AN I N TR O DUCTI O N

GEN ERAL B I O LOGY

W I LLI A M T . S E DG W I C K, PHD .

M ew
o oo
of B i l g
y i n !b e M assa c h us ett s I ns t it ut e f
o Ted no lo g ,

AND

E DMUN D B . W I LSON PH D , . .

Pr of e o
ss f Zoolog
r o
'

in Col mbi Colleg

u a e N w Yo , e rk

S E CON D E D I TI ON , R E VI S E D A N D E N L A R GE D

47139
N EW Y O R K

HEN R Y HO LT A N D CO MPAN Y

56 3 6 7
‘ 89 9
 PR E FAC E TO THE FI R S T E DI TI ON .

S E V E R A L y ea rs ago it was o ur good fo rtu ne to follow as g ,

r ad
u ate s t ude nts a co urs e of lect u re s and p ra c t ical stud y in G e neral
,

B io log u n de r the di r ectio n of P r ofe ss o r M a rtin at Joh ns Hop

y ,

kins U n iver sity S o intere stin g and suggestive was the ge neral
.

m ethod em ploy ed in thi s cou rse w hi ch in its main outli n e s had

, ,

bee n m a rked out by Hu xley and Ma rti n te n y e a rs before th at ,

w e we r e pe rs uad ed th at begi nn er s in biol o gy should al w ay s be in

t ro duc ed to the subject in s o m e simila r w ay .The pre se nt w or k
th us o w e s its o rigi n to the influe nce of the authors of the
”
E le m e nta r y B iology ,
our deep i ndebted ness to whom we
g r a tefull
y ackn owledge .

I t is still an ope n que stio n whethe r the begi nn er shoul d pu r

s ue th e logical bu t difficult course of w o r king up w a rd s fr o m the

s i m ple to the co m plex ,

or adopt the easier and m ore prac tical
m ethod of worki ng do wnw a rd s fro m fam ili a r higher for m s .

E ver y teache r of the subject kn o w s how gre at are the p ractical

d ifficultie s be settin g the n ovice who p rovided fo r the fi r st ti me
, ,

w ith a co mpo un d m icro scope is co n fr o nted w ith Y ea s t P roto

, ,

c occ u s,
or A m oeb a ; an d on the other hand how h a rd it is to sift
,

out w h at is ge n er al and e ss en tial fro m the heterogen eo us detail s

-

o f a m amm al or a flo w eri n g pl ant . I n the h O pe of le ss e nin g the

pr actica l diffic ultie s of the logic al m ethod w e ve n t ure to submit a
c o u rs e of preli m ina r st ud w hich w e ha v e used for so me ti m e
y , y
with our o wn classe s and h ave fo un d p rac tical and effective
, .

I t has n ot bee n ou r am bitio n to prep a re an exh austive t r ea

tise We h ave sought o nl y to lead begi nner s in biology from
.

fam ili a r facts to a bett er k n o w ledge of how living thi ngs a r e

bu ilt and h ow they act such as may rightly take a pl ace in g
, en

11 1
iv PR E FA CE TO THE FI R S T E DI TI ON .

e ral ed ucation or may afford a basis fo r fu rther s tudi es in G eneral

B iology Zo ology B ota ny P hys iol ogy or M edi c in e
, , , ,
.

B elievi ng th at biol ogy should follow the ex ample of ph y si cs

and c he m i stry in di sc ussi ng at the o uts et the f und ame ntal prop

erties of matter and e nergy we h ave devoted the fi rs t th ree

,

chapters to an eleme ntary account of livi ng matte r and vi al en

t
o rgy I n the ch apte rs which follo w the se fa ts are appl i ed by
.

c ,

a fairl y e xh austive study of a rep re se ntat i v e an im al a nd pl ant o f

,
consider able though not extreme complexity—a method wluc h
, ,

we believe aff ords in a give n time a bette r k no wledge of vital

, ,

pheno mena th an can be ac quired by m ore s uperfi c ia l s tud y of a

l arger number of fo r ms We are sa ti sfied th at the fe rn and the
.

earth wor m are for thi s purpose the best a v ail able orga ni sm s and ,

th at their st udy can be made fr uitful and i ntere sti ng The last .

c h apter co mpri se s a brief acco unt o f the pri nciple s and o utli n es

of classific atio n as a guide in sub seq ue nt studie s .

A fter thi s i ntroductory st udy the st ude nt w ill be w e ll pre

p ared to t ake up the one celled o rga ni sm s a nd ca n pass ra pidly
-

over the ground covered by suc h w ork s a Huxley and Ma rt in s s ’

“ ” “
P r acti c al B i o l ogy B rook s s
,
’
Ha ndbook of I nv e rte br ate
”
Zo ology ,
A rth ur B a r ne s and C o ulte r s
, l t
P an D s i sectio n
”
or
’

the second p art of thi s book which is we ll in h and and will

,

probably be ready in the c ou rse of the follo wi ng y ea r .

The dire ctions for practi c al study are i nte nded as sugges tio ns ,
not sub stitute s for individ ual effo rt We h a ve strive n to mak e
, .

the work useful as well in the c lass roo m as in the labo ra t o r -

y ,
and to thi s en d have i n trod uced
many ill ustr atio ns The ge ner .

o sit
y of a frie nd has enabled us to e nli st the skill of ou r f rie n d
Mr James H E merton who has draw n most of the o r g l
i ina
. .
,

figures from nat ure u nder our directi o n We have also bee n
, .

gr eatly aided in the preparation of the figur e s b Mr Willia

y m .

C l aus of B o ston .

SE PTE M B E R , 1886 .
 P R E F A CE TO THE S E C ON D E DI TI ON .

I '
r was o riginally ou r inte ntion to publish this wo r k in two
p a rt s ,
the fi r st wh i ch
,
appea red in 1 886 bei ng i n te nded as an
,

int r odu ctio n while the seco nd was to fo rm the mai n b o dv of the
,

wo r k and to i nclude the study of a se ries of t y pe fo rms The -

p r e ss u r e of othe r wo rk h oweve r del ay ed th e co mpletio n of the

, ,

seco nd pa rt and mea nw hile seve ral labo r ato ry ma nu al s appeared

,

w hich in la rge measu r e obviated the n eed of it N eve r theless .

the u se of the int r oducto ry volu me by te a che rs of B iology ,

a nd its sale s lowl

, y but s teadil
y in c r eased I t s oo n appe a red
.
,

h o w eve r that in so me cases the wo r k was bei ng e m ploy ed no t

,

me rel y as an int r oductio n as its autho rs i nte nded but as a

, ,

c o m lete cou r se in i ts el f ; though the w i sh was ofte n exp r es ed s

p
tha t the n u m be r of ty pe s we r e s o me wh at l a rge r The se facts .
,

a nd the m a n obv i ou s defects in the o r igi nal volu m e i nd uced

y ,

us to u nde rt a ke the p r epar atio n of a seco n d and exte nded editio n .

With increased expe rie nce ou r ideas have u ndergo ne so m e

c ha nge . We a r e as fi r mly co nvinced as eve r th at G e n e ral B iol
og y as an i n trodu cto r
y s ubject is of the ve r fi r s t i m po rta nc e ;
, , y
bu t we ar e equally per su aded th at it mu st not tre s pass too fa r
U po n the special p r o v i n ces of Zoology and B o ta ny The p r ese nt
.

e diti o n ,
the refo r e diffe r s fr o m the origi nal in these r espects
,

fi r st while the i nt r oductio n has bee n exte nded so as to in

,

c lude r ep r ese n ta tives of the u nicellula r o r ga n i sm s (A mwba ,

I nf us oria P rotoc occus Y eas t9 B a cter i a ) the publicatio n of a

, ,
-
, ,

seco nd volu m e has bee n aba ndo ned I t is hoped that the w ork .

as thu s exte nded may se r ve a double pu r pose v iz eithe r to ,

.
,

be u sed as an i nt r oductio n to sub seque n t study in Zoology B ota ,

ny o r P h y s iolog ; o r as a co m plete ele m e n tar co u r se fo r

, y y
ge ne r al s tude nts to who m the m i nuti ae of the se more s pecial sub
ec ts a r e of l ess i m po r tance th an the fu nd am e n ta l facts of v it al
j
s t r u ctu r e a n d fu nctio n We believe that a sou n d k n owledge of
.

v
 PR E FA CE TO THE SE CON D E DI TI ON
vi
.

these fact s can be convey ed by the m ethod of study h ere ou t ‘

lined ; but w e must emph atic all y i n si st th at n eithe r th i s no r any

other method w ill gi v e g o od r e sult s un le ss rightl y u sed an d th a t ,

thi s work is not de signed to be a complete text book P robabl y -

.

few teacher s w ill fi nd it desirable to go over the whole of th e

ground here laid out and we hope th at still fewe r w ill be i ncl i ned
,

to confine their w ork strictl y to it E ve n in a brief co u r se th e .

t de t m afte r goi g ove r ce r ta in portio n of thi w o k b e

r
n s s
s u n a
y ,
,

ma a de cq ua i nted with the le ad in g t y pe s of pl a n t s a n d a n nu als ;

d thi m be r pidl acco m pli shed if the i n troducto r y w ork
a n s a
y a
y ,

however li mited has bee n ca refully done I n ex te nded cour ses

,
.

we h ave so m eti me s found it de s i rable to postpo ne ce rtain pa rts of

the introductory work returni ng to the m at a l ater period
,
.

A seco nd modification co n si st s in placi ng the s t ud y of th e

anirn al before th at of the pl ant w hich pl an o n the whole a ppear s
,

de sir able especi ally for students who h ave not bee n w ell t rai ne d
,

in other br anche s of sc ience The mai n r easo n fo r thi s lie s in th e

.

greater ease with which the ph ysiology of the a ni mal can be ap

roac h ed ; for the r e is no do ubt that begin ne rs fi nd the n utritiv e
p
problems of the pl ant ab struse and diffi c ult to gr as p u ntil a ce r
tai n famili arity with vital phenome na h as bee n attai ned ; whil e
mo st of the ph ysiologic al activitie s of the ani mal ca n be readily
illust rated by well k nown ope ra tions of the h uman body
-

The third ch ange is the o mi ssio n of the l aboratory directio ns ,

these h avi ng been found u nsuitable The need s o f di ff e re n t .

teacher s diff er so widel y that it is i mpossible to d raw up a sc he me

th at sh all answer for all I n pla ce of the l abo ra to ry directio ns f o r
.

st ude nts we h ave therefo re give n in an appe ndi x a se rie s o f p rae

, ,

tic al sugge stio ns to teacher s le a v ing it to the m to w o rk o ut de

,

t ailed directi ons if de sired by the help of the st and ard l a bo ra

, ,

tory manual s The se sugge stio ns are th e re sult of a good de al of

.

experienc e on the p art of m any te acher s be side s ou r sel v e s and ,

we hope they will be found useful in proc ur i ng and prep ari ng

materi al (ofte n a matter of consider able di ffi c ult decid
y) a n d in ,
ingj ust wh at the student ma reaso nabl be expe ted to do
y y c .

For the re st the origi nal matter h as bee n thoroughl y r evised

,
,
num o s
er u erro r s h ave bee n c orrected and many additions made , ,
parti cul arly on the phy si ological side .

S E P TE M B E R , 1 89 5 .
 TAB L E OF CON TE N TS .

CHA PTE R I .

I N TR O D UCTOR Y .

L ivingth i ngs and l i f el ess th i ngs Th e co nt ras t and th e l i k eness b etween

.

l i vingmatt er and l i fel ess matte r T h e j o urney o f l ifel ess ma tt er .

th rough l i vi ngth i ngs A n l o gy bet w een a f un tai n a fl am o r a

. a o , e

w h irl poo l and a l i v i ngo rg

,
an i s m L i v i ngmatt er i l i fel ess matt er in
. s

a pec ul ia r t a t o c o ndi t i o n
s e r I t c h arac t ri t i c p ro per t i es
. B io l og
s
y e s .
,

its sc e pe a nd its s ub di vi s i o ns Th e B i l o g i ca l sci nces Th e rel atio ns

. o e .

o f B i l og o y to Z oo l o gy a n d B o t a ny Mo r ph l o g y a nd P h y i ol ogy , o s .

Defi ni tio ns and i nt er rel a t i ns o f th e bi l o g -

oi c l sci ences Psy c h o l o a .

og y , S o c i o lo g y D e fi ni t
.i o n o f G enera l B io l og y .

CHA PTE R II .

Their oc cu rrence and th eir iz e O rga nisms c ompo sed o f o rgans Func
s . .

t i ons O rga ns co mpo sed o f t i ss ues Diff er ntiat i n Ti ss ues c o m

. . e o .
~

powd o f c ell s Defi nit i o ns . Uni ll ul a r o rga ni s ms L i v i ngorg an

. ce .

is ms co n ta i n l i fel ess m tt er L i f l ss m tter o cc urs in l i vi ng

a . e e a

t i ss ues and c ll s E x ampl es L i fel ss matter i nc reases rel ati v ly

e . e e

w i th ag S ummary s ta tem ent o f th e s t r u t ure o f l i ving th i ng

a nis m as a w h o l e—
e s c

th e B ody —
. .

Th e o rg mo re i mpo rtant th an any of its

parts 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

C HA PTE R III .

P R OTOPL A S M A N D TH E CE L L .

Protopl asm th e p h y sic a l b as i s o f l i fe His to ri c a l s k etc h ”

“
Th e c om .
.

po un d m i c roscope a nd th e di sco v ery o f c ell s in co r k Th e ac h ro ma tic .

o bj ec t i v e T h e c ell th eo ry o f S c hl ei den a nd S c h w a n u
.
-
Virc h o w .

a nd Ma x Sc h ult z e M odern mea ni ngo f th e te rm cell

.
“
Th e d is .

co v ery o f pro t o pl as m and sa rco de and o f th eir essent ia l si m i l ari t y .

v ii
viii TA B L E OF cozv m zvrs .

P m

gg
n M o hl C oh n S c h u lt z r a n c e a n d s t ru c t ur e e A ppea
l‘
ffp n of c S m nt ti n f th ggdaff e entizi t m q tjl é
o
.

d l d
l
.
l
.

A t i l
ca ce ll I ts p a r t C y tO p l s m a nd th e uu le s s t
l y p i
.

r a s .
.
o

tiss ues th e gne i of th e bo dy and th ph y io l ogi es l ém

Th e orig i ll g e s . e t io n o f
e a o
”
o e e ,
t
o
t
.

s
e s
e s s ,

m o r it
,
A b
l abor Pr to pl sm t work o M a u u l r tr t s
c a n a i o n
;
az
.

: :
sc a o c a
o

l
.

i Ci r
o

t t / u t io n

?
l R o a
“
o n i n N it l l a n d A n u ”
a r ts c a
e a s t
tr ve
a s :
m
.

r t l h ir ll f d r w r t C l y o t i o n T 1 (3
r ases Vit 1
i i a
th e p o op m in as c o p a -
e s s i e o . .

sou c
r es o f p
r o t o p
l a m i c n e r g y M t a b o ls m a n d it
s p h e . e is a

m i al rel a t i ns 0f
.

ene g
r y d e not mp y i
o s l vi t l f o rc T h e c h e a
“
a e .
"
c o

r o to p l s m a p r t eid c
: a r b h y d o
t es n d f ts
s, P h y s ca l R e l at i o ns
o ra a a . i
p
,

temperature m u e i t r e l,
c t r i c i ty
o s e tc T h p ro t o
, p l as m o f p l ants a nd
e ,
. e

of ani mals similar

C HA PTE R IV .

THE B I OL OG Y OF A N A N I MA L : THE COMMON E AR T H WOR M .

A represeutativ e an rth w rm t ak en s a ty pe Th eir wi de dis

i mal . Ea o s a .

t ibu
r tio n T h e co m m o n e
.
r th w o r m I ts n m e ; h b i t t ; h ab i ts ; a . a a a

food ; asti ng ; i nflu nce on so il ; buri l o f o bj ec ts ; se nses I ts

c s e s a .

d ifferentiati n a nt er po t eri r a n d do rso v ent r l

o I ts y mmet ry :
: o s o a s
-
-
.

bi l at ral nd serial Pl n o f th arth w rm s bo dy O rgans f th e

e a . a e e o
’
. o

body and th e detai l f th ir arrangem nt in y stem ali menta ry ; s o e e s s :

cir l t r
cu a o x r t r r ir tor m to r n rv o us sens mve etc
y; e c e o y ; es p a y; o ; e ; ; . . 41

C HA PTE R V .

THE B I OL OG Y OF A N ANI MA L : THE COMMON E AR T HW OR M ( C nti o n ued )

.

Defi niti on r producti n The grm c ll S ex ual and as ex ual rep ro

of e o . e -
e s .

du t i on R eg
c en r t i n
. Th e repr duc t iv e y te m f th ea rth w rm
e a o . o s s o e o .

It pul t i n nd g
s co gl ay i n g T h p ro
a o f f r ti l iz ti o n
a nd th ee e c ess o e a a
-

.
,

seg mentat i n l v ge f the gg Th maki ng f the b ody Th e

o o r c ea a o e . e o .

g trul a Th th r gr m l y er
as . ec t o b l t nt b l t meso b l as t
e ee e -
a s : as , e o as , .

B r i f tat m nt f th ph eno men o f c ll di vi i o n

e s e e nd o f nuc l ea r
o e a e -
s , a

divi i n s ok a y ok in si Th e m ki ng f th o rg
or ansr Th e f a te o f e s . a o e .

th e g erm l ay rs The g er m pl as m
-
e .
-

C HA PTE R VI .

THE B I OL OG Y OF A N A N I MA L : THE CO MM ON EART H WOR M ( C ntinu d) o e .

l h e mi crosc o pic anat omy h i st ol gy o f th e ea rth wo rm Th e f unda

‘
or o .

mental animal t issues nd th ir c o nst i t u nt c ll ul r el me n ts E pi a e e e a e .

th li l mu c ul ar ner v ous
e a ,
ger mi n l bl oo d a nd c o nnec ti v e tiss ues
s , , a , , ,
and th ir di tr ibut i n in th
e vari o us rga ns Mic ro co pi c t ruc t ure
s o e o . s s
o f th e b o dy w ll ; o f th e al i menta r
y ca nal ; o f th e b l oo d v ess el s ; o f
a
-

th e di epi ment s ; o f th e ner v o us s t em l i

ss
y g g a et
n s , a c
 TA B L E OF CON TE N TS 1x

C HA PTE R VI I .

THE B I OL OG Y or A N M AL : THE C O M M ON E AR T H WOR M tCon tinued . )

PA G ‘
r
G ene al P h y s o l og y Th e an ma l a nd its en v o nment
i . Defi n t o ns i ir . ii .

A dap ta t o n s t uc t u a l and f unc t i o na l o f o g

i , a ni s m to en vi o n men t
r r , r r .

ri i
O gn o f ad a pta t o ns E ff ec t o f th e pe s i ste nc e a nd ac c umul a t o n
i . ir r i .

r
N a t u a l se l ec t on th o ugh t h e s u v va l o f t h e fittes t Th e need o f an
i r r i .

nc o me o f f oo d to s upp l y ma tt e a nd ene g
i y N a t u e o f th e nc o me r r . r i .

Th e f oo d and its j o u ney th o ug h th e body A l ment at o n D ges r r . i i . i

t on and abso pt o n C cul at o n Met a bo l s m Th e o ut go I nt e
i r i . ir i . i . . r
a c t o n o f t he an ma l a nd t h e e nv o nme nt
i i
S umma y ir . r

C HA PTE R VI I I .

THE B I OL OG Y OF A P L A N T : THE COMM ON B RAKE OR FE R N .

A representat i ve pl ant Ferns tak en as a t y pe Th eir w i d di t ri b ut i o n

. . e s .

Th e co m mo n b ra k e I ts name h a b i ta t s iz e etc G ener l m o r ph l

.
, , , . a o

og y o f it bod y I ts diff erent iat i n nt ero pos t eri r a nd do rso v nt ra l

s o a o e
- -
.
, .

I ts b il at ra l y mm t ry Th e underg
e r und tem O ri gi n nd arra nge
s e . o s . a .

ment o f th e l ea v es I nt rna l s t ruc t ure o f th e rh i z o me a nd t h e th r e

. e e

grea t t i ss ue sy tems Th e l m nta ry t i ss u o f pl a n ts Hi to l g

-
s .
y of e e e es . s o

t h e rh izo me R oo t s a nd b r nc h E mb ry ni t iss ue a nd t h a pi a l
. a es . o c e c

cell Ho w th e rh iz o me g
. ro w T h e f nd l a f o f H ome a nd its s . ro or e

st ruc t ure C hl o ro ph y ll bodies S t o ma t a

.
-
. .

C HA PTE R IX .

THE B I O L OG Y OF A PL A N T : TH E C O MM ON B R A KE ( C on ti n u ed) .

Th e v ario us meth ods o f reprod uc ti n in Pteris S po ro ph o re and o .

‘

o op h o re A lterc atio n o f gnera t i ns

. S p r ng ia S po res G er e o . o a . .

mi n t i o n o f th e po res
a P ro t o n ma P ro th ll i um s T h e sex ua l . e . a .

o r gn A n th eri d i a
a s . M le g er m c ell s A rc h g o ni a Fema l e gr m
. a -
. e . e

cell s Ferti l iz t i o n
. S egment at i o n Di ff eren t i a t i o n o f th e t i s ues
a . . s

C HA PTE R X .

TH E B I OL OG Y OF A PL A N T : TH E CO M MON B R A KE ( Co n tinued) .

Ph ysi o l o g
y Th e e n a nd its e n
. o n me n I ts adap a on fr A defi ni v ir t . t ti .

tion of e lif
T h e need o f an nco me o f ma e a nd ene g
.
y I nc o me i tt r r .

o f Ptoris I ts po we o f ma ng oo ds es pec a y s ta c
. T h e c irc u r ki f . i ll rh .

l i
at o n o f oo ds o ug t he p a n f ody Me a o sm th r h
Ou g o R es ~ l tb -
. t b li t . .

pira tio n I n e ac o n o f th e e n a nd th e e n
. t r ti
o n men S pec a fr v ir t . i l
 TA B L E OF CON TE N TS .

PA G ‘

h y$1010 o f t t h u sy t ms nd o f r
e
r t
iss
i o n T h
e
e q ues ti o n
s e a ep oduc .

Iiil oltd il; r l wpi th ni mal s in gen ral Th phy iol gi al im

-

r i f r W th t h e e a r th w o r m a nd o f
A c o m n f th e n a so o e i ,

a
e e s o c
an s in g ene a a .

p
ort nce o f th e c
a h lo o h
p y lless p l ant s r
p

C HA PTE R XI .

CE L L UL A R OR G A N I S MS
’

THE UN I .

b r i i t i d b t i c o mpl ete cell

Th e m u lt i cell ul ar od y I t s g n in .
n n u e u n o co , ,

i ll l r b dy I t o ri g i n t c ed t o m pl t e c ll ra o c e e
d ivi i
s o n . T h u n
lt
u
i ll
e
l r b
ce a
i ll l r
o

b o dy as
. s

m d n d t h e u n c e u a
d iv i i n
s o T h u c u e o y a
r
e
i m
a
ec ia l
”

i ndivi du l s Uni ll ul r form ph y i l gi ll y r g

.

s o o c a a
o n S p s a s s .

a ce
i m r
.

i mport nce o f th ir tr uc t ur l
a
i m ep l i c i t y s
O g n s e d u c ed to
a s .
a s

”
th eir lowes t terms .

C HA P TE R XI I .

UN I CE L L UL A R ANI MA L S .

A . A M( E B A .

rl b it t r t rt o eus a n ma c u e A i l l
“
Gene a A ccou n H a a F o m .
P . p . Th e .
o

pearance P s eu d o.
p o i
d a L oc oti
m o o n F ee d s T h e e nc y s ed s a e
. . . t
t t .

tr t r
S uc u e o f t h e un i ll l r b
ce u a o dy C y tO p I as ni N u c e u s V ac uo es . . l l
. .

r ti fi i h i l f m s o og t l h i l i l
R ep o duc o n b y ss o n P y s o o gy T h e u nd a en a p y.
ca .

r r ti
p o pe es o f p rt l
o o p a s m a si l
d s p ya e d in A m oeba T h e u es on o f . q ti
old a e g l t
R e a ed o ms
.
f r h
Th e Ri z o po da o r ps eudo pod a P ro o zoa . i l t .

s un ani m l c ul e Th e Fo ra menif era

”
A r ll
ce D j flug
a . ia Th e t .
-
a .
.

