ,
40 8
N u cl ea r m emb ra n e , 20 (fi g ) , 286 , Pa ncrea t i ch orm one 1 74 ; h orm one ,

289; sa p , 20 (fi g ) , 286 , 289; d efi ci ency 1 85 ; j u i ce 94

, ,

spindl e, 288 Pa p er ma ki ng 548

-
,

N u cl eo lus , 20 (fi g ) , 285 Pa ra m oec ium 26 (fi g ) 1 3 5 1 4 8

, , ,

N u cl e us , 20 (fi g ) , 285 ; end osp erm , Pa ra si te 27 a n ,

d h o st s rel a ti o nof , ,

33 7 (fi g ) 2 1 8; a n i m a l 1 99; li fe his to ri es
,

N u tri en t s, 91 of 20 2 ; m a l a ri a l 23 9; s tru ctu re

, ,

of , 20 1
O ccup a t iona l cl a sses , 522 Pa ra si ti c f u ngi , 2 1 0 ; pl a nts , 1 98
O cea ncu rrents , eff ect of , 4 83 ; dis Pa s te u r, Lo uis , 262
tri b u ti onof l if e i nt h e, 4 82 Pea s eed , s tru ct ure of , 3 4 1 (fi g ) ;
O cea ni c i sl a nds , a nim a ls of 4 7 6 ,
h eredi ty of , 35 1
(Edo gon iu m, 3 21 (fi g ) , 3 25 (fi g ) Pectina se , 6 7 , 26 6
O il , fusel , 26 3 ; na t u ra l , 27 Pel a gi c m a ny— cell e d a n ima ls , 4 88
O ils a nd f a ts , d ecomp osi ti onof , 266 Pepsi n, 94
Old a ge m o di fi ca t i ons , 253 P e p to nes , 268

On e cel l e d o r g a n
-
isms , 25 (fi g ) Percep t i on, O f co l or, 1 46 ; o f dis
Ora nge, s ee dl ess , 394 ; trees , 34 6 ta n ce, 1 4 6 ; o f f o rm , 1 44
564 I ND E X
Perm ea b ili t y , 52 Po lyda ctyly , 509 (fi g )
Ph a g ocyte s , 229; a cti o no f, 230 Po pes , 520
Ph e n o t ypes , 3 62 o f se co nd ge nera Po rt ugu ese ra ce , 527
t io n , 385 Pos ture m ovem e n t, 1 33 ; erect, of
Ph osph o re sce n ce , 4 90 ma n, 533
P h os ph o ru s cy cle , 27 9 Po ta s h , 272
P h o to sy n th es is , 39, 27 5 ; li mi ts O f, Po ta ssium, 38, 50 , 27 1
4 1 ; p ro d u cts o f, 43 ; rea cti on s in, Pres s ure , i nth e se a , 483 ; os mo ti c,
42 53
Ph o to tro pism , 1 1 3 (fig ) Prev en t i o n O f disea se , 2 1 3 ; of ln
Phyl um , 4 1 3 fectio n ,
228
P hysi ca l ch a ra cters , of th e o cea n, Pri nti n g , 54 8
482 ; of p rote in s , 46 ; O f p roto Pr in cipl es of d ecomp osi ti on ,
26 1 ; of
pl as m, 1 5 h e re di ty , 3 78
Physi o l o gy , 4 Pro d u cti o n , a fa cto r in p ro gress ,
Physi olo gi ca l ch a ra cte ris ti cs O f p ro 5 50
tO p la s m, 1 7 ; co l ors O f, v a l u e , 442 Pro du cts of phy tosy n th e s i s , 4 3 ; o f
P igm e n t spo t , 23 d ecomp osi ti on, 262; of p utrefa c
Pigm e n ts , c hlorophyll , 40 ; respira t i on ,
268
to ry , 1 02 Progress , m a ns f u ture, 5 52 ; the
’

P il td o wnma n ,
4 98 f a ct of, 532
P impl es , 222 Pro- ph a s e , 288 (fig )
Pin e B a rren s , 46 3 Pro te a ses , 6 7
Pi n es , see ds of , 3 40 (fi g ) Protective res e mbl a n ce , 44 1
Pi n ey s , t h e, 5 2 1 Pro te i n, 4 6, 7 7 , 92, 26 7 ; co mp o si
Pi t ui ta ry gl a nd , 1 84 ; h orm o n e o f, t i on o f , 94 ; m eta bo lis m , 260 ;
1 7 3 ; imp a i rm e t of, 1 84 n physi ca l p ro pe rt ies of, 46 ; syn
Pl a ce o f o rigi n Of ma n, 4 95 ; O f t h esis O f f ro m a mi n o-a cids , 4 7

s to ra ge i np la nts , 63 Pro to pl a sm , 5 , 4 5, 283 ; ch e mi s try

P l a gu e, 24 3 ; a n d fl ea s , 24 3 o f, 1 6 ; dis covery of, 1 4 ; f u nc
Pl a n k to n, 486 t i o n O f, 4 9; m o ti o n O f , 1 4 ;
Pla n t a dj us t m en ts , 4 39; b ree di ng , na me d , 1 5 ; o rga ni z a ti on o f, 48;
4 , 3 90 ; emb ry o , 3 90 ; f o o d , 3 8, org a n i z e d i nto ce lls , 1 3 ; physi
3 90 ; p a ra si te s , 21 0 ; rea lms , 4 56 ca l ch a ra c te ris ti cs oi , 1 5 ; physi o

Pla n ts , 1 10; ca rn i v oro us , 45; l o gi ca l ch a ra cte ris ti cs of, 1 7 ;

ch e mi ca l co rrela t i o n in ,
1 70 ; u nivers a li ty O f, 1 3
co nvergent d a p ta tions a mong
a , Pro to z oa , 1 1 9; co rre l a ti onin , 1 48;

4 3 2 ; co rrel a t i on i n 1 1 7 ; di s ,
m o vem ent i n, 1 1 9
eases O f 204 ; ,
dis trib u tion O f Psy ch o th era py , 1 87
sh ore 484 ; fl oa ting 485 ; food
, ,
P to m a in es , 268

rece ivi n g a d a p ta tio ns in 4 3 4 ; ,

P ty a li n , 93

gree n 5 3 6 ; high er a nd ni tri

, , ,
P u nn e t t squ a re, 3 62

fyi ng orga n is ms 274 ; i nse ctiv ,

P ure l i n es , 3 4 9

O t o us 43 5 ; p a ra si ti c
,
1 98 (fi g ) ; ,
Pure lin e s el ec ti o n , 3 92

re d u cti on divi si o n O f 3 4 2 ; ri ,
P u ri ty of ga m e te s , 35 2
va l ry a m ong 4 38; s eed 3 4 1 ;, ,
Pus fo rm i n
-
g o rga nisms , 222
self d efense i n 43 7 ; trea tm e nt of
-
,
Pu tre f a cti on, 6 9, 259, 265 ; ln
di sea se in 2 1 3 ,
tes tin a l, 21 7 ; p ro d u cts o f,
Pla sm a 88 ,
268
Pl a s t ids 40 ,

Po iso n ing 1 89; by f oo ds 21 6

, , ! u a ra nti ne 23 5
,

Po iso ns 2 1 6 ; o f di ph th e ria 23 2
, , ! ui ni ne 225
,

Po ll e n 3 35 ; t ube s 3 3 8
, ,

Poll u te d a ir 20 5 ,
Ra ces , la te r E uro pea n, 502
 IND E X 565

R a ci al b e tte rm en t , 55 3 ; ch a ra cte r R es u l ts , m ech a ni ca l o f root s tru e

i st i cs , h orm o ne s i n re l a ti o n t o , t u re 5 5 ; o f a s e xu a l rep ro du c
,

1 77 ti o n 295 ; of e v o lu ti o n 4 3 1 ; O f
, ,

R a di a n t en ergy , 39 f ert ili z a ti on 299 ,

R a di o , 54 9 R et i na o f ey e 1 4 5 (fi g ) ,

R a i nfa ll , dis tribu ti o nof , 1 09 R e tt i ng 26 6 ,

R a nd o m a sso rtm e n t of ch ro m o R h e um a t ism 223 ,

s om es , 3 1 3 R h o desi a ma n 4 99 ,

R a ti o , 1 : 2 1 , 352 ; 9 : 3 : 3 1 , 362 R iv a l ry a d a p ta t i ons f or 4 3 7 ;