The R adio laria

CHA PTE R XI I I .

UN I CE L L UL A R A N I MA L S ( C ntinued)
o .

B . I N FU SO R I A .

General account Hab i ta t .Th e sl i pper a ni mal c ul e Th e

. b ell “ -
.
“

ani mal c ul e Pa ra mwczum I ts f o rm s t r uc t ure and h a bi t s Cy t o

’

.
. . , ,

p l as m ; t ri c h o c y s t s ; v ac uo l es ; n u c l e i ; m o uth ; ( e so ph a g us ; a nal s po t .

T h e ency s ted st at e R eproduc t i o n b y ag . a m og enes i s ; b y c o nj ug a t io n ;

am ph i mixi s Vortzcellcr I ts f o rm st ruc t ure etc I ts re pro duc tio n

’

. .
, , .

b y fi ss i o n e ndo g ,eno us divi si o n a nd c o nj ug at i o n Mi c ro g a met e a nd , .

macro g am et e R el at ed f orms E ug lena ; Zootha mn to n; Ca rcheszum;

' ’

. .

M is ty lis ; etc h i ol gy o f th e I nf u ri Herbi v ro u ca ni

. P ys o so a . o s, r v

r
o ens , a nd o mniv r u i nf u o ri o o A nal g y w i th h i g
s h r f rm T h e s a . o e o s .

pr b l
o m of c
e hl roph y ll in ni m l Sym bi i V geta t i ng ni m l s
o a a s . os s . e a a .

Th e cl a i m o f uni ll ul r ni mal to b e r grd d as uni c ell ul r

ce a a s e a e a or

ganisms ; o
”
rga ns in th c ell ; tc e e
 TA B L E OF CON TE N TS . xi

C HA P TE R XI V .

A . P a o roc oc c us '
.

PA G B

G enera l account Hab i ta t M orph o l og . y S t ruc t ure Mot i l e and no n. . .

mo t i l e s ta tes R ep ro duc t i o n b y fi ss i o n. C ell aggrega t es P h y si .

-
.

o l ogy I nco me a nd o utg

. o T h e ma k i ngo f st arc h f ro m i no rg a ni c .

matt ers Th e f unda ment a l ph y si ol o g

. ica l pro perti es o f pro t o pl as m as
dis pl ay ed b y pl ants C o mpari s o n o f P rotococcus w i th A mazba a nd .
,

c hl o ro p h y ll beari ngpl a nt s in g e ne r a l w i th a ni mal s in

-

general O th er .

unic ell ul ar c hl o ro ph y ll b eari ngp l a n ts : dia t o ms ; des mi ds ; Ck roOcoc -

C HA PTE R XV .

UN I CE LL UL A R P L A N TS ( Con tinued) .

B . Y E A ST .

G enera l accoun t . Wi l d y eas t a nd do mes ti c at ed y ea s t . Mi crosco pi cal

x i ti o n o f a y eas t c k e Mo rp h o l gy f t h e y ea t cell C y to
e am na a . o o s .

pl as m and nuc l eus R prod uc t i n b y b udd i nga nd b y s pores . P h ys i

e o .
o

o l og y Y as t nd th e env iro nment

. e D ri d y eas t
a I nc o me M ta . e . . e

ho l is m O ut go Th e mi ni m l nut rien ts of y eas t c o mpared w i th

. . a

th os e o f Pr tococcus a nd A moeba Wh y y eas t is r grded s a pl ant

o . e a a .

To p y eas t B tt o m y eas t Wil d y e t R ed y eas t

o Fer menta t i o n . as s . .

an d f ermen ts Uni c ell ul a r pl a nts no t nec essa ri l y at th e bo tt o m o f

.

th e s ca l e o f l i fe : etc .

C HA PTE R X VI .

UN I CE L L UL A R P L A N TS ( C o n ti nued ) .

C . B A C TE R I A .

Th e s ma ll es t most , bi q ui t o us of k no wn l i v ing
nume ous r an d mo s t u
th i ngs Th ei r ab undance i n e rth i mi l k w t r tc C o mp ri so n
. a ,
a r, , a e . e . a

o f th ir w r k in s o i l s w i th th t
e o f ear th wo r ms P ra i ti c a nd sa p ro a o . a s

h
p y t i c b ac t e ri a T h i r b o t a ni c l p i t.i n S a ni t a ry a nd eco no mi c
e a os o .

i mport anc M rp h l og
e .
y S t r uc t ure C y to pl s m and nuc l eus
o o . . a .

C i l ia Th eir size S w r mi ng nd th res ti ng t ge R prod uc t i o n

. . a a e s a s . e .

E ndospo res A r th r po res

. Ph y i o l g y I nco me os Meta b o l is m
. s o . . .

O utg o Ferm nt
. Fer me nt t i o n eP ut ref c t i o n
s . Di e se O ne a . a . s a .

s pec i es c a pa b l e o f l i v i ng u po n i no rgni c m tt er R e l t ed f o rms a a . a .

W hy bac t ri e r eg
e a r ded a s pl a nt
a ar Th rel a t i o ns o f b ac t eri a to s . e

t m pera t ure moi t ure po iso ns et

e ,
S t eri l iza ti o n P s teurizi ng
s , ,
c .
, a ,

di i n f c ti o n filt ra t i o n tc
s e , , e . .
xii TA B L E OF CON TE N TS .

CHA PTE R XVI I

A HA Y I N F US I ON .

n o:

G eneral accoun t .
lts o f mic rosc o pi ca l ex ami nati on Turb idi ty
R esu . .

O d o r.
Co l o r C. o n t i t u n t s Ts h e s ene o f i mpo rt a nt p h y si
e . l c ca ,

c h e m i c al a,
n d bi l o i
g p l
o h n o mena ca P r v i ou h i s to ry o f th
e h ay . e s e

a n d t h w
e a t er E f f ec
.
t o f b r i n g i n g th e m t g th r C uses o f t ur o e e . a .

b id ity , c o l o r o,
d r t c o ,
A e r bei a n.d na ro bi c bac t eri oth ri v e c a e a .

I nf u oria mult i pl y nd dev o ur th m

s a Car ni vo ro us i n f uso ria attac k e .

t h e h e r b i v o r o us T h e . s t r ugg l e f x i t e n ce H a y a g ree n p l an t
or e s .

a nd th e so urc o f foo d
e Qui et fi nall y up r venes Ho w n ut r i ti ve
. s e .

eq ui l ibri um may b e pre r v ed o di t ur b d

se Th e h ay i nf u i o n an r s e s
-
.

epi t ome of the liv ing world

A P PE N DI X .

S UGGE S TI ON S FOR L A B OR A TOR Y S TUDI E S A N D DE MON S TR A TI ON S .

B ook s f or th e l ab o rat ory Ti me req uired f o r G enera l B i ol o g

.
y 205
S pecia l s uggest i o ns f o r l abo rato ry w o rk etc upon th e s ubj ec ts , .
. t reated
in the v
l ch apt r a o utl i ned bo ve v iz
se er
a e s s a ,

C h apt r I I nt r duc to ry
e . o

I I S t ruc t ur. f L ivi ngO rgni sms es o a . 206

I I I P rot pl m nd th e C ll o as a e

IV—
.

VI I I Th e
. .

I X XI Th e Fe
.
-
. rn
XI I . A moe ab .

XI I I . f ri
I n us o a .

XI V .

X V Y eas . t
XVI B aC leI I a
.
‘
o o o o o o o o o o o o o c o o o o o o o o o o o o o o o o o o o o ' O O O O O 0 0 0

XVI I . A Hay

I N STR UM E N TS A N D UTE N S I L S 0 0 . 0 0 0 0 0 0 0 0 0 0 00 0 0 0 0 - 0 0 . 0 0 Q 00 0 0 00 0 0 0 0 0 0 0 0

R E A GE N TS A N D TE C HN I CA L ME THO DS
 GE N E R A L B I OL O GY .

C H A PTE R I .

I N T R O D U C TO R Y .

WE from co mm o n expe r ie nce that all mate r ial thi ngs

kn o w
ar e eithe r dead or ali ve o r m ore accu ra tely that all matter is
, , ,

eithe r l ifeless o r livin g ; and so f ar as we k no w l ife ex ist s o nly ,

as a ma nifestatio n of liv ing m atte r L ivi ng matte r and l ifeles s

‘

matte r ar e eve rywhere totall y di s tin ct though ofte n closel y as ,

soc iated The m os t ca r eful studie s have o n the whole r e nde r ed

.

the dis tin ction m o r e clea r and st r iki ng and h ave de mo nst r ated ,

that li v ing m atte r n eve r a rise s s po ntan eou sly fr o m lifeless m atte r ,

but o nl y th r ough the i mmediate influe nce of livi ng matter al r ead y

existing A nd so wh ateve r may h ave bee n the case at an e ar lie r
.
,
’
pe r iod of the ea rth s hi sto ry we are ju stified in r egardi ng th e
,

p r ese nt l ine between livi ng and l ifele ss as o ne of the m ost

clea r l y defi ned and im po r ta n t of natur al bou nd ar ies .

Th e Cont rast b et ween L ivi ngM att er and L i feless Matt er is made
the gr oun d fo r a divisio n of the natu r al scie nce s i nto two g r eat
gr oups v iz : the B i ologi cal Sc i ences and the Phy si cal Sciences
,
.
,

dealing r espectivel y with li vi ngmat te r and l ifeless matte r The .

biologi cal scie nces (p 7 ) a re kn ow n collectivel y as B i olog

.
y
(fl io
'

s lif ;
,
e M5 o a disc o u rse
) which is the r,
efore ofte n de
fi n ed as the scie nce of life o r of l ivi ng things o r of livin g m at
, ,

te r B ut liv ing matter so f ar as w e kn o w is o nl y o r di nary

.
, ,

matte r whi ch has e n te r ed in to a pec uliar s tate o r condi tio n .

 I N TR OD UC TOR Y .

pre i el defi ned as tile i

s c en ce which
he ce biolog y i s more c s y
A nd n

trea ts ( f m a tte r i n tb s li vi ng s ta te .

M att er A 1
Th e R ela ti ons h i p b etw e en L ivi n g a n d L i f el e ss .

m tter pre s e n t thi s re m a rkable

t hough livi ng ma
tte r a n d lifele ss a

other the a re m o s t i n ti m a t el y r el a ted as a ,

contr ast to o n e a n y ,
The livi n g ub sta nce of the h um an
mome t n s ref
’
lectio n w i l l show .
s

od or of i l or pl t is o l the tra n s
for m ed l i
fele ss
b y a n y an m a an n y
,
a nd has
,

o which h bee n take n in to the bod y

matter of the fo d as

for ti e the livi g tate L i fele ss m atte r in

there assumed a
,
m n
,
s .

the ha d p i g out agai waste

n as the o t he r on
d n
.

one n an ass
on
thro gh the orga n i sm s o m e of th i s m a tte r e n te r s
I n its jour ney u
into the living state and li nge rs for a time as p art of the body
soo e r or l ter it d i e a n d is the n fo r the m ost
s ub sta nce . B ut n a s ,

p art cast out of the body (though pa rt a m a

y be retai ed withi n n

either cc um ul tio n of w a ste m ate r i a l o r to s e r ve s o m e

it ,
as a n a a ,

ef l p rpo e M tte r m thu p f r o m the l ifeless i n to the

us u u s
) a.
a
y s a ss

livi ng st ate and back agai n to the l ifele ss ove r and o v er in never
'

e nding cycle s A li v i ngpl ant o r ani mal is like a fo un ta in o r a

.

fl ame into which and out of which matte r is co nsta ntl y st re am

, ,

m m i i
ing while the fount ai n o r the l e i tai t ch r acte ri t c
f a a n n s s a s
,

fo rm and in dividuality I t is noth ing but the co ns ta nt fo rm of

.

a s m ai il r t r
u m oil of m a teri al m olec u le s w hich ,a re co n s ta n tl y
flowi ng into the organi sm o n the o ne s ide and st r eam i ng o ut o n
the other . I t is a s o rt of focu s to which ce rtai n m a te r i a l p a r

t icles co verge i n w hich the m o ve for a ti m e a n d fro m w hich

n , y ,

they are afterward expelled in new co mbin ati o ns The p a r allel .

bet ween a whirlpo ol in a stre am and a li v i ng be ing w hi c h has ,

ofte n been drawn is as ju st as it is st ri ki ng The wh ir lpool is

,
.

permanent but the p article s of water whi c h co nstit ute it are in

,

c essantl ch angi ng Tho se which e nte r it o n the o ne s ide a re

y .

w hirled a round and te mpor a ril y constit ute a p art of its in di

v idualit ; and as the le a ve it o n the othe r side the ir place s a re
y y ,
”
m ade good by newcomers ( Huxley )
.

Ho w then is livi ng matter d iff ere nt f r om l ifeless m atte r

The question cannot be fully ans wered by che mical anal y sis fo r ,

th e reason th at this proce ss n ecessa ril kill s li v ing m a tte r and

y ,

the result s therefore te ach us little of the che mical c o ndi ti o ns e x

i stin
g in the m atter w he n al i ve A n.a l y s e s n eve rthele
,
ss b r i n
g ,
 L I VI N G MA TTE R . 3

to light seve ral highly im po rta n t fact s I t is likely that livi ng .

m atter is a tole rabl y defin ite co m pou nd of a n u m ber of the

Che mical ele me nt s an d it is p rob abl y too low an e sti mate to say
,

th at at l east six ele m e nt s m u st u n ite in orde r that life m ay ex

ist . Moreove r o nl y a very few out of all the ele m e nt s a re able
, ,

u nde r any circu mstance s to form thi s living part ne rship , .

The m ost sign ifica n t fact howeve r is that the re is no lo ss of , ,

w eight whe n livi ng matter is killed The total w eight of the .

lifele ss p r oduct s is ex actly equal to the weight of the livi ng sub

s ta nce anal z ed and if a n thi ng h as esc a ped a t death it is im
y y
‘

po nde rable and h av in g no weight is not m ate ri al I t follo ws

, , , .

that liv ing matte r co ntains no m ate ri al substa nce peculi a r to it

se lf and that eve ry ele me nt fou nd in living matter m ay be fo und
,

also u n de r othe r circu m s ta nce s in lifele ss matter

, ,
.

C o ns ide ratio ns like these lead us to re c o g ni z e a fu nd am e nt al

fact na mely that the ter ms li v i ng and l ifele ss desig nate two
, ,

diffe re nt S TA TE S o r C O N DI T I O N S of matte r “Te do not k no w at .

,

p rese nt what cau ses thi s differe nce of c o ndi tio n B ut so f ar as

,
.

the evide nce show s the li vi ng sta te is never as umed except

,
s

u nde r the i nfl ue nce of a ntecede nt liv ing mat t er w hich so to , ,

spe a k i nfects lifeless matte r and in so me way ca use s it to as

,

s um e the livi ng s tate .

Di stinct iv e Pr operti es of L i v i ngMatt er Those p rope rtie s of .

livi ng matte r which take n togethe r di sti nguish it ab solutely

, ,

f ro m eve r y fo rm of lifeless matte r a re ,

°

1 I ts che m ical co mpo s itio n

. .

2 I ts po w er of w as te a nd r epai r and of g r owth

.
,
.

3 I ts po w er of r ep r oductio n
. .

L i vi ng m atter i n va ri abl y co nt a i ns sub sta nce s k no w n as pro

teids which a re believed to co ns titute its esse nti al m ateri al b asi s
,

(88 8 p P roteid s are co m plex co m pou nd s of C a rbo n O xy

.
,

g en ,
H y d r oge n N it r oge n S ulphu r and (in so m e case s at an
, , y ,

rate) P hos pho r u s .

I t h as been f req uentl y po i nt ed o ut th a t each o f th ese s ix el ements is

rema rk ab l e in so me way o x ygen f or its v i go ro us co mb i ni ngpo wers ;
: ,

ni t rog en f or its c h emi ca l i ner t ia ; h ydro g

,
en f o r its g rea t mo l ec ul ar ,

mob il i t y ca r bo n sul p h ur a nd ph osph o r us f o r th eir a ll o t r o pi c proper t ies

, , , ,

etc . A ll o f t h ese pec uli ari t ies may be s h o wn to be o f s ig ni fi ca nce wh en

co nsi dered as a t tri b utes o f l ivi n

gma tt er ( See Herber t Spencer P ri nc ip les .
,

of B iolo gy, v ol. i

.
)
 OD UC TOR Y
INTR
‘
.

p r ese ce of p r oteid s w hich is

not, however the m e r e n
I t is ,
—
White of egg ( a lbu m e n )
-
co n tai ns

i absolut e l y l ifel ess

l p r oteid and y et s .

istic .

co m bu tio n but co n t in u a
of i nternal s ,
cha r acte r i stic k i n d di f
g r owth of livi ng thi ng s is of a ,
over the ,
g r owth of l ifele ss thi ngs
fro m the so called
.

f eringab sol utel y

-

grow if at all b ac cretion o r

lifeless bodie s y ,
C ryst al s and other
, ,

L ivi ng m att e r
w p rticle s to the out ide s
the additi on of
.

n e a
o r the tak i n g in of ne w
i e tion
-

I n withi n b n ta s s us c p
y ,

betwee n those
fitting them i nto the i te r tice n s s

the whole m ass A n d lastl y li v

alre dy
a p r e se n t throughout
,
. , ,

repai r its o w n was t e but al s o gives

ingmatter not o l y thu n s s ,

m of livi n g m atte r wh i ch
rep r d ctio n to new asse s ,
ri se by o u
f ro m the p re nt mass e n ter fo rthwith u po n
becomi ng detached a ,

an i n depe n de n t exi ste n ce .

i these p r ope r ties a r e b y

We may perceive how ext r a o r d na r y

e gi e to po se l ike powe rs : to carry

su ppo si ng a lo c o m otive n n s ss

proce ss of s elf r epai r in order to

co m pe n s ate fo r we a r ; to
on a
-

f r o m i t self at i n te r va l s
grow and i nc rease in siz e detach ing ,

pieces of b rass or iron e ndowed with the powe r of g o i g u p r w n

g the m
'

m o f r u n n in
s tep by s tep i nto other loco otives cap able

elve of reprod c i g w loco m otive s in i

the r tu r n P re
s a d u n n e
.

s n
,

the e thi g do e b eve r liv ing th i n g a n d no

thi n
g
i
e s y
e l s n s a r e n y y ,

like the m t ake s place in the lifeless w orld .

Huxl ey h as g i v en th e bes t st at ement ex t t an o f th e d

r
i t i t iv p p
s nc o e r t i es e

o f l iv i ngma tt er a s f o ll o ws
“
,

1 I ts ch emica l co mp os itio n co n a
.
t i ni ng as—it i n v a ri ab l,y doe s ,
o ne

o r mo e r f o r ms o f a co m pl ex c o m po und o f c a r bo n h yd r o g e n o
x yg e n
,
, ,
and

a ned e cept
ni t rogen th e so call ed pro t ein ( wh i c h h as never
,
-
b bt i yet x een o
f w
as a pro duc t o f l i v i ngbodi es) uni t ed wi th a l rg pr p r t i t r
a e o o o n o a e ,
,

i m
a nd f or mi ngth e c h i ef c o ns t i t uent bt hi h ri
w i t a ry
f a s u s a n c e
o c n s p ,

unmodi fi ed s t ate is k no wn s pr o t o pl as m
,
a .

r d t i n d it co n
2 I ts uni vers al d isi n tega tio n a nd wa s b i
t o x a o n a s
.
e y ,

co mi ta n t rei nteg ra tio n by th intuss p a s c e teio n Of n e w m a t t er A p ro c ess .

t
o f was e esur lt i f r
ng o m th e dec o mpo s i t i o n o f t h e m o l ec u l es o f t h e p ro to
 L I VI N G MA TTE R . 5

l as m in vir t ue o f wh ic h th ey b rea k up i nt o m o re h i g hl y ox idat ed produc t s

p ,

wh i c h cease to f o r m any pa r t o f th e l i v i ngbody is a co ns t a n t co nc o mi t a nt ,

o f l i fe Th ere is reas o n to bel i ev e th a t ca r bo ni c ac id is a l wa ys o ne o f th es e

.

was te prod uc ts wh i l e the o th ers co nta i n th e remai nder o f th e ca r bo n t he

, ,

n i t ro g en t h e h ydro g ,
en a nd t h e o th er e l ements wh i ch may ente r i nto th e
,

co mpos i t i o n o f t h e pro t opl a s m .

The new ma t ter ta k en in to ma k e g ood t h is cons t a nt l oss is e i th er a

ready fo r med pro to pl as mi c ma teri al s uppl i ed by so me o th er li v i ngbei ng

-

, ,

o r it co ns i s ts o f th e el e ments o f p ro t o pl as m uni t ed t og eth er i n s i m l er

p ,

c o mb i na t i o ns wh i c h co ns t a ntl y h a v e to be b ui lt up i nt o pro t o pl as m by t h e
,

ag ency o f th e li v i ngma tt e r i t self I n ei th er cas e t h e addi t i o n o f mol ec ul es

.
,

to th ose wh ic h a l read y e x i s ted ta k es place no t a t th e s ur fac e o f th e l i v i ng ,

mass b ut by i nte r po s i t i o n be t wee n th e exi s t i ngmo l ec ul es o f t h e l a tt er

, If .

the proc ess es o f di s i nt eg ra t io n a nd o f rec ons t r uc t io n wh ic h c h a ra c t eriz e

l if e ba l a nce o ne a no th er t he s ize o f t he mass o f l i v i ngma t t er remai ns s ta
,

t io nary wh i l e if t h e r ec o ns t r uc t ive proc ess is t h e mo re r a pi d t h e l i v i ng

, ,

body g ro ws B ut t he i nc reas e o f s i ze wh i c h co ns t i t ut es g
. ro wt h is the
res ul t o f a process o f mo l ec ul a r i nt uss uscept i o n a nd th eref o re di ffers a lt o ,

g et h e r f r o m t h e proc ess o f g ro wth by acc ret io n wh i c h ma y be O bserved in ,

c rys t a l s a nd is eff ec t ed purel y by t h e ex t er na l addi t i o n o f new ma tt er

, so

t h a t in the well k no wn aph o ris m o f L i nnaeus the wo rd gro w as appl ied

,
-

to s to nes s ig ni fi es a t o t a ll y di ff ere nt process f ro m wh a t is ca ll ed g ro wth

in pl a nt s a nd a ni mal s .

3 I ts tend ency to underg

. o cy clic a l ch a ng es I n the o rdi nar y co urse .

o f na t ure all l i v i ngma tt er proc eeds f r o m pre e x i s t i ngl i v i ng ma tt e r

, a ,

po r t io n o f the l a tt er bei ngdet a c h ed a nd ac q uiri nga n i ndepe ndent e x is t

ence . Th e new fo r m t a k es o n t he c h ar ac t e rs o f th a t f ro m wh i c h it a ro se ;
e x h i b i ts t h e s a me po we r o f pro pag a t i ngi t sel f b y mea ns o f a n o ff s h oo t

a nd soo ner o r l a t er l i k e its pred ec ess o r ceases to l i v e a nd is r eso l ved

, , , ,

i nt o mo re h ighl y o x i da ted co mpo unds o f its el ements .

“
Th us a n i ndi v i dua l l ivi ngbo dy is not o nl y co nst an tl y c h ang i ngits
s ubs ta nce b ut its s iz e a nd fo r m are underg
,
oi ngco nt i nua l modi fi c a ti o ns ,

th e end o f wh i c h is t he dea th a nd dec a y o f th a t i ndivid ua l th e eo nt inua

t io n o f the k i nd bei ngsec ured by th e det ach ment o f po rt i o ns wh i c h t end
to run th ro ug h t h e s a me c ycl e o f fo r ms as th e pa rent N o fo r ms of ma tt er .

w h i c h a re ei th er no t l i v i ngor h a ve no t bee n deri v ed f ro m l i v i ngma tt e r

ex h i b i t th ese th ree pr o pe r t i es no r a ny a ppro ac h to th e rema r k a b l e phe
,

no mena defi ned under t he seco nd a nd th ird h ea ds ( E ncyc lop e dia B ri .

ta nn ica 9 th ed a rt ,
B io log y vo l iii p
.
, .
, . . .