°

, ,

g a m eti c, 383 ; genotyp e, 385 a mon g pl a nts 4 38 ,

ph enotyp e, 3 85 ; recombi na t ion, R oo t h a i rs 54 27 6 ; tub ercl es 2 1 2

, , ,

3 52 R oo ts 1 1 7 ; a dj us tm e nt o f 1 1 0 ; a s
, ,

R a ts , h oo d e d p a ttern , 3 98
e x cre ti n g orga ns 6 1 : s tru ctu re ,

R a tta n, 4 3 2 of 53 (fi g ) ; t ro pisms O f 1 1 2
, ,

R a w m a teri a ls of pl a n t f oo ds , 39 R o tti ng of w oo d 26 5 ,

R a ys , a ctive i nph o to sy nt h e sis , 4 0 R o y a l f a mili es 5 20 ,

invisibl e b u t ch emi ca l ly a ct ive, R udi m e n ta ry o rga n s 41 4 ,

39 R us ts 20 4 21 0
, ,

R ea l ms , Austr a li a n, 4 56, 4 7 7 ; E thi

O pi a n ,
4 7 7 ; Ho l a rct i c, 4 57 , 4 77 ; S a ge b ru sh pl a i ns , 466 (fi g )
N eo tropi ca l , 4 7 8; O ri enta l , 4 7 9; S a liva , 7 5 , 93
pl a nt 4 56 ; subdivisions , 4 80 S a ni ta t i o n, go v ernment , 24 5
R ea s o n, 1 86 S a p , a s ce nt o f , 5 7 ; nu cl e a r, 285 ,
R ea ss ort m ent , la w of, 3 5 1 ; m ech a 289
nism of , 3 6 0 S a p ro phy ti c fu ngi , 21 0
R ecep tors , 1 24 ; con t a ct , 1 36 , 1 53 ; S a u nders , D r C h a rl es E , 3 4 1
. .

dis ta nce, 1 36 , 1 4 2, 1 5 7 ; di sta nce , S ca rl et fe v er, 225

a dva n ta g e o f , 1 5 7 S ch ool , new La m a rcki a n, 4 28
R ecessiv e , 356 S ci entifi c n a m es , 4 1 3

R ecombi n a t i on , 3 50 ; ra t i o , 352 S cro ful a , 220

R ed corpus cl es , 88 S ea fl oor, a n i m a ls o f th e, 4 90 ; te m
R e du c ing divisi o n , 3 1 0 (fi g ) , 3 4 3 p era tu re o f th e, 4 83 ; w ee ds , 27 2
(fi g) S econd a ry sexu a l ch a ra cters , 1 7 4 ,
R efl e x a cti on, 1 53 , 1 6 1 ; co mpl e x , 309
1 56 , 5 3 1 ; S impl e , 1 53 S ecret i on, 6 1 ; gl a nds , 93
R e ge n era t i o n ,
295 S e cu ri t y o f li f e , 5 4 5
R e g l a t i o n, ch emi ca l , 1 69, 3 27 ; of
u See d , 3 3 4 , 3 3 9 (fi g ) ; d orm a ncy o f ,
m otion, 1 23 3 4 1 ; g ermi na ti on O f , 69; pl a nts ,
R el a tiv e a chi ev em en t , 529 3 42 ; rep ro du ct i o nby , 33 4
R e l e a s e , en e rgy i nm eta b o lism , 260 Seg re ga t i o n, 3 4 5 ; la w o f , 3 5 1 (fi g ) ;
R epa ir , 1 7 o f ge n es , 3 5 4 ; o f li nk ed genes ,
R e p ro du cti o n, 8, 1 9, 292 , 3 1 6 ; a d 3 70
j ustm ent of , 1 1 5 ; a se xu a l , 1 1 6 , S e l ecti on, 392 ; a rt ifi ci a l , 4 1 8; ef
31 6; a s e xu a l , resul ts of , 295 ; feets of, 3 97 ; na t ura l , th eory of,
sexu a l , 1 1 6 , 295, 3 1 6 ; t ypes of, 4 21 ; pure li n e , 3 92 ; s e xu a l , 4 26

3 1 6 ; v eg eta t iv e , 3 4 5 S e l f d e fe n
-
se, a d a p ta ti ons for, 4 3 6 ;
R ep ro du ct ive ins ti nct , 1 6 8 i npl a n ts , 4 3 7
R es embl a nce , f a mily , 5 1 8; p ro S e mi circul a r ca nal s , 1 3 9
te cti v e , 4 4 1 ; o f t wi n s, 5 1 8 S ense , ch emi ca l , 1 3 9 (fi g ) ; orga n s,
R e sis ta nce , dis e a s e , 3 94 ; fros t , 3 95 cl a ss es o f , 1 3 6 ; o rg a n s , i nterna l ,
R e spir a t i o n , 98
o rg a n s of , 99; pig 1 36 ; p ercep ti on, m o ti o ns o f,
ments , 1 0 2 1 33 ; te mp era tu re
1 40 ,

R es ti ng nu cl eu s , 285 ; sp o re , 1 1 7 , s s
S e n e , cont a ct , 1 3 9; e te rna x l ,
1 36 ;
3 20 i
nterna , 1 3 6 l
 66 I ND E X
Se nsi tive nes s , 1 8, 1 3 5, 1 4 8 S te ms , a dj us tm ent of , 1 1 2; s tru e
Se nso ry nerve cell , 1 53 ; nerv es , t ure of , 56 (fig )
1 54 S te ri liz a ti o n, 5 27
Se rvi ces o f s u pply to ce l ls , 82 S t im u l a tio n ,
1 9, 1 3 5
Se x, 3 24 ; d e termi n a ti o n O f , 31 3 S tim u li , 1 8, 1 1 0 , 1 1 7
‘

(fig ) , 36 6 ; diff ere nt ia ti on o f, S to m a ch , 96

3 27 ; cells , 297 ; c hrom o so mes , S to m a ta , 58 (fi g )
3 6 6 , 5 1 3 ; li n k a ge , 3 7 5 ; o rga ns , S to ra ge , 63 , 3 45
328; o rga n s , a ccesso ry , 298 S trengt h o f m u sc l es , 1 3 1
Se xua l ch a ra cte rs , seco nda ry , 30 8; S tri fe imp ul se , 55 1
re p ro du ct io n ,
1 1 6 , 297 , 3 1 6 , 3 22, S tru cture o f cells , 20 (fig ) of fern s,
324 ; se l ecti o n ,
42 6 3 30 (fi g ) ; o f l ea v es , 58 (fi g ) ; o f
S h o re pla n ts , dis tribu t io nof , 4 84 m u scle fib er 1 26 (fi g ) ; o f nerv ,

S ibs , 5 1 9 o us sys te m 1 5 2 (fi g ) ; o f p a ra ,

S igh t , 1 3 6 , 1 44 ; in rel a tio n to si tes 20 1 ; o f roo ts 5 3 (fig ) ; of

, ,

to u c h , 1 46 s te ms 56 (fig ) ,

S impl e re fle xes i nma n , 53 1

S truggl e for e xis te nce 42 3 ,

S ke le ta l mus cles , 1 25 S u bdi visio ns of rea lms a nima l , ,

S ki n , c
o l o r o f , 50 7 , 5 1 0 (fi g ) 4 80 ; pl a nt 4 56 ,

S l a vs , 5 27 S ub spe cies 4 1 3 ; of m od ern ma n

-
, ,

Sl ee p , s ummer, 250 ; wi n te r, 99,

24 9 S u ccess , co mm ercia l , 552
S lee pi n g si ckness , 237 S uga r, 40 , 92 ca n e , 96 ; i nfermen

S m a ll po x, 233
-
ta ti on ,
262 ; m a l t, 94 , 26 2 ; mi lk ,
S m e ll , 1 42 96 , 26 4
S m oo t h m u sc les , 1 25 S ulphu r b a cteri a , 27 3 ; cy cl e, 27 9
S m u ts , 21 0 S u mm er sleep , 250
Soc ia l cl as ses , 522 S u ppl emen ta ry gen es , 3 86 , 3 98

So il , 204 ; b a cte ri a , 27 5 ; c ul tiva S upply , s ervi ces o f, to cells , 82

ti o nof , 5 35 ; e xh a us ti o nof , 27 1 ; S u rf a ce , b o dy , 84 ; cell , 84 , 283
fert ili ty of, 1 09 ; i ngred ients , 38 S u rviv a l o f t he fi t te s t, 425
So u ri n g o f mi lk , 264 S w a mps , 4 7 0
So u th Ita li a nra ce , 527 S w a rm sp o res , 3 22
S pe ci a li z a ti o n,
283 S w a ll owing , 96 , 1 33
S peci es , 4 1 2 S wi ngl e , D r W T 395 . . .