Fo r th e pur poses of biological study life mu st be r ega r ded as

a p r ope rt y of a ce rtai n ki nd of com pou nded matte r B ut we .

are fo r c ed to r ega r d the p r o pe r tie s of co mpou nds as the result

a nts of the p r ope r ties the ir co nstitue n t ele men ts even though ,

we ca nn ot we ll imag ine h o w such a r el atio n exi sts ; and so in the

‘
 IN TR OD UC TOR Y .

n the prope r tie s of c ar bo n ,

fo r the prop e r tie s of liv ing

matter follo w
It fro m
og
0

Sci ences
.

s
B i l i cal
f B i l og T h e o .

S pco e o o y .

give to B iolog y
that thi s s cie nce i ncl ud es
the broad defi n itio n n
to livi g m tte r or to liv ing
wh tever er t a iu s n a
the st dy u of a p
the for m t r ucture s a n d f u n c tio n s of livi ng
things I t co n s ider s s s
, ,

thei r h bit actio s m o des of

.

s n
thi ngs in health an d in di s e ase ; a , ,

and
di trib u tio n i n s p a c e a n d t im e
nutrition ; their
sur r ou n di n g s s ,

world a d
n to o n e a n other the i r
their r el a tio n s to the lifeless ,

relatio their o r igi n and

p r oce e n d soci l a n s
sensations ment al ss s a
,
,

g d
,
othe r topic I t i ncl u de s both z o iil o y a n
their fate and ,
m a n y
s .

de l with the phe n o m e n a of ani mal an d vegetal lif e

bot any and a s ,
I
ep r tel b u t i n thei r r elatio n s to o n e a nothe r t .

n ot onl y s a a y ,

i ncl udes the medical scie e n c s a n d vegetal patholog y .

The field covered by biology as th us u nder st o od is so w ide as

to nece ssitate a subdivi sion of the subject i to u b r of p r i cip l n a n m e n a

branches w hich ar e u su a ll y ass ig n ed the r a n k of di s ti n c t s cie n c e s .

These are arranged in a tabul ar view o n p 7 .

The tabl e ho
.
s w s

of reg rdi n g the m a in s ubject a cc o rding as

i
two d ffere t n w a
y s a ,

the table is r ead from left to right o r n i ce T e rs a U de r the

n .
m o
re
us u a l arr an ge m e n t biolo g y is pr i m aril y divided i n to z o ii lo
g y a nd

a i l or pla ts r e pectivel for m t h

botany accordi ng as n ma s
,
n ,
s y e
,

su
bject of t
s u y d . S uch a divi s io n h as the g r eat adv an tage of

co ve ie ce ce m atte r of fa t m S t bi log is ts
pr actical n n n sin as
,
a c o ,
o

devote their attention mai nl y either to pl ant s alo ne o r to ani mal s

lo e Fro a scie tific poi t of view h weve r a better s ub
a n . m n n ,
o ,

division is i nto M orp holog y

A d j O s a d i
/ s co u
,
rse
)
and P hys iolog 19 n a ture; A o
'

y o s a d i s co u r s e
) The
y
.

, ,

for mer is b ased upon the facts of fo rm st r uctu re and ar r an ge , ,

e t
m n and is ss n a y s
,
e e ti ll tatic l
a ; the latt e r upo n thos e of a c tio n

or function and is essentially dynamica l B ut mo r phology a nd

,
.

physiolog are so i nti m atel y related that it is impossible to se pa

y
r ate either subject ab sol utel y fro m the othe r .

B e side s the sub scie nce s given in the table a disti nct b r an ch
-

c alled E tiolog is ofte n recogni z ed h avi ng fo r its obj ect the ih

y ,

v esti ation of the c auses of biolog

g i c al phe no m e n a B ut the sci
.

e tifi c stud of ever phe o m e o n h as for its ul t im ate o bject th e

n
n n
y y
d i scover y of its cause E tiology is therefo re i nse parable f ro m
.
 THE B I OL O GI CA L S CI E N CE S .

A na tom
y .

Th e i s c encet u of s r c
tur ; th t rm b i ng
e e e e
u u l l y pp l i ed t t h
s a a o e
coa rse r nd m r h a o e O
vi n mpo i ti n f
o s co s o o
p l nts
a a ni m l or a s .

ir e os c o p i e a na o t my
lti m t p t i l
.

Th e u a e o ca
a na ly i f tru t ur
s s o s c e
b y t h e a id o f t h e
m i c r o s c o p e ; se pa
rt
a ed f
ro m a na o my t
l
o n y as a ma e of tt r
co n v ni e e nce .

ca t io n .

Th e i fi ca t i n f
l
c ass o o
i l vi ngth i ng B ased s .

c hi fl y n ph n m na
e o e o e
o f t r u t ur
s c e .

Dis tri bution .

C o n iders
s i ti th e ou
f l ivi g
o
g n t h n s in
s nd t i m th i r a e,
E ssen t i all y d t i b ut i n v r t h
s r o o e
e
e
pr nt face
ese f th o e
ea rth nd th i r di t i
a e s r
b uti n nd u o i n a s c c ess o
t f rm r p ri d
a o as e e o s,
d i l y ed in f
s ail oss re
us .

Th e vl
sc i enc e o f
de e o p
men t fro m t he r
ge m
l ix
.

I nc udes ma ny m ed
ro b le ms rt i i
pe a n ng
r h l
t h t o mo p o o
gy
h i l
a nd p y s o o
gy A t .

r t lr l
p ese n a g e y mo r
p ho log ic a l .

P g
si o lo
k
gy .

Th e l i nce f
s pec i a sc e o
th f un t n f th
e c io s o e
i nd vi du l n h lth
i a i ea
a nd in d i ; h nce s eas e e
inc l udi ngI h th l
gy o o .

B ych olog
y .

The i
sc enc e o f menta l
p h eno mena .

Th e sc e ncei Of s oc a i l
l if e, i l if f
. s .
, th e e o
co mmuni t i wh th es , e
er o f m n r o f l o wer
e o
 I N TR OD UCTOR Y .

8
of biolog and n eed n ot be assign ed
ever l b r anche s y
any of the s a

an m( 16 )
e nde n t Pl ac e
ad m itted t
1
.

not yet ge n e rall y

o

c to m a r n d co n
and it is
a
u s y
The s e t a bli s h m e n t o i t h e

how n however that the s t u d y

clearl y s , ,

that o f biology in the ord i

able fro m
actio of the lo w e r
a n d other m e n tal n s
n ary sense ph s i o logical
.

bject s of p s
y ch o logic l a as of y
ani mal s a re a s trul y s u
co m m u t i e s as
ocial life of s u ch a n i m a l n i .

i nquir y ; the co m plex s

amo ng
the bee s a n d a n ts a re n o le ss trul y
we fi nd for i,
n st an ce ,

problems of Soc iology .

o ph log d ph
that i the sche m e m r o y a n y si
I t will be ob served
n
biologi al cie n ce s in
overl p th at is the r e a r e ce r tai n c s
ology a ; ,
t be s ep a r a ted
hich the tud of str ucture a n d of a c t io n c a n n o .

w s y co ide r the u
tr u e of e m b r
y o l o
g whi ch n s s s c
Thi s is e peci ll y
s a

The term G e n er al B i ol og y d o e s n o t de s ig n a te a p a r ticula r

e ber of the gr p of biologi c al scie n c
e s b u t is o n l y a con
m m o u ,

phr e hich co m e i to fo r the ge n e r a l in t ro d u c

v enient a s w,
h as n use

tor study of biology I t be a rs preci sel y the s a m e relat o i n to

y
.

m i ge n e r al
bi ology th at general chemi str y be ars to che try o r s

ph si cs bears to ph ysic s I t i ncl ude s an ex i tio of the g a m n a n e n

y
.

er al properties of livi ng matter as r eve aled i n the truct r

s u es a n d
ctio p rtic l r li v i g thi g an d m a s e rv e a s a b as is f or
a n s o f a u a n n s , y
b eq e t t d of m ore s peci a l br anche s of the scie n ce I t
u
.

su s u n s y
deal s with the broad ch aracte ri stic phe no me na and l aw s of life as
of
ill ustr ated by the thorough co mp arative t dy of a se ie s u r s

pl ant s and animal s t aken as repre se nta tive ty pe ; s b u t i n th is

t
s u y d the t
s u de t
n s ho u ld n ever lo se s ight of the fact th a t all the

v aried phenome na which may co me u nde r hi s ob ervatio a re

s n i n

the l ast anal ysi s due to the p rop ert ie s f

o m a t t er i n t h e l m
i in g
s ta te and th at thi s matter and these prope rtie
,
s ar e the real g oa l

of the study .
 C HA P T E R II .

THE S TR UC TUR E O F L I VI N G THI N G S . O R G A N I SMS .

LI th ings occu r in m asses of the m o st va r iou s si z es

'

FE L E S S

a nd fo r ms and m ay dif fe r widel y in s t r uctu r e and che mic al c o m

,

po sition L iving things o n the othe r han d occu r o nly in r ela

.
, ,

t iv ely s m all m asses of which pe r haps the large st are am o ng , ,

plan ts the g r ea t trees of C alifo r nia and a mo ng a ni mals the

, , ,

w hale s ; while the s malle st are the m ic r o o r ga n i sms o r bac teri a -

.

M o r eove r the i ndi vidual mas se s in which livi ng thi ngs oc c u r

,

posse ss a peculiar and charac teri stic structu r e and che mical c o m
po sitio n which have cau sed them to be k now n as o rg a n is ms a nd ,

the ir sub stan ce as org a n ic A ll organi sms are built up to a .

re ma r kable exte nt in the sa m e w ay and o f the sa m e m ate r ials ,

FI G . f t r S h ) —L ngi t ud i l t i n th ugh th gr w i g p x f y ung

1 . (A e ac s . o na s ec o ro e o n a e o a o

p in h t Th d tt d p r t i
e~s oo r pr nt t h p r t p l m t h rr w l i n b
. e o e o on e es e s e o o as , e na o es e\
in gt h p t i t i w ll m p d f l lu l ( C HW O Q ( Hi ghl y m gn i fi d )
e ar on -
a s co ose o ce os e O . a e .

a nd w e may
con ve nie ntl y begi n a stud y of livi ng thi ngs with the
large r and more c o mplex for ms w hich exhibit m o st clea r l y ,

t ho se structu ral peculiari tie s to w hich we have r efer r ed .

Or ga ni sms composed of Or g a ns Fu ncti ons I t is cha r acte r . .

i stic of any livi ng bod y— f o r exa mple a rabbit o r a ge r aniu m ,

that it is co mpo sed of u nl ike p a rts havi ng a s t r uct ure which ,

e n able s th e m to pe r fo r m v ar iou s operatio n s e ss e ntial o r acce ss o r

O y
t o the life of the w hole The pla nt has ste m r oot s branche s .
, , ,

leave s s ta m e ns pistil seed s etc ; the a ni mal h as exte r nall y

, , , ,
.

9
 C OF LI VING THI N GS
10 THE S TR U TUR E
.

e e etc and i nte r nall y stomach in

head t r unk li mb s y s c ars
, , , ,
.
,
,
,
he rt br i n and many othe r pa r ts of
testin es live r l g
un s a a ,
, ,
, ,

FI G 2 —Cross s ec t i o n thro ug
. .
- h pa r t o f t h e y o un g l ea f o f a fer n ( Pterts amwaaiw) ’
n
,

s h o wi n g th i c k w a ll ed c e ll s ; m o s t o f t h e w a ll s a re d o ub e ra n ula r s ub
-
l Th e g .

s t a n c e is p r o to p l a s m M os t o f t h e c e ll s c o n ta i n a l a r ge c en t ra l ca v i t y ( v ac uo l e )
.

fi ll ed w i th s a p th e pro to p l a s m h a vi n gbeen reduc ed t o a th i n la y er i ns i d e th e

,

p a rt i t i o n s N .u c l e i ar e s h o w n i n s o m e o f t h e c e ll s a n d l ife l e ss g r a in s o f s ta rc h
,

in o th re s : i t, n uc e l i; t
8 , s arc h; v, v l
ac uo e ; w do u
, bl e pa r ti t i on -
wa l l . x

the m ostdive r se str uc ture The se p art s are k now n as org a ns ,

.

and the livi ng body bec ause it po sse ss e s the m is call ed an or

, ,

g a n ism .

Th e wo d rgani sm as h ere used appl ies bes t to th e h igh er a ni mal s

r o , ,
.

and pl a nt s I t wi ll be seen in th e seq uel th a t th ere a re fo r ms o f l i fe so

.

s i mpl e th a t org ans as h ere defi ned ca n sca rcel y be di s t i ng ui s h ed S uc h .

l i v i ng th i ng
s are nev er th el ess rea ll y o rga ni s ms because th ey possess
parts analog ous in f unct i on to th e well de fi ned o rg a ns o f h i g h er f o rm -
.

( See p .

organi sms are compo sed of u nlike pa r ts the y are sai d

S ince ,

to be heterogeneous in st r uct ure They are al s o hete r oge neous .

in ac tion the diff ere nt orga ns perfo r mi ng d iffe r e nt oper ati o ns

,

c alled f un ctions For i nst ance it is the function of the s to mac h

.
,

to dige st food of the he art to p um p the blood into the vessel s

, ,

of the kidneys to excrete w aste matter s fro m the blood and ,

of the brain to direct the func tions of o the r orga ns A s i m il ar .

di v er sity of functions e xi sts in pl ant s The r oots hold th e .

 OR G A N S A N D TI S S UE S . 11

pl ant fas t and absorb v a riou s sub stance s f r o m s oil ; the ste m

s uppo rt s the le a ve s an d flower s

a n d co n ducts the sap ; the le a ve s

a b s orb and elabor ate po rtio n s of

the food ; and the r ep roductive

orga ns of the flowe r serve to
form and bri ng to m at urity seed s
de st ined to g ive r ise to a n ew g en

c r atio n .

Heteroge neity of the ki nd

j us t i ndicated acco mpan ied by a
,

divi s ion of la bo r a m o n g the

parts is o ne of the m o s t char
g g
gg
,
1 3

13 0 ; g fi zf f af ffifli
3 A s e ’s r s ‘ "
ac te ristic fea t u r e s of liv ing thi n gs , 1 ro e ‘

an d is not k no w n in a n
y m ass of w l d w
l d ll fr m th t m f a e oo -
ce s o e s e o

fly w
lifele ss matter howeve r la r ge and $321 2 fg3 32225ifi23
.

3 ’i or

, .

co m plex .

Or ga ns composed of Ti ss u es Differ enti ati on I n the n ext

. .

place it is to be observed th at the orga ns also whe n fully

, ,

for med a re no t h o m oge neou s but are in t ur n made up of

, ,

dif fere n t parts The hu m an h and is an o r ga n which co nsi st s

.

of m any p ar ts d iffe ring widel y in structu r e and functio n O n

, .

the outside are the ski n th e hai rs the nail s ; i nside are bo ne s
, , ,

muscle s te ndons lig

,
am e nt s blo o d ve ssel s and nerve s The le af
, ,
-

, .

of a pl an t is an orga n co nsi sti ng of a w ood y fram ework ( the

”
ve ins ) w hich supports a gree n pulp the w hole bei ng covered ,

o n the outs ide by a deli c ate t r ansp a r e n t s ki n I n like m ann er .

eve ry orga n of th e higher pl ants or ani ma l s may be r esolved

i nto diffe r e nt p art s and the s e a r e k n ow n as tiss ues The
,
.

t iss ue s of full y for med o rgans are ofte n very differe n t fro m o ne
an othe r as in the cas e s j us t m e nti o ned ; th at is they are w ell
, ,

dif ere ntia ted, but freque ntl y in adult organs and al w ays in tho se
’

whi c h are sufficie n tl y y ou ng th e ti ssue s sh ade graduall y i nto

,

o ne another so th a t n o defi ni te li ne ca n be dr a w n bet w ee n the m

,
.

I n s uch case s the y are said to be le ss differe nti ated For ex .

a m ple ih the full grow n le a f of a pla nt the w oo d fr am e w ork the

y
-

, ,

gree n cell s and the ski n ex ist as th ree plai nl y diffe re n t ti ssue s
,
.

B ut in y ou nger le a ve s the se sa m e ti ss ue s a re le ss differe nt an d ,

in ver y y o ung leave s s till in the bud there are no vi sible di ffer
, ,
 THE S TR UC T UR E OF L I VI N G THI N G S .

12

hole o rgan is ve ry n ea rl y ho m oge n eou s I n this .

eren tia d though pot e ntiall y capable

c ase the tissue s are u n d i
f te ,

the sa e
m w y the ti ssue s of the e m b ry
In a ,

FI G . 4 .
—
Cr o s s s ec
- ti on thr ou gh dea d w oo d -
l ikll fr m th und r gr und t m f
e ce s o e e o s e o a

r
f e n ( Ptems
‘

i
aq u il na ) . ll
Th e w a s a re un mm
co l y th i k d th pr t p l m h
on c an e o o as as

di sa pp ea r ed . Th e c h l h wn
a nn e s s o se rv d i l if t k p t h ll in vi tal n
e n e o ee e ce s co

n ec t io n . (x

onic human h and are i mperfectl y differe ntiated and at a v e ry ,

e arl y stage are undiffere ntiated .

Ti ssues composed of Cells Fi nall y mi c roscopica l ex amina .

,

tion shows every ti ssue to be co mpo sed of m i nute pa rts k no w n

as cells which are nearl or q uite i m il r to n n othe r th rou gh
, y s a o e a

o ut the wh o le ti ss ue and for m the ulti mate units i nto which the
,

ti ssue s and organs and he nce the whole organi sm be c o me mo r e

, ,

or less perfectl y divided so me wh at as a natio n is divided i nto ,

states and these into countie s and to wn ships .

 CE L L S . 13

It w ill be sh own bey on d that these ulti mate u nits o r cells

possess eve ry whe r e the sa m e fu ndam e ntal str uct ure ; but they
diffe r i mm e n sel y in fo rm s iz e an d m ode of actio n not o nly in
, , ,

diffe re n t animal s and pla nts but eve n in difi erent p arts of the ,

sa me i ndividu al A s a r ule the cell s of any give n tissue a r e

.
,

c losel y sim ilar o ne to a n other and a r e devoted to the sa m e fu nc

tio n but d iffe r fr om those of othe r tissues in for m si z e a rr ange

, , ,

m e nt an d e s peciall y in fu nctio n
,
I n deed the dif fere nce s be .
,

t w ee n ti ss ues a re m erel y the outc om e of the diffe r e nces betwee n

the cell s compo si ng the m The ski n of the ha nd diff e r s in ap
.

eara nce and uses f r o m the m uscle which it cove rs becau se ski n
p ,

cells diffe r fro m mu scle cell s in fo rm s i z e color functio n etc

-

, , , , .

H e nce a t issu e may be de fi ned as a g ro u

p fo s im ila r ce lls h as

i ng a si mila r f u n c tion *
A s a rule each orga n co n si st s of
.
,

seve ral s uch g r ou p s of cell s o r ti ss ue s but as s t ated above o un g

y , , ,

or gan s a r e nea rl y or q uite homoge neou s ; that is all of the cell s ,

a re nea r ly o f qu ite alike I t is o nl y whe n the organ gro w s .

olde r that the cell s beco m e differe nt and ar r ange the mselve s in
diffe r e n t g r oup s — a p r oce ss k no wn as the dif eren tia tio n of the
’

I n the case of so m e orga ns —for i nst a nce the l e a f o f a

,

tiss ues
m oss —
.

the cells re mai n pe rma ne ntly n e a rly alike so me wh at as ,

in the e m b ry o nic co nditio n a nd the whole orga n co nsi s ts of a ,

si ngle tissue .

Wh at has bee n said thu s f ar appl ie s o nl y to highe r plant s

a nd ani mals B ut it is an in teresti ng and sugge s tive fact th at
.

the re a r e also i nnu m er a ble i sol ated cell s both veget al and ,

a ni mal w hich a r e able to car ry o n an i ndepe nde nt exi ste nce as

,

o ne celled
-
pla nt s o r a ni mal s P h y s iologica lly the se m ust c er .

tainly be rega rded as in di v idu al s ; but it is n o le ss cert ai n th at

the y a r e equivale nt mo rphologicall y to the co nstitue nt cell s of
, ,

o rd ina ry m a ny celled orga ni sm s I t will appear hereafter th at

-
.

the study of such u nicellula r o r ga ni sms for ms the logic al gr ou n d

wo r k of all biological scie n ce ( S ee p . .

S i nce o r ga ni s ms m ay be resolved succe ssivel y i nto organs

ti ssues and cell s it is evide nt th at cell s mu st co ntai n livi ng
,

matter A nd a cell may be defi ned as a s ma ll mas s of liv in g

.

ma tter either livi ngap a rt or f orm in gon e o f th e u lti m a te u n i ts

Ti u ss f req uentl y n t i n m tt r d p i t d b t w n ll b ut th
es co a a e s e os e e ee ce s es e

h a v e u u ll y b n di r tl y d ri v d f r m th ll nd v ry s th cell v ry
s a ee ec e e o e ce s, a a a e s a .
 OF L I VI N G THI N G S
14 THE S TR U TUR E C .

o thefi
m The cell is an org
an i c i n div id ua l f st

f
o a n or a n is
g .

0rder
”
( L a ng ) m S ce
i v i ngO g
.
.

l M att r i n t h e L r a n i s i n o u r .

L ivi ng a nd L i f e e ss e
m
f lower a n i m a l s an d o f pl a n t s a r e co

f r o m what h as bee n a id
osed of m a tter it m a
y be su ppo sed
s ,
h g
.

p th t
,
the are c o m po s ed of v1 n
in the l a st ch apter a y ,
true l i n p a t I t is
Thi h o w ever is o n y r hv m
.

matter s ,

g
. , .

tr e th t ever pl ant o r i m l co n t n
a n i ns
an a i s
stri c tl y u a y
little refle c tio n w ill s how th a t i t co ta
matter b u t a
ess m at
,
hf l
.

l o I the hu m a n bo d y e
lifele ss m a tter a s n .

f n an d
ter is found in the h ai rs the e d n s o the a il s ,

—
,

l er of the ski m s tru c t u re s which are

the outer y a s

devoid of feeli ng ever o n e k n o w s t he m

not simpl y
as y ,

re ll lifele i eve r se n se although

to be b u t a r e a
y ss n y ,

p r t of livi g bod N o r i s lifel e ss m a t

formi g n a a n y .

to the exte r ior of the b od

y The m i n e r al
ter c onfi ned .

tter of the b e s is no t alive ; a nd thi s is t rue

ma o n ,

though ss le obvio us y l of m a n
y othe r p a rt s
,
such as
,

the liquid b asi s or pl asma of the blood the f at ( which ,

ever wholl b e t d vari ther f r m of m t

is n y a s n ) a n o u s o o s
,
a

ter oc curri ng in many part s of the bod y .

In lo w er a n i m a l s exa m ple s of thi s t ru th occu r o n

ever y h and The c al ca reo us sh e ll s of a nim al s like the.

snail and the o y ster ; the s keleto ns of

co r al s and s po nge s ; the h a rd o ute r c rust

of i nsects lobs ter s and rel ated a ni m al s ; , ,

the scale s of fi sh and reptile s ; the

feather s cl aws and be ak s of bird s ; th e , ,

f ur of ani mal s the se a re a few of th e -

countless i nsta nc e s o f st ruct ures co m

p osed wh o ll y o r in p a rt of l ifele ss mat
F I G5 ( A ft r R
. . vi r ) — Mu
e
ter
an
W hi C h n eve rthele ss e nter I nto the
e . s

l ll
o e ce -
A f r m ths . i t ,
o e n es

ti ne of d g i r a o compo sitio n of livi ng ani mal s

, n c o ss s ec
- .

t i ; B i gl i l t d ll
g
on s n ce

f m th i t t i
ro
,

e f b
n es
e
A
so a
nl o n
ne o
pl e
an ts
a ra
hl e
,
f ac tS a re ev en ‘

b it vi w d f r m th
, e ei d more c o nspic uo us
o e s N o o ne ca n d o ubt
e. .

x 320) .

th at the outer b ark of an oak is devoid

of life The heart wood o f a tree is e nti rel y de ad and e v e n
.
-

in the so called live w ood thro ugh whi c h the sa p flo w s no t o nl y

-

, ,

is the solid p art of the wood lifeless but al so the sap its elf ,
.
 L I FE L E S S MA TTE R B E T WE E N CE L L S . 15

FI G . 0 (A f t r S c haf r )—
e e H um
. an c a rt il a ge f ( ro m h ea d o f met a ta rsa l b
o ne) . 0, ce ls l
m , l if l
e ess ma trix . ( x 6m) .