S p ec tru m , 3 9 (fig ) S ymbi osis , 7 , 1 96 (fig ) , 2 1 8, 27 5

S pe e d i n h o rse s , i n h eri ta nce of , S ymp to ms o f dise a se , 1 80
40 2 Syn a psis, 3 54

S pe rm , 29, 298 (fig ) , 3 25 ; m o th er Syn da ctyly , 50 9 (fig )

ce ll , 3 1 0 , 33 6 S ynth esis o f p rote i n s , 46
S pi n dle , nu clea r, 288 (fi g ) S ys te m , co llo id a l , 4 9; fl a tw orm
S pi ro ch e te s , 2 1 9; dis e a se du e to , t ra n spo rta ti on, 85 (fig ) ; nervo us ,
1 1 0, 1 4 8 (fi g ) ; t ra c h e a l of

S pli t ti ng of ch rom osomes , 288 i ns ects , 1 0 1 (fig ) ; mod era tely

S po n ta n eo us ge n e ra ti o n, 29
2 co mplex tra nsp orta tion 86 ,

S po ra n gium , 3 32 S ys te ms tissu es a nd o rga n

,
s 32 ,

S po re 330 ; l a rge , 333 ; p rodu ct i o n

,

i n fe rn s , 3 3 1 ; res ti ng , 1 1 7 , 3 1 8; T a ngib le b a rriers 4 53 , , 4 74

sma ll 3 3 2 ,
T a nni n 20 9 ,

S po ro phyte 3 30 ,
T a pe wo rm 200 (fig ) ,

S po rts 552 ,
T as te 1 4 1
,

S ta rch 44 64 92
, , ,
T a xo nomy 4 4 1 1 , ,

S ta tis ti ca l m eth o d 5 1 6 ,
T ee th ul cera ted 222
, ,

S ta tu re hum a n 5 1 2
, ,
T el eg ra ph 54 8 ,
 IN D E X 67

T e leph one 54 8 , Ul othri x 320 (fi g ) 3 24 (fig )

, ,

T eloph a se 289 (fi g ) , U ni cell ul a r a nim a ls f erti l iz a ti o no f , ,

T emp era tu re 1 09 1 39; a s a b a r , , 30 1 ; imm orta li ty o f 2 54 ; a n d ,

ri er 4 55 ;
,
of th e sea 4 83 ; a s a , b o tto m d ep osi ts 4 87 ,

s t im ul us 1 35 ; sense 1 4 0 , , U nsui ta bl e envi ro nment 204 ,

T endrils 1 1 3 ,
U rea 26 6 27 2
, ,

Th eory cell 1 5 ; of d ea th W eis

, , , U rea s es 6 8 ,

m a nns 252 ; D a rwi ns f a ctors

’
,
’
, U ti liz a ti o nof f ood 7 3 ,

of , 421
Th eories of evo lu ti on 420 , V a cci na ti on 23 3 ,

Th erm a l ra ys 39 , V a cu ol e, 21 (fi g ) ; co ntra cti l e 26 ,

Th erm otropism 1 1 2 , (fi g ) 1 0 3 ,

Thigm otro pism 1 1 3 , V a ri a bili ty i ni nh eri ta nce 5 1 6 ,

T hi rst 1 3 7
, V a ri a t i on 3 4 9 4 24 i n ca rb on di
, , ,

Threa d w orms 200 (fi g ) 208 , , o xid 4 2 ; , qu a nt i ta tive 50 5 ,

Thumb fl e xibili ty of 53 4
, , V a ri ety 4 1 3 ,

Thy roid gl a nd h orm one of 1 7 2 , , V e ge ta t i on typ es 4 52 ; o f th e ,

(fi g ) ; o v er a ct ivi t y of 1 84 ; -
, U ni te d St a tes 4 5 7 ,

under a ctivi ty of 1 82 (fi g ) V e geta t iv e rep ro du cti on 34 5

'

-
, ,

Thyro xin 1 7 3 1 83 , , V erteb ra tes d ev el opm ent of 30 4 , ,

Ti cks a nd dis ea se 24 4 , (fi g ) , 4 1 6

Tilla ge 20 5 , Vess els lymph a t i c 89 (fi g )
, ,

Tissu es 3 2 ,
V es t igia l org a ns 4 1 4 ,

T o a ds tools 21 0 , V i ne ga r fly see D ros ophil a ,

T onsils infecte d 222

, ,
V irul ence 227 ,

T ou ch 1 39; i nrel a t io nto sigh t 1 4 6

, ,
V irus dis ea s e due to 225 ; fi l ter
, ,

T oxi n 23 2 268
, , a bl e 220 ,

T ra ch ea l syste m of i nsects 1 0 1 ,
V i ta mins 91 , ,
1 88; d efi ci ency i n ,

(fi g) 1 82
T ra i ts hum a n 50 5
, ,
V o i ce p ro du cti on 1 33 ,

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V o li ti on 1 6 3 1 86 54 0, , ,

T ra nsp orta tion 54 9; system fla t , ,

V o li t i ona l a ct ivi ty 1 6 2 ,

w orm 85 (fi g ) ; systems mod

, ,
V o lunta ry a cts 1 62 ,

e ra tel y compl e x 86 (fi g ) ,
V o lv o x 28 (fi g ),

T rea tm ent of disea s e i n pl a nts Vo n B a y er s hyp o th esis of ph oto

’
,

21 3 synth esis 4 3 ,

T revi ra nu s , Go tt f ri e d , 1 4

T riple f usi on, 33 9 Wa r 54 6

W a rm —
,

T ro pisms m ech a nisms ,

of , 1 1 7; of bl oo d ed a ni m a ls 7 9 1 4 0 , ,

ro o ts 1 1 2 ,
W a rning col ora ti on 4 4 3 ,

T rypsi n 94 ,
W a s te 81 ; a nim a l 272 ; decomp o
, ,

T sch erma k , E . V .
,
351 s i t i o n of n i trogenous 266 ; dis ,

Tub ercul e b a cteria , ,

27 6 ; roo t , 2 1 2, p osa l of o rga ni c 259; i n f u nc ,

27 6 ti on a l m et a b oli sm 81 ; i ng ro w t h ,

Tub ercu losis 220 ,

m eta b olism 80 ; solid ni troge ,

Tub es p o ll en 3 3 8
, ,
no us 26 7 ,

Tu rgor 6 1 ,
W a ter 38; conta mina ti on of 235 ;
, ,

Twi ns resembl a nce of 5 1 8

, ,
m ovem ent of inroo ts 56 ; f ro nt , ,

Typ e Alpine 50 2 ; M edi terra nea n

, , , a ge , 83
50 2 ; N ordi c 502 ; v e get a t i o n , ,
W eeds sea 27 2 , ,

4 52 W eigh t of ma n 5 1 1 ,

Typ es of rep ro du cti on 3 1 6 ,

W e ism a nns t h eory of d ea th 252’
,

Typh oid fe ver 221 ; a nd fli es , ,

Wh a l e convergent a d a p ta ti o ns i n
,

23 7 th e, 4 3 1
568 I N D EX

Wh ea t , Ka nre d , 3 91 ; Ma rq u is , Wo u nd sh oc k , 1 89
3 91
bl oo d corp us cl es
-
, 30 (fig ) , Y ea s ts 262
,

88, 229 Yell ow fev er a nd m osq ui toes 24 1.

Wi ne disea ses of 262

, , Yo gu rt 265
,

Wi nte r slee p 99 24 9 , , Yo u ng no u rishm ent of th e 30 8

, ,

Woo d 44 ; rotti ng o f 265

, ,

Woo d s s tu dy of h ere di ty 520 Z oo lo gi ca l rea lms , 4 7 7

’

Worms buddi ng i n 294 (fig ) ;

, , Z ool o gy , 4
p a ra si ti c 200 208 , ,
Z o os po res , 3 22
Worry 1 66 ,
Z ym a s e , 26 3