( M o di fi ed vi r ) —
f ro m R a n e B l d f fr g h w in gt w f r m
. oo o f o , s o o o s o ce ll s ( c or
p usc le s ) , o ne tt
fl a en ed a nd v l n b r n h d fl t i ngin th l if l
o a , o e a c e , oa e e es s p la sma .

(x
 c r owded u n it s o r cell sep a r ated b y ve r y s
n ear l y si milar
clo sel y g
m ra nula r
,

E ach ce ll co ns ists of a ass of ,

narrow p ar titio n s
m l
a o e so id
.

k o w as r o top la s m an d r ,
v i scid livi ng sub sta n ce n n p ,
,

the u cleu s
rounded bod y
n .

rather the whole m ass 0

,
co po ed of h v mgmatte r ( p r otoplas m)
m
.

w hich is al mo st e nti r el y
s

ho i Fig 2 I n eve r y c ase th e p

m p o
s ed o fa rti t io ns a re co
s w n n
d b
. .

d d d t y
lif e less m a tter w h i c h h a s b e en m a n u f ac tu r e a n ep os i e

i
od e f h ll I n
the liv ingp rot p o la s m co n s tit u t in g th e b s o t e ce s .

ce r tai n of the li feless wa lls m ay

s till older p art s of the pla t
n

p r ot o plas m e nti r el disappea r s an d

beco me extremel y thick ,
the y ,

the whol e tissue ( wood ) co nsists of

lifele ss matte r e nclosi ng s pac es fill ed
with air o r water ( Figs 3 and .

A mo ng a ni m al s a nal o gou s cases

are co mmo n The mu scles of the .

s mall i nte sti ne fo r i nsta nce ( Fig , ,

.

consi st of bu ndle s of elo ngated

cell s ( fibres) e ach of w hich is co m
posed of livi ng matte r su rr o unded
by a ve ry thi n coveri ng ( s hea t h) of
lifele ss m atte r I n cartilage o r .

gri stle w hich cove r s the e nd s of ,

many bo n e s ( Fig the ov al c ells .

a r e v er y w idel y sep a r ated by th e

depo sitio n betwee n the m of large
quantitie s of solid lifeless m atte r
FI O 8 ( M di fi d f r m S h
. . o k ) —S for m i ng wh at is k no w n as the
e o c en . ec
t i f b f rom th h um f m ur
on o one e an e

s h w i gth livi gb r h i gb
o n e 7 na tr i1
n I ll blood ( Fig 7 ) the
a nc n o ne ’ '
.

ll l y i g i th b y l i f l m
,

ce s n n e on e es s a
fl tte ed or rreg ul r ce lls ( co r
trix Di g m t i
. a ra a
a c
n ,
i a

p as cles
) are s ep ara ted b y a lifel ess

fl ui d ( p las ma) i n which they fl o at I n bo ne ( Fig 8 ) the cells . .

 L I FE L E S S MA TTE R WI THI N CE L L S . 17

have a b r a nchi ng i rr egula r fo r m and are sepa r ated by soli d

, ,

cal car eo us m atte r which is u n mi stak abl y l ifele ss The se ex .

a mples sho w that the lifele ss matter s of the body ofte n occu r in
the fo r m of depo sits bet w ee n livi ng cells by which they hav e
bee n p r oduced I n all such cases the e mb ry o nic ti ssue co nsi st s
.

at fi rst of liv ing cells in di rect co nta ct o r sep arated by o nl y a ,

v e ry sm all q ua ntity of lifeless m atte r I n la te r stage s th e .

cells may manu factu r e addi tio nal li fele ss sub stance whic h
appear s i n th e fo r m of fi rm p a titio n walls bet w ee n the cell s
r
or as a mat ri x solid o r li q uid i n w hich the cell s lie Whe n
,

, , .

soli d wa ll s a r e p r ese nt the

y a r e ofte n pe r fo r ated by na r row ch an
n el s th r ough whi ch the p r otoplas mic cell bodies r e mai n in c on -

nec tio n ( S ee Figs 4. 8 and .

, ,

L if eless Matt er wit h i n L ivi ngCells E q u ally i mpo r ta nt with .

the depo sit of li fele ss m atte r between cell s is the fo r matio n of life
less matte r within cell s eithe r (a ) by the depo sitio n of various sub ,

s ta nce s in th e p r otopla s m o r b b
( ) y the direct t r a nsfo r matio n o f ,

the whole mas s of p r otoplas m E xamples of the fi r st ki nd are .

Fm 9 . .
—A gro u p o f c ll fro m th e s t e m
e s of a gera n i um FI G . 10 . (A ft r e R a n v ie r) .

( P elammuu m) s h w i gl if l ub t n
, ( t r h
o n e es s s s a c es s a c G r up fo ip o
“
ad o se ce ll s

t i th i th p t p l m A i Fig 2
a n d c ry s a ls ) w n e ro o as . s n .
, fr m th t i u b
o e ss e en e a th t h e

ea c h ll t i ce l r g n t r l v u l fi ll d
co n a ns a a e ce a ac o e, e sk i ( ub utn
“
u s c a n eo s con

wi th p; 0 gr up f ry t l f l i u m x l t ;
sa , o s o c s a s o ca c o a a e n ti
ec ti uve f n m ss e o a e

i. c i nt r ll ul p ; nu l u ; gr n ul f
.
, e ce ar s ac e i t, c e s 8, a es o b y r l f h w i gd p f
o ca , s o n ro s o

s ta r h ( x c . f t in t h p r t p l m f f t
a e o o as . , a

dr p o ( bl k )
s n nu l i ac , c e

(X

cry stals (Fig

mi n e ral grai ns of st arch (Fig drops of . .

w ate r and many other sub s ta n ce s fo un d wi thi n the cell s of

,

plants A mo ng a nimals drops of fat (Fig 1 0) and calca r eou s

. .
 T UR E OF L I VING THI N GS
H S TR UC
.

18 T E

depo it e s i m il arl y produced I n deed the r e is ,

or siliceous a r
.
s s

li m it to the n u m ber of li feless s u b s ta n ce s whi ch

s carcel y a yn
ithi the cell both of pla n ts and a ni mal s
ppe r w n s .

may th u s a a

c e i of le ss i m porta n ce tho u gh of co mm o n
The seco d n a s s ,

good ex m ple is fo un d in the li n i n g m e m bra ne

o ccur r ence A .
a
whi h like the hu m a n
of the e oph g
o s a u s of the dog ( Fig c .

e ti el m de up of lo el c r owded cel ls Tho se

m
skin is al ost n y a c s y .
r

11 — li vin g
FI G S ti ec on th
ro ugh th e i nn e r c oa t of th e g ll t
u e of a dog. sh o w i n g:
ll rl y r li f l ll rfi ia l l
. .

e e ss c e s of th e s upe c a y e rs ; n , n uc le us
d eep e
.

ce s of th e a e s ; a.

in the deepe t p r t co ist chiefl o f livi ng p r otoplas m ve ry

s a n s y
i
s m a il r to th t of the ou ng pi ne s hoot ( co m p ar e Fig
a y
.

A bove the m the cell s graduall y beco m e flatte n ed u n til a t the

rf ce the have the fo r m of fl a t scales A s the ce ll s beco m e
su a y
.

fl attened their sub stance cha nge s The protopl as m di mi nish es .

in q uantity and die s ; so th at n ea r the su r face the cell s are

w holl de ad a nd fi na ll y fall o ff I n a s i m il a r m a nn er are
y .

for med the lifele ss p ar t s of nails claw s be ak s feathers and , , , ,

m any rel ated str uctu re s A h ai r is co mpo sed of cells esse ntiall y
.

like those of the ski n A t the r oot of the hai r they are alive
. ,

but as they are pushed outw ar d s b y co nti nu ed g r owt h at the

root they are transfor med bodily i nto a dead hor ny sub stance
, ,

for mi ng the free portio n of the hai r Feathe r s are o nl y a co m .

plic ated ki nd of h air and are for med in the sa m e w ay .

I t is a s ignific ant fact th at the quantity of lifeless matte r in

the organism te nd s to increase with age The ve ry y oun g plant .

or ani mal probably possesse s a maximu m pro po rtio n of proto

pl sm and as life progre ss es lifele ss matte r gr ad uall y acc umulates
a ,

W i thi n or about it — someti mes fo r s upport as in t r ee tru nks and -

, ,
 THE C
S TR U T UR E OF L I VING THI N G S . 19

bo ny skele to ns ; so m etim e s for prote c ti on as in oy ster an d snail .

s hell s ; so m eti me s app a re ntl y fro m sheer i nabilit o n the p a rt of

y
the p rotopl asm to g et xrid of it Th us we see that y outh is lit
.

e rall
y the pe r iod of life g
an d vigor an d a e the pe riod of c o m
,

a rativ e l ifel ess ne ss

p .

S umma ry . The bodies of hig her ani mal s and pl ants are
s ubdi v ided in to va r i ous pa r ts ( org
a n s havin g d iff e r e n t st r ucture
)
a nd fun cti o ns . The se may be re solved i nto o ne or m ore ti ss ues ,

e ach of w hich co ns i s ts of a m ass of si m il a r cells ( o r thei r de r i va

tiv es) hav ing a si m ila r f unctio n The cells are small masses of
.

l i ving mat ter or protoplas m which depo sit more or le ss lifele ss

, ,

m atte r eithe r arou nd ( outside) the m o r w ithi n thei r substa nce .

I n the forme r case the protopl a m m ay co nti nue to live or it s

,

m ay die a nd be absorbed I n the latter case it may like w i se live

.

o n for a ti me or m ay die either d isappea ri ng altogethe r or leav

, ,

ingbehin d a re s idue of lifeless matter .

Th e Or g a ni sm as a Wh ol e Up to this poi nt we have con

.

s idered livi ng o rga ni sm s f ro m an a nato mical a nd anal tical sta nd

y
po int and h ave ob se rv ed the ir natu ral subdivi sio ns i nto orga ns
, ,

t issues a nd cells We have no w o nly to r e ma rk th at the se p a rt s

,
.

a re m utu ally i n te rde pe nde nt and th at the orga ni sm as a whole

,

is g re a te r tha n a ny of its pa rts P r eci s el y as a chro no meter is

.

s u pe rio r to a n agg regate of wheel s and s p ri ng s s o a livi ng organ ,

ism is su pe rior in the solida rity of its p a rts to a me re aggregate of

o rga ns ,
t issue s and cells We shall soo n see th at in the livi ng
,
.

body these have had a co mmo n a ncest ry and s till sta nd in the
close st rel atio ns hip both in respect to st ructu ral co nt inuity and
 C H A P TE R III .

PR OTOPL AS M A N D THE C E LL .

I has been shown in the last chapte r that life is inhe rent in
T

a peculia r sub sta nce p rotop las m occur ri ng in defi nite masses o r
, ,

cells . I n other w o rd s p rotoplas m is the ph y si cal basis of l ife

, ,

and the cell is the ultimate visible st ructu ral u nit P rotoplas m .

and the cell de serve the refo re the m os t ca reful co nside ratio n ;
but because of the tec hnical difiic ulties i nvolved in the ir study
only such cha racte ri stics as are eithe r obviou s o r in dispe nsable to
the begi nne r will he re be dwelt u po n .

Hi st or i cal Sk et ch O rgans and tiss ue s a re re a dil y visible but

.
,

in orde r to resolve tissues i nto cells so m ethi ng mo re than th e

naked e e was necessa r The co m pou nd m ic ros co e came i nto
y y .
p
use about 1 6 5 0 and in 1 66 5 the E nglish bota nis t R obe rt H oo k e
,

annou nced th at a fa mili a r vegeta l ti ss ue co rk is made u p of

, ,

little bares or cells d is tinct f ro m o ne a no ther ”

Many oth er .

ob server s desc ribed si milar cell s in sec tio ns of wood an d oth e r

vegetal ti ssues and the w ord soo n ca me i nto ge ne ral use I t
, .

was not until 1 8 3 8 however and as a co nse q ue nce of a m ost

, ,

i mpo rtant i mproveme nt in the co mpou nd mic ros cope viz the ,
.
,

i nve ntion of the ach romatic objective that cell ular st ructu re
,

ca me to be recogn iz ed as an i nva ri able and fu nd am e nta l cha rac

teri stic of li v i ng bodies A t thi s ti me the bota nis t S chleide n
.

brought fo rwa rd p roof th at the higher pl ants do no t simply co n .

tain cells but are wholly made up of the m or thei r p roducts ; and
about a e a r later the z oologi st S chwa nn de m o n t rated that
y s th e
same is t rue of a nimal s This g reat gene raliz atio n k nown as
.

,
“ ”
the cell tlzeo ry of S chle iclen a nd Schwa n n laid the bas is fo r
-

,
all s
ubsequent biological study The cell theo ry was at fi rst de
.
-

v elo ed
p upo n a pu rel y morphological bas i s I ts applica tio n to .

the phe no me na of phy siologica l actio n was fo r a ti me reta r ded

 HI S TOR Y OF “
CELL "
AN D “
P R OTOPL A S M ”
21

by the mislead i ng
ch r ac te r of the ter m cell a The wo r d itself “
.
”

s ho w s that cell s we re at fi rs t rega r ded as c a vities like the cells

(
of a ho ney co mb o r of a p ri so n) sur rou nded by solid wall s ; and
eve n S chleide n and S c h wa nn had no accu rate co nceptio n of their
t rue natu re S oo n afte r th e p ro mulgatio n of the cell theory
. -

however it was sho w n th at both the wall s and the cavity might
,

be wa nti ng and that therefo re the re mai ni ng po rti o n namel y

, , ,

the p rotopl as m w ith its nucleu s must be the active and es se nti al ,

p a rt The cell w as acco rdi ngly defi ned by Virchow and Max
.

“
S ch ult z e as a mass of p r ot o pl as m su rrou ndi ng a nuc le us and ,

i th
n is se se the word is used to day
n *
The word c ell bec ame -
.

the reafte r as i nappro p r iate as it w ould be if a pplied to the ho ney

withi n the ho ney co mb or to the livi ng p r i so ne r in a pri so n cell -
.

N eve rtheles by a cu ri o u s co nse r va t i sm the te rm w a and is re

s
, ,
s

tained to de sig nate thes s t ruct ures w hether occu r ri ng in m ass e s

e

as seg me nt s of the pl ant or a ni mal body or leadi ng i ndepe nde nt ,

live s as u n icellula r orga nis ms .

Pr ot oplasm w as obse rved lo ng befo re its signific ance w as

unde rstoo d The di scovery of its e sse nti al ide ntity in pla nt s a nd
.

animal s and ulti mately the ge ne ral recog ni t io n o f the ext re me

, ,

impo rta nce of the mile whi c h it every whe re pl ay s m ust be reck ,
~

o ne d as o ne of the greate s t s c ie ntifi c achieve m e nt s of thi s c en

t ury I t was Duj a rdi n w ho in 1 8 3 5 fi rst di sti nc t l y c alled atte n

.

“ ”
tio n to the i mpo rt a nce of the p ri mary a ni mal sub s t ance o r
“ ”
sa rcode w hich for ms the bodi e s of the si m ple s t a ni mal s .

Without clea rly recog ni z i ng this subs t anc e as the s e at o f life or ,

u s i ng the w ord protopl as m he n everthele ss described it as en ,

d o wed w ith the po w ers o f s po nta neou s move me nt and co n

tractility The w o rd protopla sm
. fi r st ; ” A do /rm ,

fo rm) was a ppa r e ntl y fi rs t u se d for a ni mal s ub st anc e by P urki nje

in 1 8 3 9 — 4 0 a nd next by H Vo n M o b ] in 1 8 4 0 to de sig nate
,
.
, ,

the gr anul a r visc id sub sta nce occ u r ri ng in pl ant cell s altho ugh -

bo th w orke rs w ere ignora n t of its f ull sig nific anc e I n 1 8 5 0 .

C oh n defi nitel y mai nt ai ned no t o nl y th at a ni m al sa rcode a nd

vegeta l protoplas m were e ss e nti ally of the sam e n atu re but ,

also th at thi s s ub sta nce is the r eal s e a t of vit ality and he nce to

be regarded as the ph y sical basi s of l ife To Max S chult z e .

It i p ib l th t in m f th l w t nd i mp l t rgn i m v n t h
s os s e a so e o e o es a s es o a s s e e e

n uc l u m y be w n t i n
e s a gas d i t i nc tl y diff r nt i t d b dy S p 1 9 3
a a s e e a e o . ee . .
22 PR O TOPL A S M A N D THE CE L L .

0 i ge er ll ig n ed the c r edit of h a v i n g fi nall

y placed
( 1 8 6 ) s n a
y a ss

h i m m
thi s conclu sion upo n a s ecu r e b as i s ; an d b y the e a n i ng of

the word P ot opla r sm w as s o exte n ded as to i n cl u de all l ivi ng

im l or vegetal I n th is s e n s e the wo r d i s
matt er whether an ,
a .

n ow unive rsall y e mpl oy ed .

r nd S t r u c tu r e P r otopl as m a n d c ell s d iffe r

A ppea a n ce a .

greatl in appe aranc e in different plant s and a ni mal s as w ell as ,

y
i n d iffere n t p a rt s a n d di ffe r e n t s t a ge s of develop m e n t of the
same i ndivid ual The a ppea r a n ce .
of p r otopl as m a n d the co n st i

tutio n of the cell a r e as a rule

m o st ea sily m ade out in ve ry
y o u ng struct u re s such as the ,

eggs of so me a n im a ls o r in
t h e cell s of y ou ng vegeta l

shoot s The egg of the s ta r .

fi sh fo r ex ample ( Fig is , ,
.

a s i ngle isolated cell of n ea rl y

t ypical fo rm and st ructu re .

I t is a minu te nea rl y s pheri ,

F I G 12 —
. S li g htl y d i gr m m t i fi gur f cal body (BLO i nc h dia ln eter)
. a a a c e O '

th gg
e e vum f t r fi h h w i gth
or o o a s a s s o n e

t u tur f t y p i l ll m m m br n ; "1 W I U C h th r ee pa r ts may be

-
,

s r c e o a ca ce . . e a e

n, n u l u ; pr t p l m ( y t p l
c e s m) o o
di sti ngui shed V iz 2 ( 1 ) the
as c o as .

.
,

cell body which for ms the b ulk of the cell ; ( 2 ) the n u cleus
-

,
a ,

r o unded vesic ul ar b od y suspe nded in the cell bod y ; ( 3 ) the mem -

bra n e or cell wa ll which i mmedi atel y su rr ou nd s the cell b od y

- -

O f the se th r ee the nuc leu s and cell —

.
,

,
bod y are mai nl y co m pos ed
of protopl asm while the me mbr ane is a lifele ss depo sit upo n the
,

exter i o r The protoplas m of the cell body is ge ne rally c alled

.
-

c ell pl a m or c to la s m that of the nu c leus n u c leo la s m

s th a t
y p p ;
-

, ,

is the livi ng m atter of the cell is diffe r e ntiated i nt o two difi erent
,

but clo sel y r el ated for ms of p r otoplas m cy toplas m and nucleo ,

pl asm .

The Cy t oplasm appears as a clea r semifi uid o r v iscid sub

s t ance co nt ai ni ng numero us m i nute g ra nule s and of a wate r
, y
a ppea rance tho ugh it shows n o te nde nc
, y to m i x w i th wate r .

U nde r very high powe rs of the microscope e speciall afte r t r ea t

y ,

m e nt w ith suit able re age nt s the clea r sub s tanc e is fou nd to hav e ,

a defi nite structure the preci se natu re o f w hich is in di spute

, .

B y so me ob server s it is de sc r ibed as a fibr o u s m es h w o rk o r retic

 THE M IN UTE A N A TOM Y OF THE CE L L . Q3

ulu m like a S po nge ; by othe rs as m ore n e a rl y like an e mul si o n

,

o r foa m co nsi sti ng of a m ore so lid fr am e w ork e nclo si ng i nnu

,

merable m i nute sep a r ate s pherica l c avitie s filled with liquid ; by

o t he rs s t ill as co mpose d of u nb ra nched th re a d s r un ni n g in all

di r ectio ns th r ough a more li qu id basis ; but its r eal natu r e is still

u nk n o w n .

I t is evide nt that the v is ible str ucture of protopl asm give s no

hi nt of its ma rvellou s po w e rs as the seat of v ital actio n and we ,

a re therefore c o m pelled to i nfe r th at it is e ndo w ed with a che m i

c al and m olecula r co ns titutio n extre m el y co m plex and prob abl

y ,

f ar exceedi ng in co mple xity th a t of any l ifele ss s ub sta nce .

Th e N ucleu s is a r ou nded body suspe nded in the cell sub -

st a nce ; it is dist ingu ishable fro m the l atte r by its highe r ref rac
tive powe r and by the i nte nse colo r it as sum e s whe n tre ated
,

wi th s tai ning fluid s I t is su rr ou nded by a v ery thi n me mb ra ne

.
o

a nd co ns is ts i nte rnall y of a clea r sub sta nce ( ac hro ma tin ) through ,

which exte nds an irregula r network of fib re s It

these fib res which a r e s tai ned by dy es I n the .

FI G . ( A ft er S h )
ac—Y s . o un ggr w i n g ll fr m
o ce s o th e ex tre m e t ip o f a s on ewo t rt
( Char a). m m m br n
, e a e ; 71,u l i ;
n c epr t p l o o as m; v, v ac uo e l fi ll ed w i th sa p .

me she s of the n etwork is suspe n ded in m any cases a second

rou nded body kn own as the n u cleolu s which stains eve n m o re ,

d eepl y tha n the n etwo rk itself .

Th e Memb r a ne or W all of the ce ll fo rms a r athe r tt k sac ,

24 PR O TOP L A S M A N D THE CE L L .

composed of a soft lifele ss mate rial closely su rr ou nd ing the cell

,
*
sub st ance .

As a seco n d ex am ple w e choose the g r o win g poi n t of a c o m

w ter pl t Fig 1 3 Thi s st ructu re is co m po sed
mo n a an ( C
-
h a ra
) ,
. .

of cell s which are more o r le ss angular in outli ne as a res ult of

mutu al p re ssure but show othe rwi se an u nm i stakable similarity
,

to the egg cell j ust desc ribed They diffe r mai nl y in the fac t
-
.

that the protoplasm of the l arge r ce ll s co nt ai ns rou nded ca vities ,

k nown as v a c uoles filled with sap also in the che m ica l c o m

,
“ ”
position ef the cell walls (here c o nsi sti ng of cellulose a sub
-

st an ce of rare o c curre nce amo ng ani mal s) .

Or i gin of Cell s and Genesi s of the B ody The body of every .

”
higher pl ant o r a nimal ari se s fr om a si ngle ge r m cell egg -

“ pore ” etc m ore or le ss nearl si mila r to that of the s ta r

s , ) .
y
fi sh desc ribed above and o ri gi nall y fo n n i ng a pa rt of the pa re nt
, ,

body The ger m cell therefore in s pite of e ndle ss va ri a tio ns in

.
-

.
,

det ail sho ws us the model afte r which all othe rs a re b uilt ; f o r
,

it gi v e s ri se to all the cells of the bod y by a co ntinued p rocess

of segme nta ti o n as follo ws
The first step (Fig 1 4 ) co nsi sts in the divi s io n of the eg .
g
i nto two si mil ar h alve s which differ f ro m the o rigi nal cell o nl y ,

in l acki ng me mb rane s both bei ng s urro unded by t he me mb ra ne ,

of the origi nal cell E ac h o f the h alve s divide s i nto two ma k

.
,

ingfou r in all ; the se agai n i nto two m a ki ng e ight a nd so o n , ,

throughout the ea rlie r part of the develop me nt B y th is p r ocess .

(k nown as the cle avage o r seg men ta tio n of the egg) the ge rm
cell gives ri se s uccessively to 2 4 8 1 6 3 2 6 4 etc de , , , , , ,
.
,

scendants fo r mi ng a p ri mitive body compo sed of a mass o f

,

nearl
y si milar cell s out of which by still fu rthe r divi sio n and
, ,

growth the f ully fo rmed body of the f ut u re a nimal is to be

,
-

built up The se cell s a re o nl y slightly m od ifie d but diff e r in

.

most ani mal s from the ty pical ger m cell in h a vi ng at fi r s t n o sur -

r oundi ng membrane s The me mb rane of the o rig . in al ge rm

cell meanwhile di sappea rs .

r
Th e w o dll h b n u d in Ch p I nd l se w h re t d n t th e
ce as ee se a . a e e o e o e
h mgm tt r w i th i n th m mbr n th l tt r b i n i r r t
g
v a e n d d p
e d uc f e a e, e a e e co s e e a o o
th e ce
ll r th r th n n int gr l p rt f i t I t i m r u u l t i n l ud th
a e a a e a a o . s o e s a o c e e
m mbr n in a d fi ni t i n f t h e c ll and as m tt r f c n v ni n
ce i t i
e a e e o o so e , a a e o o e e s
i n l ud d h r
c e e e .
 D E VE L 0P M E N T A N D D I FFE R E N TI A TI ON CE L L S
‘
OF . 25

The e mbry o nic body or embryo of ever y higher pl ant and ani
mal is de rived f ro m the ge rm cell by a proce ss e sse ntially like th at -

s t desc ribed though both the fo rm of the cells a nd the order of

j u ,

d ivi sio n a r e u su a ll y m o r e o r les s ir r egula r I n a nim al s the cell s .

FI G . IL -
Cl v
ea a ge or se gm e n t a t i o n of an ovu m . s h ow n i gs uc c es s iv e d ivi s i o n o f th e
germ -
l
c el (a )i t t w ( b ) f ur
n o o d ig
. ht o an e L a ter s ta ge s a re s h o w n at 6
a nd f . Th e fi r t f ur fi gur
s o di gr mm ti ;
es a re a a a c e and ft r
f a re a e Ha ts c h ek
‘
s fig
u es r o f th e d v l p m nt f
e e o v ry i m p l v rt br t
e o a e s e e e a e ( A mp h to r us ) .

t hu s fo rmed are u su all y naked at fi rst th o u gh they ofte n ao ,

q ui re a m e m bra ne in l ate r st ages A m o ng pla nt s o n the co n .

,

t rary the cell s u suall y po sse ss m e m bra n e s f ro m th e fi rst prob

, ,

a bl y be cau se thei r need fo r a fi rm oute r su ppo rt is greater tha n

the need fo r f ree m ove me nt de m a nded by a n i mal s *

.

Modifi cati on of th e E mbr y oni c Cell s Differ entiati o n Th e . .

c los e s i mil a rit of the e m br o nic cell s doe s no t lo ng pe rs i st A s

y y .

develop m e nt p roceed s the cell s co nti nu all y i nc rea si ng in numbe r

,

by divi sio n beco me modified in differe nt w ay s o r d g f eren tia ter

l ,
/

to fit the m for the many difi erent kind s of w ork w h ich they h ave
to do Tho se w hich are to beco me mu scle cell s grad uall y a ssu m e
.
-

a n e nti rel differe n t for m and s t ructu re fro m tho s e which are to
y
beco me sk in cell s ; and the f utu re n erve or gl and cell s take
- -

o n s till othe r for ms and s tru ctu r es The e mbry o nic cell s are .

gr aduall y co n ve rted i nto the ele ments of the differ en t ti ssue s

t hi s p r oce ss bei ng the def eren tia tion of tire tiss u es which h as

Fo r a mo re pr c i e s e a cc o un t o f ce ll div i i
-
s o n see p 83 . .
26 PR O TOPL A S M A N D THE CE L L .

bee m e ntio n ed o n p 1 1 — a n d are in this w a

y e na bl ed
alre ady n .

i i i of l b
to effect a p hys iolog ic a l d v s o n a o r .

The v ri tio
a a n s i n fo r m and structu r e which th u s a ppea r a re

d i ve r ified C ell s m a
y assu m e al m ost an
y co nceivable
e ndle s l y
s s .

d there are eve n cell s A m o b a or the colo r l ess e

for m a n ,
,

cor puscle s of the blood) whi c h co nti nuall y change thei r fo r m

o e t to m o m e t The v r i tio n in s t r uctu r e m a
y ih
f r om m m n n a a s .

volve any or all of the three ch ar acteri stic part s of the ty pical
cell bei ng at the same t i me acc o mpanied by v ar i atio ns of fo rm
,
.

e to u der t d the r efore h w cell m va r e n dl essly

I t is asy n s a n o s
,
a
y y ,

ppe ra ce while co n fo r m i n g m o r e o r less cl o sel y to the sa m e

in a a n ,

general type .

ea
M nw hile th e p r ot o pl a s m itse lf u nde r go e s ex t e nsive al te r a

tion E ven in young cell s o r in the ge r m cell i tself it may

. ,
-

contai n an ad mixtu r e of othe r substance s ,

and these may e ntirel y cha nge th ei r

ch ar acter o r (as is e s pecially common in
plant cells) may beco me mo r e abu n -

d ant as the cell gro ws olde r taking th e ,

shape of fl uid so lid o r eve n gas eous de , ,

posits C o mmo n examples o f such de .

po sit s ar e d r op s of w ate r oil and r e si n , , ,

granule s of pigme nt s ta r ch and solid , ,

proteid matte rs and c ry sta l s of mi ne ral

Fm 15 R n vi r )
,
. (A ft . er a e .

P rt f
a in g l fi br
o a sf l sub sta nce s like calciu m ox a l ate
e e o vo phos ,
u t y mu l f m th l g
B ub
ph ate an d carbo nate an d si l i ca
n ar sc e ro e e

p p m wp m m ;
.
,
o f bb it
a ra .
' s

n ,u l u n ( x 700)
c e s . bles of g as so meti m e s appe a r in the p r o
.

toplasm but thi s is exceptio nal

,
The livi ng sub sta nce itsel f .

often change s in appea rance as the cell s beco me di ffe r e ntiated .

The protopl asm of volu nta ry mu scle s ( Fig 1 5 ) is fi rm clea r .

, ,

n on gr anul ar
-
highly refr acti v e and a r ra nged in al te r nati ng
, ,

b ands o r stripe s of darke r and ligh te r subst ance I n so me cas es .

the outer portions of the ski n or o f a hai r as e xpl ai ned , ,

in C h ap I I ) the modific atio ns of the cell sub sta nce bec o mes so
.
-

great that both its ph ysi c al and che mical co nstitutio n a r e e nti r el y
altered and it is no lo nge r protopl as m but so me fo rm o f lifele ss
, ,

matter .

Pr ot oplasm in A cti on We may now b r iefly co nside r proto .

pl asm fr om the d ynamical or ph y siological poi n t of vie w We .

 PR O TOPL A s a ne M O VE M E N TS . 27

k now that livi ng thi ngs are the se at of active changes whic h ,

ta ke n togethe r co nstitute thei r life I n the l ast anal ysi s the se .

c ha nge s a r e u ndoubtedl y che mic al actio n s taki ng place in th e

pr o toplas m which may or m ay n ot p r oduce vi sible re sult s

, .

The re is no doubt that exte nsive and p r ob abl y ver y co mplex

m o lecul ar actio ns go o n in the protopla sm of y ou ng gro wi ng
cell s though it may appea r a bsolutel y quie sce nt to the ey e eve n
, ,

u nder a powerful m ic r oscope I n other c ase s the che m ic al .

,

actio n p r o duce s pe r ceptible ch anges in the protopl asm — for in ,

sta nce s o m e f o r m of
,
m otio n — j ust as t he i nvi s ible c he mi c al ,

actio n in an electrica l b atte ry m ay be m ade to pr o d uce vi sible

effec ts ( light loco motio n etc ) th r ough the age nc y of an electric al
, ,
.

machi ne .

A fa milia r i nsta nce of p r otopl as mic move m e nt is t he co ntr ae

tio n of a mu scle Thi s p r o c e ss is m o s t likel y a ch ange of mc lee
.

ula r a rra nge me nt ca usi ng the mu scle while keepi ng its ex ac t

, ,

bulk to cha nge its fo r m the tw o e nd s bei ng br o ught ne are r

, ,

togethe r ( Fig The vi s ible ch ange .

of for m is he re su ppo sed t o be due to an

i nvi sible cha nge of m olec ul a r a rr ange
m e nt a nd thi s in tu r n to be c oi nc ide nt ,

with c he mic a l actio n t aki ng pl ac e in the

livi ng subst ance .

A striki ng and be autiful ex am ple

of m ove me n t in protopl asm o cc ur s in
the s i mple orga ni sm k no wn as A mccba
( Fig 8 4 p The e ntire bod y of .
,
.
.

thi s ani mal co nsi st s of a m ass of naked

protoplas m e nc lo si ng a n u cleu s o r ,

so meti m e s tw o ; in o ther w o rd s it is a
A ,

_ i gle n ked cell The protopl asm of

p m 1! , Ch n g f f rm in a
s n a
e o o a
.

co nt r ti gac “ A ~m u an active A m a ba is in a st ate of ce as e

n " N SC . s“

l i th
c e nrd i r y e oxt d na or e en

e d t t s a 3 th m m u l les s m ov e me n t co ntr ac t i ng expa nd i ng

e ; , e sa e sc e , , ,

Wh mm wd ( Di gr m ) flowi ng and ch angi ng the fo r m of th e

en o o ‘ c ' a a
,
“ l ’
ani mal to such an exte nt th at it is k n o w n as the Pro teus
ani m alc ule The w hole m ove me nt is a ki nd of flu x A portion
.
.

of the pro topl as m flo w s o ut f r o m the mass maki ng one o r mo r e ,

prol o ngatio ns ( p seudop ods) i n to w hich the r emai nde r of the

p rotoplas m fi nall y pas ses so that the whole body adv ance s in the ,
 T
PR O OPL A S M A N D THE CE L L .

28

di rection of the flo w .

m fi w arou nd
p r otoplas o s
o w r d l avi g the r efu se
asm fi w n a e n
withi n the body the p r otopl o s ,

o wmgmay
,
d d d a
y tl l l S fl
behind H o r .
u a fte r hou r a n a
y
go o n a n d there i s perhap s no
,
d sugge s ti ve
more fasci nati g n a n

spect acle
k n ow n to the biologi st .

ch ge of fo r m is ex
A sm i il ar a n

hibited by the colorle s s co r pu scles

d othe r blood in
of amphibian a n ,

whi ch it may be ob se r ved though ,

f ar le ss sa s ti f acto r il y if A m oe b ce
,

cannot be obtai ed n A m o n g pl a n ts . ,

prot opl asmic m ove m e n t s of perhaps

equ al beauty may be obse r ved .

O ne of the si m ple st is k n o w n as the

rota tion of protoplas m which m a

y ,

FI G —A ll f t n w r t ( N it l
. l7 . ce o a s o e o e

l ) h w i n g th r t ti f pr t
—Tw ll
a s o e o a on o o o

pl m ; th
as rr w h w th di
e a o s s o e rec FI G . 17a . p rt o ce s a. a

t i n f th fl w m m mb r n f
o o e o . , e a e o th i r fr m t h t i p f
d o l f e o a
“
ea
“
of a

th e ll ; n n u l u p p i t t
ce , c e s, o os e o t n w t h wi n g r t t i n f
s o e or , s o o a o o th e

wh i h i c nd p pr t pl
s a sec o m; r , o o as . p r t p l
o o m i n t h d
asi t i n f e rec o o th e

l g
ar e cen tr l v u l fi ll d w i th
a p
ac o e e sa . ar orw s .

be studied to advant age in r ather y ou ng cell s of sto neworts ( Ch a ra

or N itella ) These cell s h ave the for m of sho r t or el o ngated
.

cyli n der s which are ofte n poi nted at o ne end ( Fig Th e .

 P R O TOPL A S MI C M O VE ME N TS . 29

p r o t oplas m is su rr ou n ded b
y a delicate m e m b r ane which t hus

for ms a sac e nclos i ng the pr o toplasm I n ve ry you ng cells the .

p rotoplasm en ti r ely fills the sac ; but as the cell gr ow s olde r a

d rop of li quid appears n ea r the ce nt r e of the mass and i nc r eases
in s i z e u n til the p r otoplas m is r educed to a thi n l ay e r (p rimor
d ia l u vriele) linin g the i nn e r su r fac e of the m emb r ane ( co mpa r e
,

Fig . I n favo r able cases the e nti r e m ass of p r ot o plas m is

see n to be flowi n g s teadil y a rou nd the i nside of the sac as in ,

dicated by the a rr ow s in Fig 1 7 I t m oves u pwa r ds on one

. .

side down wa r d s o n the O ppo s ite side and in O ppo site di r ectio ns
, ,

a c r o ss the e nd s fo r mi ng an u nb r oke n ci r cuit

,
The flow is ren .

de r ed m o r e co nspicuou s by va r iou s granules and other lifele ss

m asses flo ati ng in the p r otoplas m and by the l a rge oval nucleus
o r nuclei all of which are s wept o nwar d by the cu r rent in its
,

c easele ss r o und A s i mila r rotatio n of protoplas m oc c ur s in many

.

othe r v egetal cell s o ne of the best exa mple s bei ng the leaf cells
,
-

of A na c/ca ris .

A s eco nd and s o m ewhat m o r e i nt r icate ki nd of m ove m e nt in

vegeta l p r otoplasm is kn o w n as c irc ula tio n Thi s diffe rs fro m .

r otatio n chiefl y in the fact th at the protopla sm tr a v el s n o t o n l y in

a pe r iphe r al s t r ea m but also in st r and s w hich run ac ross th r o ugh
the ce nt r al space ( vacuole) and thus fo r m a loo se netwo r k C ir .

6
E 1 0 18
. .
—Fl ow e r—
c l us t r ( ) n d i n g
e l t m n (b ) f
a a s e s a e o a cu lt iv t a ed s p iderwo rt Tr ades

ca nt ia ) . h, h ir up n th t m
a s o li ghtl y r
e s a en . a. s ed uc ed ; b, s l i ght ly nl rg d
e a e

culatio n is well see n in cell s co mpo si ng the h air s of v ar ious pl ants ,

s uch as the commo n n ettle ( Urtica ) the s pide r wo r t ( Tra des ,

30 PR OTOP L A S M A N D THE CE L L .

ca n tia) the hollyhock

ce r tai n s pecies of gou rd s
( A ltiuea ), an d
,

I t may be co ve nie n tl s t u died in the h a i r s u po n

curbi ta ) y
n
( Cu .

the stamens of the c ultiv ated spider wo r t ( Tradesca n tia ) The .

fl ower of thi s plant is shown in Fig 1 8 a and one of the .

, ,

stamens with its h air s at b E ach hai r cons i sts of a .

Frot lil —E n a ged lr cell s of the h i r fr m th t m n f th p i d rw rt A

a s o e s a e s o e s e o fi ve
é ellé s o mew a
. .

h t en la rgd
p ro oe , t p l m t h w ; B d C l l mu h m r
as no s o n an , ce s c o e en
la l g
a
‘
ed . s h o wi ngt h e c “c ula o
’

t i n f pr t p l m i nd i t d b y th rr w
O o o as as ca e e a o s ; n,
nuc eus l .

of elongated cell s covered by delicate me mb ranes and co nn ected

b y the i r e nd s A s in N itella the p r otopl as m doe s no t fill the
.
,

c av i ty of the sac but for ms a thi n li ning (p ri mo rdial utricle)

,
 CI L I A R Y A C TI ON . 31

on its i nne r face ( Fig . Fro m thi s lay er delicate th read s of

p r otopl asm reach i nto and pass through the ce ntr al c avity where ,

they ofte n b r anch and are co nnected together so as to for m a

ver y loo se net w o rk The nucleu s (n ) is e m bedded either in the
.

e r iphe r al l a e r or at so m e poi nt in the n et w ork an d the thre ad s

p y ,

of the latte r al ways co n verge m ore o r le ss regularl y to it I n .

active cells cu rre nts co n ti nu all y flow to and fro th r o ughout th e

whole m as s of protoplas m I n the th r ead s of the net work gr an
.

ules are bo r n e rapidly alo ng glidi ng no w in o ne di rection n o w

, ,

in anothe r ; and although the flow 1 s u suall y in o ne di rectio n in

an p a r ticu la r th r ead no sy ste m c an be di sc o v e r ed in the co m
y ,

plicated m ove me nts of the whole I n the larger th r e ad s the .

‘

cu r iou s spectacle ofte n appea rs of two rapid c urre nt s flowi ng in

opposi te d ir ectio ns o n oppos ite side s of the sam e thread The .

cu rr e n ts in the thread may be see n to joi n c u rren t s of the pe

riphe ral laye r which fl o w he r e and the r e but without sthe regu ,

larity ob served in the p r otopla m of ZVitella s The prot o pl as mi c .

n et w o r k al so as a whole u ndergoe s a slo w but stead y ch ange of

, ,

fo rm its deli ca te s tr and s slo wl y

,

sw ayi ng hi ther and thither w hile ,

the nucleu s t ravel s slowl y f r o m

poi nt to po i nt .

Fi nall y we may co nside r an

,

example of a fo r m of pro to plas

mic move me nt kn o wn as c ilia ry
actio n whi c h pl ay s an i m po r ta nt
,

ré le in ou r o wn live s a nd tho s e

of lower a ni mal s and of so me

pl ants The i nte r io r of th e tra
.

chea o r wi ndpi pe is li ned by

, ,

cell s h a ving the for m sho w n in

Fig 2 0 A t the free su r face of
. .

the 08 11 (tu r n ed to ward s the C 3 V “ F 20 ( A f t r K l i n )—Th r i l t d

I G . . e e . ee so a e

ty O f the trache a) the protopl as m i l i t d ll c f r m at he i t r

cei r sf t h
o e n e o o e

w i d pip f th n t the o i l i t th e ca . c, e c a a e

1 8 p r oduc ed lI l tO deh ca te v i br a fr d; th u l u ; p th
ee e n 71, pr te n c e s . e o o

tor y fil ame nts havi ng a sickle P l m ( Hi g h ly m g

as ni fi d ) a e .

sh ape whe n be nt ; these a r e k now n as c ilia ( cilium an eyela sh) ,

.

The y a re so small and lash so vigorousl y as to be nearl y o r quite

i n vi s ible u nt il the m ove me nts are in so m e way made sluggi sh .
32 P R O TOPL A S M A N D THE CE L L .

The movement is then seen to be m ore r apid and vigo r ou s in one

th i the other all the cili a worki ng togethe r like
d irection a n n ,

the oar s of a r ow bo at acti ng in co ncerted m otio n B y this

- .

a ctio n a defi n ite c u rre n t is p r oduced in the s urrou n di n g m ediu m

thi ca e the m uc of the tr che ) flowi n g i n the d i r ectio n
( u a a
in s s s

the m ore vigoro m ove m e t I n the t r ache a thi s m ove m e n t

of u s n .

is upw a rd s to w a rd s the m o u th a n d m ucu s

,
d u s t etc a r e thu s
, , .
,

re moved fro m the l ungs and w i ndpi pe I n ma ny lo w er an i m al s

.

a nd pl ant s e speci all y in the e m b ry o nic s tate

,
c ilia are used as ,

o rgans of loco motio n se r i ng as o ar s to d r ive the o r ga n is m

v

through the water The male reprod uctive ge rms of plants and
.

a ni mal s are al so propelled in a si mil ar fas hio n .

I n all the se for ms of vital ac tio n the p r o toplas m is v isibly at

w ork . I n m o st c ase s ho wever no move me nt s of the pr o to pl as m
, ,

in cell s can be detected B ut it is ce r ta i n f ro m i ndirect evide nc e

.

t h at pr o topl asm is no le ss ac t ive in tho se m ode s o f physiological

a ctio n th at g ive no vi sible out ward sig n as f o r ex a mple in an ,

acti v e ner v e cell or a secreti ng cell Thi s ac tivi ty bei ng mel ee

-
.

ul ar and che mic al is beyo nd the re ach o f the mic ro sco pe but it ,

is no ne the le ss real ; and the pl ay o f these i nvi s ible molec ul a r

a ctio ns is do ubtle ss f ar m ore t umul t uou s and c o m plica t ed th a nthe

vi sible move me nts of the pr o topl as mic mass di spl ay ed in N itella

o r in a nettle h air -
I t is of t he ut mo st i mpo rta nce th at the stu
.

de nt should at t ai n to a ful l a nd vivid se nse of the rea li ty a nd

e nerg
y of thi s i n vi sible ac t ivi t
y eve n i n p r otoplas m wh i ch ( as is

o rdi naril
y the ca se ) de r the clo se st sc r uti ny appea r s to be abso
u n

lutely quie sc e nt .

The Sources of Pr ot opl asmi c E nerg y Whe nce co mes the .

p ower required for protopl as mi c actio n a nd ho w is it expe nded ? ,

The a nswer to thi s q ue sti o n c an be give n a t thi s poi nt o nl in

y
ver y ge ner al ter ms I t is certai n th a t p rotoplas m w o r k s by
.

m e ans of c he mi c al actio ns taki ng plac e in its o wn sub s ta nc e

;
a nd it is further ce rta i n that the se ac tio ns are br o adl s pe aki ng
y , ,

processes of oxid ation o r co mb ustio n ; f o r in the l o ng run all

f o r ms of protoplas mic ac t io n i nvolve the taki ng up o f oxyge n
a nd the liber atio n of c a rbo n dioxide
E nergy is the refo r e se t
.

i
free n livi ng active protoplas m so me wh at as it is in the co m
,

b ustion of fuel under the boiler of a steam e ngine a n i th

d n is -

process the p r otopl as m like the coal is g r adually u sed up disin

, , ,
 CHEM ICA L R ELA TI ON S OF PR O TOP L A S M . 33

te g rates , and w astesa w ay givi ng off as waste matter the v ariou s

,

che mic al p r oduct s of the co mbustio n and liberati ng e nergy as ,

he at and mech anic al work The lo ss of substa nce is however .

, ,

c o nti nua ll y m ade good ( much as the coal is r eple nished ) by th e

ab sorptio n of new sub st ance in the fo r m of food which m ay ,

co nsi s t of actu al proto plas m de r ived fro m othe r li vi ng bei ngs;

,

or of sub sta nce s co nvertible i n to it The se substa nce s are in .

so me u nexplai ned w ay co nverted i nto protoplasm an d thu s

built i nto the livi ng fabric .

“
To thi s du al p r oce ss of w a ste ( ka ta bolimn ) and r ep ai r
a nd bo lis m
/ is applied the te r m meta bolis m which m u s t b e ,

co nsidered as the most c ha r acte r i stic and fu nd ame ntal p r operty

of livi ng matte r I t is ev ide nt fro m the fo regoi ng th at met a
.

holi s m i nvolves o n the o ne ha nd a de st r uctive actio n (An te bel

is m) thro ugh which p r otopl as m di si ntegr ate s and e nergy is set
f r ee a nd o n the other ha nd a co nstru c t ive actio n ( ( m a bolis m)
,

w he reb y ne w prot plas m is built u p fro m the i nco me of food a nd

o

fre sh e ne r gy is sto r ed I t is a m o st r e mar ka ble fact th at as f ar

.

as k no w n the co ns tructive acti o n r es ulti ng in the f o r m ati o n o f

n e w pr o t o pl as m n eve r take s place ex c ept t hrough the i m m edi at e

age nc y of p ro to pl as m a lre ad y exi s ti ng I n other w ord s there is .

,
“ ”
no evide nce that sp o nta ne o u s ge ner atio n or the pro d uctio n
of li v i ng fro m lifele ss mat te r w itho ut t he i n fl ue nce of antecede nt
life ever ta ke s pl ac e N o r is the re a ny evide nc e th at any e nergy
.
’

“ ”
can be ge ne ra ted but ra ther that the vit al e nergy o f livi ng
,

thi ngs is o nl y the t ra nsfor med e nergy of their food and th at ,

“ ”
vita l force havi ng an origi n el se w here than in s uch e ne rgy
doe s n ot exi st .

Ch emi ca l R elati ons We k now n othi ng of the p r eci se che m i

.

cal co m po sitio n of li v i ng protopl a m bec au se as has bee n said s

, ,

p livi ng p r otopl as m c a nnot be s ubjected to che m ic al analy

( .

without de s troy i ng its life B ut the r e su lts of che m ic al ex

.

am inatio ns le a ve no do ubt th at the m olec ule s of proto pl as m a re

highl y co mplex and a r e p robably sep arated f r o m one a nothe r by

laye rs of wate r .

A P aormns
. I t has alr ead y bee n stated ( p 3 ) th a t th e
. .

char ac te r i stic products of the analysi s of protopl as m are the

group of closel y rel ated sub sta nce s k no wn as p roteids B ut pro
-
.

te ids fo r m o nly a sm all pa r t of the total w eight of any plan t o r

34 P R O TOP L A S M A N D THE CE L L .

i
a n ma
l bei g al w a s associ a te d wi th q u a n titie s of othe r sub
,
n y
v e the white of egg w h ich i u s uall ta ke n fo r
s t ance s . E n a n s y ,

t pi l proteid c n t i n o n l t w elve per ce nt of ac tual proteid

a y ca o
,
a s y
m a tter the re m a i n der co n s i s ti n g chiefl y of w ate r The fo l low .

t
ing a l s b e ho w s the perce n ta g e of proteid s a n d othe r m att e rs in

a few famili ar organi sms and thei r prod ucts

P R O X I M A TE PE R C E N TA G E C O M P O S I TI O N O F S OM E C O MM ON
S U B S TA N C E S .
‘

A rr g
a n ed acc ord n i gto ric h n ess i n P rotc ids .

A ll proteid s ha v e ne arl y the same ch e mi cal c om po s ition and

S i mi l ar ph sic al propertie s ho weve r differe t
.

y n ma be the for ms
y ,

of protopl asm in whi ch they o cc ur The anal y s i s o f proto pl as m .

are its basi s teache s us reallv

o r r ather o f the proteid s which
,
n
othi ng of its vital p r opertie s but se r ve s o nly to sh ow the ,
c he mi c al
co mpo sitio n of the materi al b asi s b v whic h th ese are
man i fe sted .

j
I roteids are so called fr o m thei r r esembla e
nc to p r otein
( n p mr o s firs t) a h ypothetical sub sta nce fi r s t de sc r ibed and
, ,

iti
g g gg
r
mp o s m
m lé flz bx d dii
c o on b w
f or t h e S m i th er

I ii—
t t t i n th ugh
.

so n i a n I ns i u of w, o a e e xa e
n um e s 2, 10
e a 24
br . 11 , 12. fr m J h n n H w Cr ps G r w N Y 1883
o o so
’
s o o o v 0
w .
 P R O TE I D S . 35

n am ed by Mulder A cco r ding to Hoppe S ey ler they have

.
-

a
p
p r oxi m atel y the followi n g perce ntage co mpositio n

A s mall
quan tity of pho spho r us is also very freq ue ntl y pres e nt .

A ssoc i a ted w ith the se ele m e nt s are al way s s mall qu antitie s of

va riou s mine ral sub sta nces which r e mai n as the ash when proto
pla s m is bu rn ed ; but the natu re of thei r rel atio ns to the other
ele m e n ts is u nce r tai n The ash v a r ies both in qua ntity and
.

chemical co m positio n in diffe r e nt a ni mals an d pl ant s I n th e .

white of eg
-

g the
-
chief co n stitue nt s of the ash are potassiu m chlo

ride ( KC l) and sodiu m chloride ( N aC l) the for m er bei ng much ,

in exce ss The re mainde r co ns i sts of pho sph ate s sulph ate s and
.
, ,

ca r bo nates of sodiu m an d potassiu m w ith mi nute q uantitie s of ,

c al c iu m m agn e siu m and i r o n and a t ra ce of silico n

, , , Ma ny .

o the r min e ral su bsta nce s occu r in associatio n w ith othe r ki n d s of

p r oteids bu t al w ays in ve ry small p ropo r tio n These salt s are in

, .

so me w ay esse ntial to the activity of p rotopl asm as we k no w by ,

familia r e xpe r ie nce Man like othe r a ni mal s and the pl ant s
.
, ,

r eq u i r es ce rt ain mi ne r al sub s ta nce s (e g co mmo n salt) but we . .

,

ha v e no kn o wledge of the p a rt the se pl a y in protopl asm .

I t is i m po rta nt to note the clo se c he mica l si mil a rity of an i mal

and vegetal p r ote id s b eca u se this is o n e r eas o n for rega rdi ng
,

veg etal and a nimal p rotoplas m as es se nti all y si milar in other re

s ec ts
p . The follo w i ng table fro m J oh n so n a fter G o r up B esanez
,
-

and R itth ausen s hows the perce ntage co mpo sitio n of v a r iou s p r o
,

t eids and p roves that the di ffe r e nce bet wee n vegeta l and a n i mal
,

p r oteid s is che micall y n o gr eater tha n th at betwee n d iffe re nt

kin ds of vegeta l o r diff e re nt ki nd s of a ni mal proteid s :
PE R C E N TA G E C O M P O S I T I O N O F P R O TE I DS .
36 PR O TOP L A S M A N D THE CE L L .

h r
T e e is a c o rr e sp o n d i n g l i k e n e s s i n t h e g e n e r a l p r o p er t i es a nd r eac t i ns o

r
o f p o e ds
t i . T h e y a r e c o ll o i d a l o r n o n d ifi u s i b l e i e-
t h e
y
w ill no t pas s
,
. .
,

h r h m m b r f d i l r r o nl y W i th g r ea t d iffi c ult y ; th ey
t o ug t h e e a n e o a a y se o ,

a re a
r r e l y c r y s ta ll i n e th e y r o t a t e t h e p l a n e o f p o l a ri z e d l i g ht to t h e l ef t .

h
T oug no h t a ll s o l u b l e in w a te r th ey m a y,
b e d i ss o l ved by th e a i d o f h ea t

t
in s o g r n a c e t i c a c i d a n d in c a u s t i c a l k a l i es b u t a re i nso l ubl e in co l d a h
,

l u t e a l c o h o l a n d in e th e r T h e y m a y be pr ec i pi t a t ed f ro m s o l ut i o n by
so .

s trongmi nera l acids etc ,

Many pro t eids are pr eci pi t a t ed b y h ea t ( a pro
.

h i h ll w r th o t e th a t t e mpera
c ess w i d g u l t i o n ) a n d i t is o y o f n
—
c s c a e co a a

t ur es wh i ch produce coagl t i 0 7 5 C ) p rod uce a l s o t h e

°
pr t i
°
u a o n o f o e d s ( 4 .

A mo ng s t t h e o rg a ni c p ro x i ma t e pri ncipl es
“
dea t h o f mos t org a ni s ms .

wh ic h ent er int o th e co mposi t ion o f th e t iss ues a nd o rg ans o f l ivi ng bei ng s ,

those bel o ngingto th e cl ass of p roteid or a lb umi no us bodies occupy quite

a pec ul i ar pl ace a nd req uire an ex cept i o nal t rea t ment f o r th ey a l o ne are ,

nev er ab sent f ro m th e act i v e l i v i ngcell s wh i c h we rec o g ni ze as t h e p ri

mordia l s t r uct ures o f a ni mal a nd v eg et ab l e o rg ani s ms I n th e pl a nt wh i lst .

,

we recog nize th e wide dis t ri b ut i o n o f s uch co ns t i t uent s as c ell ul ose a nd

c hl o roph y l ,
a nd ac k no wl edg e th eir re ma r k a b l e ph y s i ol o g ica l i mpo r t a nce ,

we at t h e sa me t i me are fo rced to admi t th a t th ey cc upy a lt g e th er a o o

di ff erent posi t i o n f rom th at o f th e pro te ids o f t h e p ro t o pl as m out o f wh i ch

th ey were ev ol v ed We may h a v e a pl a nt wi th o ut c hl o r ph y l and a v eg
. e o ,
o

t ab l e cell wi th out a cell ul ose wall b ut our very co ncept io n o f a l i v i ng , ,

f unc ti onall y act i v e cell wh eth er v eget a bl e o r a ni mal is necessa ri l y asso

, , ,

c iated wi th th e i nt eg ri t y of its pro t o pl as m o f wh ic h t he i nv a ri a b l e o rganic ,

c o ns t i t uent s a re pro t ei ds .

I n th e a ni mal t h e pro t ei ds cl ai m even mo re s t ri k i ng

,
l y o ur att ent ion
th an in th e v eget abl e in th a t th ey fo r m a v ery muc h l a rge r pro po rt i o n o f
,

th e wh ol e o rg ani s m a nd o f ea c h o f its t i ss ues a nd o rg

, a ns We ma y i ndeed .

say th a t th e mat eri a l s ubs t r a t um o f th e a ni ma l o rg a ni s m is p ro te i d a nd ,

th a t it is th rough the agency o f s t r uct ures essent i a ll y pro te i d in nat ure

th a t th e ch emic a l and mec h a ni ca l proc ess es o f t h e bo dy a re eff ec ted I t is .

tr ue th a t th e protei ds are no t th e o nl y o rg a ni c co ns t i t ue nts o f th e t i s s ues

and or g a ns and th a t th ere are o th ers

, p res e nt in mi nute q ua nt i t i es wh ich
, ,

prob a bl y are al mos t as wi del y di s tri b uted such as f o r i ns t a nce ph os ph o rus ,

con t ai ni ngf a tt l ycogen yet a v o wedl y we ca n ( a t th e mos t )

y bodi es and g , ,

o nl y s ay p roba bl
y and ca nno t in r ef erence to th es e a ffi r m th a t wh ic h we
, , ,

may co nfi dentl y affi rm o f th e pro t ei ds th a t th ey a re i ndi s pe ns a b l e c ns t it -

o

neut s o f ev er y l i v i n
g ac ti v e ani ma l t i ss ue and i ndisso l ubl y co nnec ted
, , ,

wi th ev ery mani f es t a t io n o f a ni ma l ac t ivi t ”

y ( G a mg ee P h ys io log
. ica l ,

Ch emis try C h ap I )
, . .

The molec ul ar i nst ability of proteid s is prov ed by the ease

W i th which the ma be deco m po sed i n to s i m ple r co m po und s
y y ;
the i r complex constit ution by the n u merou s co m pou nds the m ,

y co mplex whi ch may thu s be de r ived o r

sel v e s ofte n highl
,
s plit off fro m the m .
 UA R B OH Y D R A TE S A N D FA T8 . 37

A m ongst the othe r m atte r s fou n d in protoplasm o r clo sel y

associat ed w ith it those of m ost f r eque nt occu rr e n ce and greate s t

ph ysiological i m po rta nce a r e two gr oups of less com plex sub

sta nces v iz ca rboh y dr at es an d f at s These con tai n ca r bo n h y
,
.
,
.
,

droge n and oxyge n but no nit roge n ; they do n o t appea r to be

, ,

closel y relate d to pr ot ei ds in che m ical co nstit utio n but they ,

o ccu r to so nre exte n t al most eve ry whe r e in li vi ng organ i sms and ,

in m any i nst ances a r e k now n to be of g r eat i mpo rta nce espe ,

c ially in n ut ritio n They are r ich in potential e ne rgy and mo

.

bile in m olecul ar a rr a nge m e nt ; he nce it is not st range that they

figu re l a rgely in food an d a r e ofte n l ai d by as r ese r ve food,

mate rial s in the o rga nis m .

B C A R B O H Y D R A TE S
. The se sub sta nces are so called becau se
.
,

be side s carbon the y co nta i n h yd roge n and oxyge n u nited in the

,

sa m e p r opo rtio ns as in water They i ncl ude starch va r ious .

,

ki nd s of su ga r cell ulo se and gl y coge n S t ar ch

, ,
is of .

ve ry fr eque n t occu rre nce in pl an t cell s whe r e it appea r s in the -

fo r m of granules e mbedded in the protopla sm ( Fig C el .

lulo se havi ng the sa me che mica l fo r mula as sta rch but quite
, ,

diff ere nt in ph y sical prope rtie s al mo s t i nva ri abl y fo r ms the basis ,

of the cell m e m bra ne i n pla nt s

-

0 F A TS . These are of e s peci al i mpo r ta nce as r e se r ves of

.

food mate ri al s
-
in adipose ti ss u e and in seed s) They co n .

tai n much less oxy ge n tha n the c arbohy dr ates ; are therefo re
m ore o xidi zable and r iche r in pote nti al e ne rgy,
*
They com .

m oul y occu r in the fo r m of d r ops s uspe nded in the protoplasm

( Fig . a nd a r e e s peci all
y co mm o n in a ni m al cell s though b
y ,

no m ea n s co nfi ned to the m .

Phy si cal R elati ons The appea ra nce consi stency etc of
.
, , .
,

protoplas m have alread y bee n de scribed ; but it still r e mai ns to

s pe a k of ce rtai n of its othe r ph sical p r ope rtie s an d especiall
y y ,

of th e ma nne r in which its activity is co nditio n ed by va rious

ph ysica l agent s .

R ela tio ns o f Vi ta l A ction to Temp era tu re I t is a gen e ral .

law that withi n ce r tain li mits heat accele rates and cold di mi n ,

i shes the activity of p r otopl asm We kn o w th at cold tend s to

,
.

A cco d r i ngto
rf l r rh n p ca e u es ea c es , o e o und o f b utt er c nt i nso a 5 65 4 f t oo

t n
o s, a nd a p und f s ugr 2 7 5 5 f t t n
o o a oo -
o s , of e ne rgy A p . o und o f p ro t id is
e

nea rl y q uiv l ent in th i s r pec t t p und

e a es o a o of ca r b h y dr t
o a e .
38 P R O TOP L A E M A N D THE CE L L .

benumb ou r own bodie ( s prov i ded the y beco m e r e a ll y chi lled ) an d ,

lower a i
nm s al the he a r t beats m ore s lowl y the m ove m e n ts be
in ,

i beco m e s low an d
come sl uggi sh or cease b r eath g n s

—
,

a word all of the vit a l a ctio n s beco m e dep r e ss ed w he neve r ,

,
I
the ordi nary m te perature is su ffi cie n tl y lo w e r ed f we c
hi ll .

g l
.

the rot ating protopl as m of C li o/ m o r N i tella the V i b r ati n c r ia

,

of c ili ated cell s or an actively flo w ,

ing A moeba the m ove me nts ,

become slower and fi nall y cease altogether

,
.

the other h d m oderate w ar m th favo rs protoplas mi c

On an ,

action B .
e nu m bed fi n ger s beco m e o n ce m o r e n im ble befo re the
war mth of the fire I n a hot roo m the f r og s he a rt beats m o re
.
’

rapidly cilia l as h more e nergetica ll y the A moeba flo ws m o re

,
,

rapidl y and the p rotopl asm of 01 mm cou r ses mo re s wif tl y I n

,
.

the w i nte r month s the p rotopl asm of plant s and of many anima ls
is in a st ate of comparative i nac tivity Mo st plants l ose the ir
.

leave s and stop gro wi ng ; ma ny ani mal s b ury the mselves in the
mud or in bu r rows and pass the wi nter in a deep sleep ( hiberna
,

tion ) duri ng which the vital fi r es bu rn low a nd see m well nigh

-

exti ngui shed The war mth of sp r i ng re esta bli shes the activity
.
-

of the protoplasm and in co nseque nce a ni mal s awake f r o m thei r

,

sleep and plant s put fo rth thei r leave s .

B ut thi s law is tr ue o nl y w ithi n ce rt ai n li mits E xt re me .

heat and cold are alike ini mical to life and as the te mpe ra tu r e ,

app roache s the se extre mes all for ms of vital actio n g rad uall y o r

suddenly ce ase The li mit s a re so va riable that it is no t at

.

present possible to fo rmul ate any exact law which sh all i nclude
all k now n ca se s Fo r i nst ance many orga nis ms a r e killed at
.
,

the free zi ng poi nt of water ( 0

-
but ce r tai n for ms of life
°

h ave with stood a te mpe ratu r e of 8 7 C 1 23 F

°
an d re
.
°

cent expe ri ment s show t hat frogs and rabbits may be ch illed to
an u nexpe c ted degree without fat al re s ults .

The upper li mit is al so i nco nsta nt th o ugh less so than the lowe r
, .

Most organi sms are de stroy ed a t the te m pe r atu r e of boi ling

wate r but the spores of bacte r ia h ave bee n e xposed to
a much higher te mpe rat ur e w ithout de str uction ( 1 2 0 —1 2 5 ° °

A s a rule p r otopl as m is killed b a te mpe ratu re v a r i ng f r o m

, y y
4 0 to 5 0 C
°
the i mmedi ate c ause of dea th bei ng appa re ntl y
°

,
.

due to a sudden coagul atio n ( p 3 6) of ce rtai n sub s ta nces in the

.

pr otoplasm Thu s if a br ai nle ss frog be gr aduall y h ea ted ,

.
,
 P R OTOPL A S M A N D P H Y S I CA L A G E N T8 . 9

death e nsue s at abou t 4 0 C and the bod y become s sti ff and °

.
,

r igid (r ig
'

or ca lo r es f
) u
ro r the coagul a t o n of the m u cle sub
'

i s -

st a nce The lo we r for ms of ani mal life ag r ee well with pl ant s

.

”
in the ir fatal te m pe r atu re s which in many cas es lie bet w een ,

40 50 C
° °
a nd .

L astl y , it
ppears to be t ru e th at the r e is a certai n m o st
a

favor able or op ti m um te m pe ratu r e fo r the p r otoplas m of e ach

S pecie s of pla n t and a ni m al this opti mu m diff e ri ng co nsider ably ,

in diff ere nt s pecie s P r obably the highe st li m it occ ur s am o ng

.

the b ir d s whe re the u nifor m te mpe r at ur e of the body may be

,

as high as 4 0 C The lowes t occu rs amo ng the m ari ne pl ant s

°
.

an d a ni mals of the A r ctic seas o r of g r eat depth s w he r e the , ,

te mpe ratu r e seldo m r ise s mo r e tha n a degree o r two above the

fr ee z i ng poi nt B etwee n these li mit s there appea r s to be great
-
.

va r iatio n , but 3 5 C may pe r h ap s be take n as the ave rag

°
e op.

Mois ture . P ro to pl as m
t i a l arg e a mo unt o f wa t er o f
al ways co n a ns ,

wh i c h i ndeed t h e l i fel ess o f l i v i ngth i ng s c h i efl y co nsi s t s

po r t i o n ( Se .

ta b l e o n p . A ll pl ant s a nd a ni ma l s a re bel i e v ed to be k i ll ed b y co m

pl eta d ryi ng th oug h s o me o f th e s i mpl e r fo r ms res i s t pa r t i a l dryi ngf or a

,

l o ngt i me beco mi ngq ui es cent a nd re v i v i ngaga i n wh en mois t ened so me

, ,

ti mes ev en a f te r t h e l a pse o f yea rs Hence wa t er a ppea rs to be a n essen .

t i a l co ns t i t uent o f pro to pl as m a lth o ugh as in th e cas e o f mi nera l ma tt ers

, , ,

we do no t k no w th e na t ure o f its co nnec t i o n wi th t h e o th er el ement s or

co mpo unds pr ese nt .

E lectric ity I t h as been sh o wn th a t many f o r ms o f vi t al ac t i on are a o

.

compa ni ed by el ec t ri ca l dis t ur b a nces in t h e p roto pl as m I t is th erefo re .

no t s urpri s i ngth a t th e a ppl i c a t i o n o f el ec t ri c i t y to l i v i ng p ro t opl as m s h o ul d

h a v e a mark ed effec t o n its ac t i o ns I f th e s t i mul us be v ery s l ight pro t o .
,

pl asmi c mo v ement s are f a v o red C o l or l ess b l ood co rpuscl es creep more -

ac t i v el y a nd c i l i a ry a c t i o n i nc reas es in v i g
,
or S t ro ng er s h o c k s c a use a .

s pas modi c co n t r ac t i o n o f t h e pr o t o pl as m ( teta nus) f ro m wh i c h it ma y o r ,

may not be ab l e to r ec o v er acco rd i ngto th e s t reng th o f th e s h oc k

,
.

Po is ons To wa rds cer t a i n ag

. e nts pro t o pl as m is i ndi ff e ren t or seemi ng ly
s o b ut t o wa rds o th ers it be h a v es in a very r ema r k ab l e ma nner
,
Th e ma t .

ters k no wn as po i s o ns modi f y or des t ro y its ac t i v i t y as is well k no wn f ro m ,

t h e fa mil i a r effec t s o f a rs eni c o pi um etc Disease may a l so i nte r fere wi th

, , .

its no r ma l ac t i v i t y ; b ut th e co ns i dera t i on o f th es e ph ases o f th e s ubj ect

bel o ng s to th e mo re exclus iv elv med i ca l s ci ences s uch as t o x i co l og y and ,

pa th o l o g y .

Other Ph ys ica l A g ents The mo re h i g h l y spec i al i zed fo rms of prot o

.

pl as m a re affec t ed by a g rea t va ri et y of ph ysica l agents s uc h as l igh t , ,

40 PR O TOPL A S M A N D THE CE L L .

u n d r ess u r e e t c a n d u po n th i s s u sce p t i b i l i t y d e pe nd ma ny o f th e
so , p ,
.
,

i h r m i f t t i f l i f F i t w a v es o f l ig h t or o f so und
h ge a n e s a o n s o . e o r n s , a nc e ,

t i i l r t l m i t r t r in th e ey e a nd ear ca ll f o r t h
ac n gp u o n s pe c a p o op a s c s u c u es ,

a c t io ns wh i c h ult i ma t el y res ul t in t he se
nsa t io ns o f s ig h t and h ea ri ng
.

S mi i l a r co n s i d e r a t i o n s a pp yl t o t h e se n s es o f s,
m ell t as te a nd to uc h ; b ut
,

th e dis cuss i on o f all th es e spec ia l modes o f pro t opl as mi c ac t io n mus t be

deferred .E noug h h as been sa id to s h ow th a t l i v i ngo rganis ms ( t ha t is ,

th e protopl as m wh ic h is th eir ess ent i a l par t ) are a b l e to res po nd to ma ny

i nfl uences proceedingfrom th e wo r l d in wh ic h th ey l i ve Upo n th is prop .

erty depe nd th e i nt i ma t e rel a t i o ns bet wee n t h e o rg a nis m a nd its enviro n

ment a nd th e po wer o f adapt ab i l i t y to t h e enviro nment wh ic h is one o f th e

,

mos t ma r vell ous and c h a ract eris t i c propert i es o f l i v i ngt h i ng . s

N on d ifl us ibility L i v i ngpro t opl as m l i k e mos t o f th e v a ri ous pro teid

'

-
.
,

ma tt ers wh ic h it yi el ds ( p . is i nd ifl us ible. I t wi ll be seen event uall y

t h at os mo t ic processes pl ay a l ea di ngrole in th e l ives o f pl a nts a nd ani mals ,

s i nce th ey are in l a rg e pa r t th e means b y wh i c h nut ri ment is co nv eyed to

t he l i v i ngs ubs ta nce . I n vi ew o f th i s fac t the no n di ff us i b i l i ty o f proto

,
-

l as m as w e ll as o f o rdi na ry pro t ei ds is a f a ct o f muc h s i g ni fica nce

p .

Veg et a l and A nn ual Protop las m . The pro t o pl as m o f pl ants is es

s entially i dent ica l wi th th at o f a ni ma l s in c h emi ca l a nd ph ys i ca l rel a t i o ns ,

a nd mani fes t s t h e s ame f unda menta l v i ta l pr o pe r t i es . B ut it wo ul d ma ni

f es tly b e a bsurd to suppo se th is i denti t y a bso l ut e f or if it we re so pla nts
, ,

a nd a ni mal s woul d a l so be i dent i ca l ; a nd f urth er mo r e , th e pro to plas m

o f ev ery s peci es o f pl a nt a nd a ni ma l mus t di ff e r mo re or l ess f ro m t he

pro t opl as m o f ev ery o th er s peci es . Wh at is mean t is t hat th e diff erences

bet ween th e ma ny k i nds o f pro to pl as m are f ar l ess i mpo rtant t han the
f undament al resemb l ances whi ch underlie th em .
 C HA P T E R I V

THE B I O L OG Y OF A N A N I MA L .

Th e C o m m o n E a rt h w o rm .

( L umbricus terres tris L i nn us )

, ae .

WE n ow advan ce to a m ore p r ecise exa mi natio n of the livi ng

bod y co nside r ed as an i ndividu al I t is a fam ilia r fac t that
.

livi ng t h i ngs fall i nto two great grou ps k nown as pl a nt s and ,

anim al s. We shall the r efore exa mi ne a r ep r e se ntative of each

o f these gr an d divi sio ns of the livi ng world and i nqui r e how ,

they r ese m ble each othe r and how they differ A ny highe r .

a ni m al wo uld se rve as a t y pe bu t the co mm o n ea r thwor m is a

,

peculiarl y favo r able object of st ud y because of the si mplicity of

,

its st r uctu re the clear ness of its r el atio n to othe r an i mals st an d

,

i nga bove an d below it in the scale of o r ga n i z atio n and the ease ,

with which it may be procu red and di ssected E ar thwor ms of .

,

w hi ch the r e a r e many ki nd s are fou n d in all pa rts of the w orld

, ,

e xte ndi ng eve n to i solate d ocea nic i sla n d s I n th e U nited S tate s .

the r e are seve r al s peci es of w hich the mo st co mmo n are L

, .

co mm un is ( A llo lo bop fiora m uc osa E ise n) L terrestria and L

, , , .

aetidus ( A llo lop ob/wm f e ntida E ise n)

f The
,
fir s t two of the s e
.

a r e fou nd in the soil of ga r de ns etc L terres tris bei ng th e

,
.
,

large r and stouter s pecie s and r ea dil y di sti ngui sh able by th e

flatte ned sh ape of the po ste r io r r eg io n L f aetzdus a smalle r
'

,
.

red species ,
transversel y striped and havi ng a characte ri s tic
,

odo r occu r s in an d about co m po s t heaps

,
-

Mode of L if e etc ,
E a rth wo r ms live in the ea r th bu rr ow
.
,

ingth r ough the soil at a depth va ryi ng f r o m a few i nche s to

s eve r al feet .H e r e they pass the d ay ti me c r awli ng out at ,

n ight or afte r a showe r The bu r r o w s p r oceed at fi r st straight

.

do wn war ds and the n wi nd about ir r eg ularly so meti m e s r each

, ,

41
42 THE B IOL O G Y OF A N A N I MA L .

ing a depth of s ix or eight feet The ea r th w o r m is a n octu r nal

.

an m
i al and d u ri ng the d a
y lies q uiet i n it s bu r r ow n ea r the sur
,

face extended at fu ll le ngt h head uppe r m ost A t ni ght it

,
,
.

becomes ver y active and thrusti ng the fore end of the body
, ,

explore the vici it in ll di r ec t io n s though still c li ngin g

fa or u t ,
s n y a ,

fast as a r ule to the m outh of the bur r ow by the hi nde r e nd

, ,
.

I n this way the wor m is able to fo r age sei z i ng leave s pebbles , , ,

and othe r sm all object s and draggi ng the m i n to the bu r row

,
.

S o me of the se are devo ur ed ; the re mai nder ( i ncluding the peb

bles etc ) are used to line the uppe r pa r t of the bu rro w a nd to
, . ,

plug up its ope ning when the wor m r e ti r es fo r the day B e .

sides bit s of le ave s and ani mal matte r e arthwor ms s wallow l a rge ,

quantities of earth which is pass ed slo wly th r ough the ali me nta r y
'

ca nal so that any nutriti o us subst ances co ntai ned in it m ay be

,

dige sted and ab sorbed This earth is ge ne r all y s wallowed at a

.

conside rable di sta nce below the sur face of the gr ou nd and is ,

finall y voided at the surface ne a r the ope n i ng of the bu rr ow .

I n this w ay ari se the small piles of e ar th ca s t ing s o r f woes )

which every o ne has see n e speci ally in the mo r ni ng whe r eve r , ,

earthwor ms abound Ve ry l arge qua nti t ies of ea rth are thu s

—lll so me ca se s acco r d
.

brought to the surface by ,

ingto Da rwi n s e sti mate s m o r e th an eightee n to ns pe r ac re in

’
,

a si ngle year I n fact m o st soil s are co nti nuall y bei ng worked

.
,

ove r by wor ms ; and Dar wi n h as sho wn th at the se hu mbl e

creature s in the c ourse of ce ntu rie s have helped to bu ry hu ge
, ,

rock s an d the rui ns of ancie nt b uildi ngs *

.

The ea rth worm h as no ea rs ey e s o r any other w ell ma r ked -

organs of speci al se nse N eve rthele ss—and thi s is a poi nt of

, ,

gre at physiological i ntere st —

.

the fore end of the body is se ns i

tive to light ; for if a s tro ng light be sudde nl y flas hed u po n this
p art of the wor m as it lie s stretched fo rth it w ill ofte n dash ,

i
l ke a r bbit i nto its bu rrow
a ”
The ani mal has a kee n se nse of.

touch as may be proved by tickli ng it ; and its se nse of taste

,

must be well developed si nce the wo r m is so me wha t fas tidiou s,

in its choice of food E arth wor ms appe ar to be q uite deaf but

.

possess a disti nct though feeble se nse of s mell

, , .

Da w in Veg eta ble Mo uld a nd E a r th worm l t

r
a l so W h i te s
, . A pp e on, N . Y 1 882 . Se e
N a tur al His tory of Selbo r ne I ndex
’

, , r fr
e e ence s to Ea rth w rmo .
”
 A N TE R O POS TE R I OR -
DIFFER EN1 I A TION . 43

G E N E R A L MO R P H O L OG Y .

A
tte ntio n will fi r st be directed to ce r t ai n featu r es of the
B O D Y see mi ngl O f little i m port an c e but really full of m e ani ng
y ,

whe n co m pa red w ith like fe atu re s in oth er

animals higher or lo we r in the sc ale of
o r gan i zatio n .

A nt er o post er i or Di ff er enti ati on

-
The .

body ( Fig 2 1 ) has a n elo ngated cy li nd r ic al

.

for m tape ri ng to a bl un t poin t at o ne end

, ,

ob t usel y ro unded a nd fl atte n ed at the other .

A s a r ule the poi nted end m ove s for ,

war d s in loco m oti o n and the m o uth o pe ns ,

n ea r it For the se and other reas o ns

.

th e poi nted end m ight be called the head

en d a nd the othe r the t a il end
,
B ut the -

worm has re all y neithe r he ad no r t ail and ,

he nce the t wo e nd s m ay better be di sti n

g u is h ed as the
f ore en d and th e /cin de r en d
,

o r sti ll bette r as a nterio r a nd p os ter ior .

A nd in scie ntifi c l a ngu age the fact th at the

wo rm h as a nterior a nd po sterior e nd s
whi ch differ fro m each othe r is st ated by
'

sayin g th at it sh o ws a ntero p os terzor clg

’

f er -

en tia tio n Thi s si mple fact acqu i r e s great

.

im po rta nce in the light of co mpar ative

biology ; f o r it may be show n th at the
antero pos te rior dilf e re ntia tio n of the ea rt h
-

wor m ins ig nifica nt as it see ms is o n ly the

, ,
”

begi ni ng of a se rie s of i mporta nt m odifi ca

tio ns exte ndi ng u pw a rd s th ro ugh m ore and
mo re co mplex stage s to c ul mi nate in man
h i mself .

FI G 21 —E nl r gd vi w f t h
. . a n t ri r d p t ri r
e e o e a e o an os e o

p t f t h b d y f n r th w rm
ar s o e o fr m th o a ea o as see n o e
v t l p t m u ; li t ll um ; gp gl d ul r
en ra as ec . a an s e, c e . .
, an a

pr mi o n th 26t h m i t ; m m u th ;
n e n c es o d xt r e so e , o o .
, e e

n l p
a i g f th id u t ; p p r t m i um ; t ;
O en n s o e ov e s a , os o 8 , s e ee

sa x p ni g f th
, o e mi
n lr pt
s o l ; d xt r le se na ec e a c es s . .
, e e na
o p i g f t h p r m d u t Th f rm f th b d y
en n s o e s e -
e s . e o o e o

v i gr tl y i lif
a r es ead i gt t h t t f p
n e ac c o r n o e s a e o ex an

s iO n Th S p
. i m h r h w i fr m
e ec l h li en e e s o n s o a n a co o c

pr p r t i
e a ( S li g
a ht ly nl gd )
on . e ar e .
44 THE B IOLOG Y OF A N A N I M A L .
.

Dorso v ent r a l D
-
i ff er e n tiati on I n li vi n g or w ell
.
prese r v ed spe -

the bod is t perfectl c li n d r ic al b ut i s s o m e wh at

e imens y y
n o
, y ,

fl attened pa r ticul,
a rl y n e a r the po s terio r
pri smatic four sided for m O ne of the fl atte ned si de s sl i ghtl y
end an d h as a sl ghtl y
i ,

,
-
.

d arker in color th an the other is habit uall y tu rn e d up war d s and , ,

therefore c lled the b ck the oppo site o r lo w er s ide co m m o nl y

is a a , ,

dow w ard bei g the bell For the sake O f acc u rac
t ur ned n s n , y .
y ,

however biologi st s are w ont to spe ak of the dorsal a sp ect (back )

,

bell of the bod d the fact th at an a ni mal

a nd v en tr a l asp ec t ( y) y; a n

h as a back a n d bell y d i ff e ri n g f r o m e a ch other in s t ructu re o r

f unction or both as in the earth wo rm is expressed by say i ng

, , ,

th at the body exhi bits do me v en tra l dif eren tefa tiou T hi s l ike
-
.
,

a ntero po ste rior differe ntiatio n is ve ry feebl y exp ressed in the

-

exter nal features though cle arly marked in the a rra nge me nt of
,

the inter nal p art s of the e a rthwor m I n higher ani mals it .

becom e s one of the most co nspi c uou s fe atu re s of the body .

B i lat eral Sy mmet ry Whe n the b o dy is placed in the nat ural

.

position w ith the ve ntr al as pect do w nwar d s a ver tica l pl ane

, ,

passi ng lo ngitudi nall y through the m iddle w ill divide it i nto

exactly si mil ar right and left halve s Thi s si mil a rity is called .

two sided like ne ss or bila tera l sy mmetry

-

,
Tho ugh not very .

O bvious ext er nall y thi s sym metr y char acteri z es the a r ra nge me nt
,

of all the i nter nal part s ; and it m ay be gr ad ually t raced up

w ards in higher ani mal s u ntil it beco mes as striki ng and pe rfect
,

as in the hu man body .

Thus a very superfici al exami natio n r ev eal s in the earth

wor m two fundame nt al l aws O f o rga ni zatio n v iz dif eremzw
'

.
, ,

tion or the law of differe nce and s y m me try o r the law O f like ,

n e ss . A nd the se l aws are of i ntere st for the r eas o n a mo ng

many oth ers th at e arth wor ms like other orga ni s ms have as a , ,

r ace h ad a hi stor y h ave come to be by a gr ad ual process (of

, .

p . A nd biol ogy m ust stri v e to answe r the q ue stio n s [lo w and

why certai n p art s h ave beco me s mmetri ca l and othe rs difl er
y
entiated Without e nte r i ng i nto a f ull di sc u ss io n of the q ue s
.

tion at this poi nt it may be said th at the mai n ca use o f sy m

,

metr y or d ifferenti atio n has probabl bee n like n e ss o r u nli ke n

y
of function or of r elation to the e n vi r onm ent E a r th w o r ms
, .

show antero po sterior and dor so ve ntr al diffe r e ntiation becau se

-
-

the anterior and posterio r ext re mities o r the do r sa l and ve ntral ,

 ME TA ME R I S M . 45

a spects have bee n differentl y used and exposed to di ffe r ent con
,

ditio ns of e n viro nm e nt A nd o n th e othe r han d the organism is

.

b il ate r ally sym m et r ic al becau se the two side s h a ve bee n si mila r l

,
y
u sed and have bee n expo sed to like co n ditio ns of e n viro nme nt .

Metamer i sm A nothe r gen eral fe atu re of the earth w o r m is

.

O f gr e a t i m porta nce in vie w O f the co n ditio n s exi sti ng in other

a n i mal s i ncludi ng the highe r for m s

,
The body is m arke d off .

by transverse grooves i nto a serie s of si m ila r part s like the joi nts
O f a b am boo fi shi ng rod or like the joi n t s of fi nge r s ( Fig
-

, .

The se p a rt s are c alled meta meres o r m ore ofte n s omi tes and , ,

the bod y is co nseque ntl y sai d to h ave a meta memc st ructure o r

'

to exhibit m eta meris m F r o m the out side the so mite s appea r to

.
,

be produced si mpl y by r egul a r folds in the skin like the ,

w ri nkle s betwee n the joi nt s O f ou r fi nge r s B ut as the wri nkles .

o f the fi nge r s a r e o nl the e xte r nal exp r e ssio n of a m ore f un d a

y
m e ntal joi nted str uct ur e withi n so the exter nal fold s sep ar ati ng
,

the so mite s r epre se nt an i nter n al divi sio n i nto succes sive p a rt s

, ,

which a ff ect s all the organs of the body and is a result of so m e ,

o f the m ost i m portan t phe n o me n a of develop m e nt .

Th e ex pl anati on o f met amerism or s eria l sy mmetry is o ne of th e

mos t di ffi cul t pro b l ems o f mo rph o l o g y B ut it wi ll be seen f a rth er o n th at
.

meta meris m so c l ea r l y a nd s impl y ex pressed in t h e ear th wo r m c a n b e

, ,

t r ac ed upwa rd in ev er i nc rea si ngco mpl ex i t y to t h e h ig

-
h es t f o r ms o f l i fe ,

a nd s ug ges ts s o me o f th e mo s t i n t eres t i nga nd f undament al pro b l ems wi th

wh i c h bio l o g y
— a nd es pec i a ll y m o rph o l o —
gy h as to dea l I ndeed th e .
,

c o mpa r a t i v e s t udy o f t h e a na t o my o f mo s t h i g h er a ni mal s consi s t s very

l a rgel y in t raci ngout the ma ni fol d t ra ns for mat i o ns o f th eir c ompl i ca t ed
s o mi t es wh i c h u nder many di s g
,
ui s es ca n b e r ecog nized as f unda men t a ll y

li k e t h e s i mpl er so mi t es o f the ea r th wor m .

The so mites diffe r co nside r abl y

Modi fi cati ons of th e Somit es .

in dif fe r e nt pa r ts of the body The extre m e a nte r io r end is .

fo rm ed by a s m oothl y r ou nded k n ob called the p ros to miu m

-

w h ich is show n by its m ode of develop m e nt not to be a tr ue

so mite I t fo r ms a ki nd of ove r hangi ng uppe r lip to the mouth
.
,

which lies just behi n d it on the ve ntr al aspect B ehi n d the .

m outh is the fi rst s o m ite in the fo r m of a ri ng *

,
i nte rr upted ,

above by a b ackw ar d p r olongatio n of the pro sto m iu m .

In beri ngth e so mi t
n um es th e p r t os o mi um m us t v r be r k
ne e ec o ned, th e
fi r t so m i t b i ngbehind the m
s e e ou th .
 THE B IO L OG Y OF A N ANI MA L
46
.

The so mite s fro m the 1 st to the 2 7 th are r ather broad ,

i cre e i n s i e A v a ri a ble n u m be r o f t h e
grad ll
‘

n as z
an d u a y
.

o ite l i g betwee the 7 th a n d 1 9 t h are ofte n s wolle n o n

s m s y n n
eno us g
the ventr al side for mi ng the
,
so c a lled c ap s u lo g la nds - .

the 2 8 t h n d 3 5 t h ( the n u m be r a n d po sitio n var y

B et wee n a

i differe t peci m e n s
) the so m ite s are s w oll e n
ing slightl y n n s

bove the ide an d the fold s bet w ee n the m are

a a n d o n s s ,

s c a rcel y
de fi n ed ex c ept
co
o n the
picuou
ve n tr a

gird
l aspect
le c alled
T a ke n to ethe r

the c lztellu m
.

g ,

they for m a broad n s s

,

Fig 2 1 whose function is to secrete the c a p s ule i n w h i ch

( .
,

the eggs are l aid and al so a nutritive m ilk like fl ui d for the use
,
-

The clitellu m i t p r e e t i
of the developi ng e mbr yo s ( s n o .
s n n

B ehi d the clitell u m the s o mites are

i mmatu r e speci mens ) n .

m fo r ided i n cro s ec t io n a n d fl a tte n ed

n arro wer so ewhat u s ss -
-

,
,

do w w rd Thi fl t t e n i n g s o m eti m e s beco m e s

fro m above n a s s a .

very co nspic uous t o w a rd s the po s terior e n d Toward s the ver y .

last they dec re as e in si z e r ather abruptl y and they end in th e ,

a na l s m
o ite whi c h ,
i s perfor a ted b y a vertic a l s lit the a n u s ,

( Fig .2 1 an
) ,
A ll the .
s o m ite s a r e perf o r ated by s m all O pe n i ng s
leadi ng i nto the i nterior of the bod y and fo r mi ng the outlets of ,

num ero us org ans ; the po sitio n o f the s e o pe n i n g s w ill be de

scribed in t r eati ng of the o rgans E ach so mi t e ex c eptin g the .
,

anteri o r tw o o r th r ee a nd the last ,

give s i nsertio n to fou r gro up s of

s hort and m in u te br is tles o r setce ,

w hi c h a re arr anged in four lo ngi

tudinal row s alo ng the body Two .

of the se ro ws run alo ng the ve nt ral

Fm 22 Di gmm t i llu t t t h
,
_ , a o
aspect s ra e
t w o a r e m ore e
upo n the
,

“ t i “ f th? o w Th d tt d s ide s
e se e e
The set ae exte nd outwards
o e

ut l r pr t th p i t i f
.

o ine e e s en s e os on o

th e t d it
se a mu d
an wh s fro m the i nterior O f the bod y
s es en ,
b ent m th pp t m m “
e O os’ e “ whe r e the are s upplied w ith small
e o 7

mu l ; s c es t ; w b dy w ll
8, se a ,
o -
y a .

m uscle s by w hich they can be

tur ned some wh at eithe r forward s or b ack wa r d s and can also be ,

protr uded or w ithdr awn ( Fig The set as ar e of gr eat use .

in l o co m otio n When pointed b ackward s they support the wor m

.

as it cr aw l s for ward s
; w he n the y a r e t u r n ed for w a rd s the wo r m
c an c reep b ack w ard s _They are of in tere st therefore as repre ,
.
,

se nti ng an extre mel y si mple and pri mitive li mb l ike o r gan -

.
 GE N E R A L PL A N OF THE B OD Y . 47

Plan of th e B ody
The body of the earthwor m ( Fig .
.

l ik e t hat of all hi ghe r an i m al s consi st s of two tube s o ne (a l) , ,

with in the othe r and separated fro m it by a co n siderable sp ac e

o r cavity The inn er tube is the a liment a ry ca na l O pen ,

ingin f r o nt by the mo u th an d behi n d by the a n us the oute r

t ube is the body w all and its cavity is the body ca v ity o r coelom
-

,
-

“f 0 a d 7 1,

FI G .
—A di , a r th w r m
gra m o f n in l ngi t ud i l t i n f th b dy
th e ea o as se e a o na s ec o o e o ,

s h wi n gth t w t ub t h
o e o l m d th d i p i m t B d i gr m f r
es , e c oe o , an e ss e en s .
, a a o c os s

sec tl t
on :l i m t ry t u b :
a . a en a u ; l m ; m m uth C d i gr m e a n, an s caz, c oe o , o .
, a a

s h wi n gth rr ng m n t f m f th p ri i p l r g n m m u th ;
o e a a e e o u ; so e o e nc a o a s : , o a n, a n s

a t
. l i m t ry
a en al ; i di p i m t ; L n d r l bl d v l ;
c a na ( s, nt l s se en s ( i , o sa oo -
es s e v,
’

ve ra or

s ub i t
-
ti l v l ;
n es na ir ul r v l ;
es s e ph i d i
c v .t y rg
. . c
; c a es s e s n, ne r a o r ex c re a r o a ns

c ere br l g ngl i ; g v tr l h i f g gl i ; d vi du t ; d v ry Th
a a a r . .
, en a c a n O an a c . .
, o c o . .
, o a . e

arro w i nd i
s t th u ca e f th i r u l t i f th bl d
e co rs e o e c c a on o e oo .

The coelo m is not however a free co nti nuous space exte ndi ng, ,

f ro m end to end but is di v ided tr ans ver sely by a se rie s of thi n

,

m usc ula r pa rt itio ns the d issep imen ts i nto a se rie s of n e a rl y, ,

closed cham be r s tr aver sed by the ali me n ta ry ca nal E ach com .

p artm en t cor r e spo n d s to o ne so m ite the dissepi me n t s bei n

g ,

Op po site the ext ern al fur row s m e ntioned on p 4 5 A ll th e . .

organ s of the body are origi nally developed fro m the wall s of
these cha mber s an d so m e of the m ,
the organ s O f exc r etion)
project into the c avities of the ch am be r s th at is in to the c oelom ,
.
48 THE B I OLOG Y OF A N A N I MA L .

In the m edian do isel li n e of each so m ite ( exceptin g the fi r st

two or three) is a mi nute po r e ( the dorsa l p ore) which per fo
r ate s the body wall and th us places the coelom in connecti on
-

w ith the exterior *

O ther po res th at pass th r o ugh the bod y
.

wall i nto the c avitie s of var io us o r gans will be descri bed fu r

ans of th e A ni mal B ody

Org The body of . Sy st ems of Orga ns .

the e arthwor m co nsi st s e sse ntially of proto plas m and in o r de r that ,

s o l a rge a mas s of livi ng matte r m ay co nti n ue to ex ist and ca rr y

o n the o r di nary life of an earthwor m it mu st be able to obta in

a sufficie nt suppl y of food ; to dige st and ab so r b it an d dis ,

tribute it to all p ar t s of the bod y ; to build up new p r otoplasm

a nd re m ove w as te I t mu st be se nsi t ive to exte r nal and i nte r nal
.

i nfl uence s ; c ap able of motio n and loco motio n A bove all eac h .

,

p art mu st act w ith refere nce to and in ha r mo ny with eve ry , ,

o ther part s o th at the orga nis m may no t be m e r el y an agg r ega te

,

o f organs b ut o ne bod y ac ti ng as a u nit o r a whole

,
.

The se f u nctions are fulfi ll ed by the O R G A N s r e s pectivel y OF ,

.

A L I M E N TA TI O N ,
DI G E S TI O N ,
A B S O R PTI O N ,
C I R C UL ATI O N ,
E X C R E TI O N ,

S E N S A TI ON ,
M O TI O N ,
the se minis ter to
an d C O OR DI N A TI O N . A ll O f
the welfare of the i ndivid ual The R E P R O DUCTI V E fu nctio n o n .
,

the othe r hand and its co r re s po ndi ng orga ns se r ve to pe rpet

,

uate the specie s thu s mi nisteri ng rathe r to the r ace than to the
,

individu al .

S ets of o rgans devoted to t h e sa me fu nction co nsti tute sys

tems ; as the a limen ta ry sy s te m the c irc ula tory sy s tem etc , , .

Those which are mo re i mmedi atel y co nc er ned w ith the i ncom e

a n d o utgo O f matter— n amel the ali me n ta r digestive ab so r
y y p , , ,

tive c irc ul at o r y and excreto ry sy ste ms are so metim e s ca lled the

, ,
-

g
e e eta tive s s tems or s s tems
y y f n utritio n while those which
o
h ave to do more i m m diatel with the r el atio n of the b o d to e
y y
its e nviro nme nt rather th an the in dividual it s elf are ca lled
, ,

tems o f rela tion E x ample s of t he l a tte r a re the s s te ms of

y
.

o rga ns of suppo rt m otio n i ncl udi ng loco motio n se tio

( ,
) nsa n an d , ,
c o ordi nation
; and eve n the reproductive sy s te m as r el atin g chiefly ,

to othe r i ndivid ual s fi nd s a pl ace he re , .

Ifl ivi ngw rms be i rri t t ed th y w i ll f t n x t r ud

o a e O e e e a mi lk y fl ui d f ro m
t h ese por es ,b ut th use f th l tte r is n t w el l und rs to od
e o e a o e .
 A LI ME N TA R Y s rs rrm
’
. 49

A . S YS TE M S OF N UTR I TI V E ORGA N S H E I R SP E CI A L MOR

T

P H O L OG Y AN D P H Y S I O L OG Y .
( For B see p .

A li ment ary Sy st em ( Or g
wo rms
ans of A limentati on) . E art h -

feed m ain l y u po n leaves o r decay in g vegetable matte r but ,

will also eage rly devour meat fat and other an i mal sub , ,

stance s They also swallow large quantities Of ea rth from

.

which they ext ract not o nly any o rg a nic mate rial s that it m ay
co ntai n but p robably also moistu re and a small amou nt of v a ri
,

o us salts The m ost e sse ntial and cha racte ristic p art of thei r
.

food is de rived fro m vegetal o r ani mal matte r in the fo rm of

va rious orga n ic com pounds of which the most im po rta nt are ,

p roteids ( protoplas m albu m e n ca rbohy dra tes

, ( s tarch , ,

ce ll ul ose ) and f a ts These mate rial s a re u sed by the ani mal in

,
.

the m anufactu r e of new p r oto plas m to take the pl ace of that

which has bee n used up I t is howeve r impossible fo r the ani .
, ,

mal to b uild the s e mate rials directly i n to the s ub sta nce of its
own bod y They mu st fi rs t u nde rgo ce rt ai n p reparatory che m i
.

cal cha nges k no w n collectivel y as dig es tio n

; and o n l y afte r the
co mpletio n of th is proce ss can all the food be ab sorbed into the
c ircul atio n Fo r this p ur pose the food is take n not i nto th e
.

bod y prope r but in to a kind of tub ula r che mical labo r ato ry
,

c alled the a lime nta ry ca na l th r ou gh which it slo w l pas ses

y
bei ng subjected m e anw hile to the actio n Of ce rt ai n che m ical sub
sta nce s o r r e age nt s k no wn as dig
,
es tive
f ermen ts
,
The se sub .

sta nce s which a r e di ss olved in a wate ry liquid to form the dig

,
es

t/ r
ve fl uid are sec r eted by the w alls of the ali me nta ry tube
,
.

T hr ough thei r actio n the solid po r tion s are liquefied and the foo d
is r e nde r ed c a p able of abso rptio n in to the proper body .

The alim e nta ry ca nal is divi sible i nto several differe ntl y c on
st r ucted po rtions play i ng differe nt pa rts in the proce ss of alimen
tation .G oi ng b ackwa rds f ro m the mouth these a r e as follow s
1 The p ha ry nx ( Fig 2 4 p h) an elo n gated b a rr el shape d
. .
, ,
-

pouch ext e nding to about the 6th so m ite I ts w all s are thic k .

and m uscular an d fro m their coelo m ic su rface num e r ou s s m al l

,

muscles radiate on eve ry s ide to the body wall Wh e n these -

.

m u scles con t ract the cavity of the pha ry n x is expanded ; and if

,

the m onth has bee n p rev iou sly applied to any solid object such ,

as a leaf o r pebble the pharyn x acts upo n it like a suctio n pu mp

,
-
.
FI G . 24 .
—
D r
o s al vi
t ri r p rt f th b d y f L umb i u
ew o f th e an e i t ppe r
o a o e o o r c s , as a a s
wh l i d p n l ng t h d r l
en a o e
p t a o rt i r h ; r p ;
e o sa br l as ec . a o, ao c a c c, c o c ere a
g ngli ; o l
a l if r u gl nd ; d d i p i m n t ; d d r l v l ; a gi zz d ;
a c ; ca c e o s a s , sse e . c, o sa es se ,
'

ar
u ; p h p h ry x ; p t m i um ; t t m h in t t in Sh w i g
pr
w ph g
, oeso a s a n a, os o a s o ac es o n
,
,
-
e.
th l t r lp u h
e a e a m in l ro
pt
c les ; s r, se
; .
th th r e a f ece a c es e e O
l t r l m in l v i l
a e a se a es c es .
 OR G A N S OF A L I M E N TA TI ON 51
'
.

I n thi s way the a ni mal l ay s hold of the v ario us O bject s nut ri ,

tious and other w i se which it devou r s o r d raw s i nto its b urr ow

, .

E m be dded in the muscular wall s of the phar n x are a

y
“ ”
n u m be r of small sa liv a r gla n ds of w hose fu nctio n n othi ng
y
is defi nitel y k now n though they doubtless pou r a digestive fluid
,

in to the phary nge al cavity .

2 The (esop ha g
. us ( as ) a sle nder thi n walled t ube exte ndi ng
, ,
-

f ro m the 6 th to the 1 5 th so mite Th r o ugh this the food is .

s w allo w ed bei ng d r ive n slo wl y alo ng by w a velike (p eris ta ltic)

,

c o n t rac tio ns ( p . I n the r egio n of the 1 1 th and 1 2 th

s o m i tes are th r ee pairs of s mall pouche s O pe ni ng at the s ide s of

the Oeso pha gu s The se are the ca lcif ero us g

. la nds They
c o nta i n solid masse s of calciu m c a rbo nate an d Da r w i n co njee
,

tu re s that thei r use is pa rtl y to aid dige stio n by ne ut rali z ing the
acids ge ne ra ted du r ing the digestio n of leave s and perhaps ,

pa rtly to se rve as an outlet fo r the exce ss of li me in the body ,

e speciall y whe n w o rms live in calca r eou s soil .

3 The crop about the 1 6 th s o m ite

.
; a thi n walled sac -

lik e d ila tatio n O f the ali m e ntary ca nal which se rve s as a re s e r ,

v oi r to r eceive the s wallo wed food .

4 . The g izz a rd a bo ut the 1 7 th s o mite

; a c y lin d r ic al
fi rm and muscula r po r tio n li ned by a ho rny me m bra ne
, I n thi s .

t h e food is r olled abo ut squee z ed and grou n d to p repare it fo r

,

d ige stio n in the follo wi ng po rtio n v iz ,

5 . The s toma ch i n tes tine -

which corre spon d s phy sio
logically to bo th the sto mach and i nte sti ne of higher a n i mal s .

This is a st ra ig ht thi n walled tube exte ndi ng fro m the gi z z a rd

-

to the a nus w ithout con volutio ns no t diff ere nti ated i nto stomach
, ,

a n d intestin e and devoid of di s ti nct gl an d ul a r appe nd age s such

,

as the live r or pa nc re as exi sti ng in the h igher a ni mals The .

digesti v e fluid is sec reted by the w all s of the ali me nta ry ca nal
i tself the su rf ace of which is m uch i ncrea sed by the pre se nce of
,

late ral pouche s or dive rticula o ne o n eithe r side in e ach so m ite

, .

I n fro n t the se are l a rge and co nspicuou s but beh ind they g radu ,

a ll
y di m i n i sh in s i z e u ntil sca rcel y perceptible .

i nner sur face o f th e s t omac h i ntes t ine is f urt h er i ncreas ed by a

Th e -

deep i nwa rd fo l d call ed th e typ h losole r unni ngl o ng

,
i t udi na ll y al o ngth e
,

do rsa l medi a n l i ne The t yphl osol e is not vi si b l e o n th e ex t eri or but is

.
,

s een b y openi ngth e s to mac h i ntes ti ne f ro m th e s i de or bel o w or upon

-

,
52 THE B I OL O G Y OF A N A N I MA L .

m a k i ng a c r oss s ec-
t i on I t is
. ri c hl y s upp l i ed wi th blood vess els th a t pass -

i t i v i t f r m t h d r l v l ( Fig nd its mai n f unc

dow n n o ts ca y o e o sa es s e a .

tion is probab l y to i nc rease the sur face f or the a bso rpt ion of food ( c f the .

spira l v a l ve in th e i n t es t i ne o f s h ark s ) .

The outer s ur face o f th e s tomac h i nt es t i ne is covered wi th pi g mented

-

e ll o w i s h b r o wn c hl o rag og ue cell s -
Th ese were fo r mer l y s upposed to be
”
y
- .

c oncer ned wi th th e sec ret i o n of th e dig es t i v e fl ui d a nd h ence a re o f te n ,

ca edll “ h
epa t i c cell s Th is h o wev er is pro bab l y a n erro neo us i nte rpreta
.
”
, ,

t i on and th ey are now beli eved to be concerned wi t h the proc ess of excre
,

tion ( p .

D ig
estion Dige stion
begi ns eve n befo r e the food is tak e n
.

in to the ali me nta ry ca nal ; before bei ng sw allowed the leav es , ,

etc are moi ste ned by dige stive flui d po u red out f rom the
.
,

mouth s of the wo r ms The mai n action howeve r doubtless goes

.
, ,

o n in the a nte r i o r pa rt of the sto mach in te sti ne and d im in ishes -

as the food passe s back wa rd I t has bee n p roved by expe rime nt .

th at the dige stive fluid act s on a t least two of the th r ee p ri ncipal

v arietie s O f organic food stuffs v iz on proteid s and o n sta rch
-

, .
,

r re se mbles the pa nc reatic fl uid of

( c a rboh y drate ) a n d in so f a ,

higher ani mal s which it further re se mble s in havi ng an alkaline

,

reactio n A nalogy lead s us to believe th at the digestive fl uid

.

h as so me actio n al so on fat s ; b ut thi s has no t bee n proved .

K u r k enbergand Fré dericq h a ve s h o wn th at the diges t i ve fluid of the

ea r th wo r m co nt a i ns a t l eas t th ree ferments a nd a cco rd i ngto th e fo r mer
auth or th es e occ ur o nl y in th e s t o mac h i nt es t i ne Th ey are a s fo ll o ws
-

1 P ep tic f erment wh i c h has th e ro er t in a n a c i d medi um o f co n

.
, p p y
vert i ngpro t eids int o sol ubl e a nd di ff usi bl e p ep ton es ; th i s is th erefo re
a na l og o us to th e pe s i n o f th e as t ri c j ui ce in h i h er fo rms
p g g .

2 Tryp ti c f ermen t h a v i nga s i mi l a r ac t i o n o n ro tei ds b ut o nl in an

'

p y
.
,

a l k al i ne medi um—h ence a na l o o us to th e t r

,

g yp s i n o f pa nc rea t i c j ui ce .

3 Dia s ta tic f erment wh i c h c onv ert s s t a r c h i nto l ucose rape s ug

g (g a r)
in an al k al i ne medi um—h ence ana l o o us to the t al i n o f s a l i v
.
, -

g p y a a nd the .

amy l ol yt ic fer ment o f

panc reat ic j ui ce .

A bsorp tion The fe rme nt s of the digestive fl uid co nve rt the

.

solid proteid s i nto soluble and d iff u sible pepto nes the s ta rchy ,
matters i nto sugar gl uc o se
( ) These products d issolve in the .

liquids p rese nt and a re the n gradually absorbed by the w alls of

the I ntesti ne as the food passe s alo ng the alim e nta ry ca nal The .

precise mechanism of absorptio n is not y et tho roughl y unde rstood ,

but t is probable that much of the nut ri me nt passes by diff u sio n

I

( osmosi s) into the wall s of the stomach i nte st ine and the nce into -
 OR G A N S OF CI R C UL A TI ON . 53

the blood for dis tributio n to all p art s O f the bod y The refuse .

re mai ni ng in the ali me nta ry ca nal (and which has n ever bee n a
p art of the bod y proper) is fi nally voided through the anus as
c as ting m
s or
f ceces Thi.s proce ss of def aecatio n u st n o t be

co nfou n ded with that O f ea cretion which will be de sc ribed later , .

Cir cu lat ory Sy st em The food h a vi ng bee n ab sorbed is

.
, ,

d ist ri b uted th rougho ut the bod y by tw o device s .

1
. Coelo mic C ircu la tio n The cavity of the owlo m is filled
.

with a c o lorle ss fluid coelo mic fluid w hich m ust be regarded as a

ki nd O f l y m ph or blood B y the co n tractio ns O f the body w all as
.
-

the w o rm c rawl s about the co lo mic fluid is d rive n b ack and fo rth
,
e

th rough all pa rts O f the coelom ,

th rough ir regula r ope ni ngs in the

d isse pi me nt s A s the dige sted
.

food is absorbed fro m the sto mach

i n test ine a co ns ide rable pa rt of it is
believed to pass i nto the coelo m ic
fluid and is thus co nvey ed d irectly
,

to the orga ns which thi s fluid

bathe s The c oelo mic flu id is com
.

po sed of two co nst itue nt s v iz a ,

.
,

colorle ss fluid c alled the p la s ma ,

an d colorle ss i sol ated cell s or co r

u s cles w hich float in the plas m a

p ,

a nd are re markable fo r the fact

th at they u ndergo co nst ant though
slow cha nge s o f for m I n fa ct they
.
3
clo sely re se mble ce rta m k i nd s of f m th ro e co g

A mcebce a nd w e should cert ai nl y l m i fl u i d f th th w rm A o c o e ea r o .

.
,
l m ti n f h f t
ffm u d i g f i igfi fifdfi ;
a o e ra o e
co nsi der the m to be such I f we S n n a o e

fou nd the m occu r ri ng free in st ag i n gl Ph g y t w i th v u l

s e a oc e, ac o es.

( A f t r M t h ik ff ) e e sc n o
n an t w ater We k no w ho wever
.

.
,

that they live o nl y in the p las ma and have a co mm on origi n ,

with the othe r cell s of the body ; he nce we mu st r egard them

no t as in di vidual a ni mal s but as co nstit ue nt cell s of the e a rth
,

wo rm The coelom ic fl uid is in fact a ki nd of tis s ue consi sti ng

.

of iso lated colo rl ess cell s fl oat ing in a fluid i n te r cellul ar sub st ance .

These free flo ati ng cell s are prob abl y the scavenger s ( p hag ocy tes )

of the bod y devo uri ng and de str oyi ng waste matters S o m e

, .
 B I OL O G Y OF A N ANI MA L
THE
.

in v a di n g p a r as ite s s u ch as b a cteri a .

su ppo s e th a t the y a l s o a tt a c k
cu l i B e ide s the c o elo m ic circulati on
2 V a s c u la r C i r a t o n s .

m co m plic ated c i r c u lator y app a r atu s c on

there is another a n d ore
t be the b lo od c s e s e l s which for m a c o m
f br hi ng u s
-

,
s i sti g n
n O a c ,

r m if i g thro u gho u t the bod y Thro u gh these

pli c ated sy stem n
.

a y
t ube s is dri v e n a red fl uid a n a logo u s to the r ed blood of h igher
i l like it co i ti n g of p l a s m a a n d c o r
p u s c le s the l a
tte r
an ma s and
n s s ,
,
pi dle h ped The r e d colo r is
bei ng fl attened a n d s o m ewhat s n s a -
.

due to a sub t ce s an h ae m o g,
lo b in di ss olved in the,
plas m a a n d n o t ( as

c t i ed i the c rp u cle s w hich a re colorle ss

in higher for m ) n n o s .

s o n a ,

e rth or p r ovided with peci a l pu m p i n g

The a w m is n o t a s

organ or heart for the pr o p ul sio n O f the blood s u ch as w e fi nd

,

higher i l pl ce of thi ce r t i n of t h e l a r ge r blood

in an m a s I n .
a s a
”
ve ssel s v iz the “ do rsal ve ss el a n d the a ortic arche s
( .
,
i
h ave mu c l r s u a
co n tract i le w all s w hich propel the
,
bl o od n a c o n

t t directio ave like o tr c t i n th a t run alo ng the

s an n by w c n a o s
-

ve ssel fro m one end to the o ther ( pe ri t ltic “

s a co n tract i o n s cf ,
.

p 5 1 ) at r egul ar interv al s and th u

“ ”
s give ri se to a pulse .

The co ntr ac tile ve ssel s give O ff other n o n co t ra tile t ru ks

n c n -

which divide and subdivide i nto tube s of ext r e mel y s mall calib r e
a d havi g ver thi n w a ll s The ulti m a te b r an ch es kn ow n as
n n y . ,

pe r m a e te e rl all the org n a n d ti ss ue s in which

cap illari es n a a s
, y ,

they for m a close n etw ork The tre s a.

m of bl d fte r pas i g
o o a s n

thro ugh the c apill aries is gathered into suc c ess ively l a rge r vessel s
w chi h a fter a lo n ger or s horter c o u r se fi n a ll y e m pt y i n to th e

origi nal c ont ractile tr unk s and c o mplet e the ci rcuit Thus the .

v ascul ar system is a cl osed syste m O f t ube s and there is r e as o n to ,

believe th at th e bl o od follo ws a pe r fectl y defi nite co u rse tho ugh ,

*
thi s is not yet preci sel y deter m i ned .

We may now co nsider the ar rang e me nt O f th e pri ncipal

trunk s The l arge st of the m which is also the mo st im po rta n t
.
,

of the contr actile ve ssel s is : ,

a The dorsa l ress el ( Fig 2 4

. a l ong m u sc ular tube
.
,

lyin g upon the upper s ide of the ali me nt ar y ca n al I n the liv .

ingw or m it m ay be di sti nctl y see n thro ugh the se mi t ra ns p a re n t -

h ul d b n t d th at in th b n f h rt i t i di ffi ul t to di t i n
It s o e o e e a se ce o a ea s c s

gu h b t
is w n rter i ndee v i“
n a W m
e y m
es r a n v n i ntl y dis t i n
e s .
”
e a o e co e e

gu i h sff r nt v a l rr i
y g
e en b l d to w rd
esse s . t h ”
p i ll r
ca i nd fi nt oo a s e ca a es , a e ere

v s l rry i ngb l d w y fr m t h m
e se s ,
”
ca oo a a o e .
 B L OOD VE S S E L S .
. 55

s k in as a da rk red ba n d -
which is tolerably st raight when the
,

w o r m is exte nded but is m ade z ig z ag by co ntr actio n of the body

, .

I f it be closel y ob s e r ved a sort of wavel ike co ntractio n is ofte n

,

see n r unning fro m behi nd forward s Thi s m ay be very clearly .

o bse r ved in a wo r m s t upefi ed by chlo r ofor m e s peci all y if it has ,

bee n l aid ope n along the dor sa l side The dorsal ve ssel the n .

a ppe a r s as a deep —
red so m ewhat t wisted tube ru nning alo ng the
, ,

uppe r side of the ali me nta ry c an al W avelike contr actio ns .

c o n tin ually s tart f r o m its hin de r en d a nd run r a pidl y for w a rd s ,

o ne afte r a nother to the ante ri o r end where the do rsal ve ssel

, ,

fi nall y b r e aks up o n the ph a ry n x into a la rge n umber of b ran che s

( Fig.

The r e sult of these o rde rly p rogr e ssive co ntractio ns is that

the fl uid withi n the t ube is p ushed for wa rd s —ve ry mu c h as the
fluid in a rubbe r tube is forced alo ng w he n the t ube is stripped
th r ough the fi nger s I t is still better ill ust ra ted by the actio n
.

of the fi nge rs in the O pe ratio n of milk ing Th is action of the .

v essels is a ty pic al ex am ple of p eris ta ltic con tra ction .

b S u b i n tes tina l v es sel

.
-
This is a st raight ve ssel which
.

ru ns alo ng the middle li ne o n the lower side of the ali m e n t a r

y
ca nal pa rallel to the o ne ju s t desc ribed
,
I t r etu r n s to the .

hin der p a rt of the body th e fl ui d which has bee n ca r ried

fo rwa rd s by the do r sal vessel O n the ph ar yn x it bre ak s up .

in to many b ran che s w hich r eceive the fluid f r o m cor r e spo nd ing
,

b ra nches of the dorsal ve ssel .

c . Ci rc ula r or comm is sura l v essels m eta m e r ically r epeated ,

t r unks which run fro m the dor sal ve ssel do wnw ards aroun d the
al ime ntar y canal and ulti matel y co nnect w ith the ventral ve ss el .

They a re of seve ral k ind s of which the m ost impo r tant a r e as

,

follow s
1 . The aortic a rches o r circumcesop liag ea l ves sels ofte n ,
“ ”
k n ow n as heart s s i nce like the dor sa l ve ss el they are co n
,

t ractile and with the l a tte r furn i sh the e ntire p r opulsive fo rce
for the ci r cul atio n These a r e five p airs of la r ge vessels en
.

c irc ulin
g the m p g
S O ha us in s o mite s 7 to 1 1 i n clu s ive Thes e .

vessels pass directl y fro m the do rsal to the ve n t ral vessel givi ng ,

o ff n o b ra n che s D uri ng life they pe rfo rm powe rful pe ris ta ltic

.

cont rac tions r eceivi ng blood fro m the do rsal vessel and pum ping
,

it i n to the sub i n testi nal or ve nt ral

-
.
 THE B I OL OG Y OF A N ANI MA L
56
.
.

i i l l passi ng f ro m the do rsal vesse l

2 . D ors o n tes t na- v ess e s ,

n the regio n of the stoma ch i nte sti ne

to the wall of the g u t i -

in
.

the e ve el the r e are two o r th r ee p a i r s in each so m ite

Of ss s
.

s
cove r ed like the do r sal vessel in thi s regio n)
They are thickly ( ”
chlo ragog e cell that the i r r ed colo r is
with pigmented u s s o -

u suall y not a ppare n t U n li ke the.

aortic arche s the s e vessels

break up on the wall of the inte sti ne i nto capilla ries which ar e
conti nuous with branche s from the ve nt ral vessel .

m l p i g fro m the do r sal vessel

3 D . or so te g u en-
ta /ry v es se s ass n ,

along the di ssepime nt into the body w all o n e ach side Th ese -
.

are small ve ss el s th at pass di rectly arou nd the bod y to co nne c t

with a longitudinal t runk sub neu ral ly i ng below the v en-

tr al n e rve cord (see belo w) an d giving off b ra nches to the body

-

w all di ssepi ment s and neph ri di a

, ,
.

h B l d The p recise cou r se of the blood in

C o u r se o
f t e oo .

L u b m r i cu s is s till in di s p u te though its m o r e ge n e

,
r a l fe atu re s a re

k no w n I t is c e rt ai n th at the bul k of the blood p asse s for ward in

.

the d o rsal vessel dow nward ar o u nd the gut thro ugh the ao rtic ar c he s
,

i nto the ve ntral ve ss e l and the nce b ack wa rds to ward s the po s
,

terior region I ts p ath the nc e i nto the do rsa l ve ssel is doubtful

.
.

The most probable vie w is th at the blood proceeds fro m the v en

'

tral ve ssel thro ugh ve ntro i nte sti nal vessel s to the c a pill a rie s o f
-

the i nte sti ne and the nce to the do rsa l ve ssel through the do rso
i nte stinal ve ssel s I t is po ssible ho wever that the r etu rn path
.
, ,

is through the dor so tegume nt ar y ve ssel s and th at the do rso-

i nte sti nal c arry blood f rom the dorsal ve ss el to the i nte sti ne .

I n th e fo reg o i nga cco unt o nl y th e mo re o bv i o us f ea t ures o f th e b l oo d

v essel s h a v e b een ment i o ned a nd ma ny i mpo r t a n t det a i l s h a v e bee n pas s ed

,

o v er. Th e circ ul a r v essel s o f th e s t o ma c h i nt es t i ne ca n be fo ll o wed f o r -

o nl y a s h o r t di s t anc o ut f ro m t h e do rsa l v ess el wh ere th ey s eem to b rea k

e ,

up i nt o a l a r g e numb er o f s mall pa r all el v ess el s l y i ngc l os e t og eth e r a nd

r unni nga round to th e l o wer s ide The efl ereut v ess el s do no t direc tl y j oi n .

'

t h e s ub i nt es t i na l b ut empt y i nt o a si n us o r v essel wh i c h r uns par a ll el to

-

t h e l a tt er c l osel y i mbedded in t h e wa ll o f t he s t o mac h i nt es t i ne

, The s ub -
.

i nt es t i na l v essel proper is q ui t e separa te f rom th e s t o ma ch i nte s t i ne a nd -

c o mmuni ca t es by s h o r t b r anc h es ( usuall y t wo in eac h so mi te ) wi th t h e

v essel l yi nga bo v e it Th is may be cl earl y seen in th e regi o n o f th e gizza rd

. .

O n th i s th ere is a v a riab l e number o f small l a t era l v essel s wh i c h b r ea k up ,

pa r tl y int o a b ra nch i ngnet work a nd are pa r tl y re so l v ed i nt o ex t remel y

,

fi ne parall el v essel s surroundi ngth e org an O n t h e c r p a re th ree o r f o ur . o

pairs o f l a t eral b ranch es f ro m t he do rs a l v ess el wh i c h b ranc h o ut i nt o a .

 B L OOD VE S S E L S -
. 57

fi ne net wo rk ,
i nt o para ll el v essel s as on th e gizzard
b ut do no t b reak up .

I n t he t wo so mi t es ( 1 3 th a nd 1 4 th ) in f ro nt o f t h e c ro p th ere a re us ually
t wo pa irs o f v essel s r unni nga ro und th e oeso ph a g us I n t h e 1 l t h and 1 2 th .

s o mi t es a s ma l l b r a nc h is g iven o ff to eac h ca l ci fero us gl a nd Th e most .

a nt er i o r pair o f c irc ul a r ves s el s a re in t h e 6 th s o mi t e a nd a re very s ma ll , .

I n f ro n t o f th i s th e do rs a l vess el b rea k s up i nt o th e ph a ryng ea l net wo r k .

I n f r o n t o f th e 1 1 th somi te th ere a re th ree s ub i nt es t i na l vessel s Th e two -

a ddi t i o na l vess els lie o ne n ei th e r s ide o f th e pri ma r y o ne a nd b r ea k up

,
o

i nt o b ranc h es at th e s ides o f th e ph ar ynx Th e aor t i c arch es empt y i nt o .

th e middl e v esse l a nd a t th e p i n t o f j unc t i o n th ere is a c o mmuni c a t i on

,
o

wi th t h e l a t era l vess l o f th e co rres po ndi ngsi de e .

B esi des th e do rsa l a nd s ub i n t es t i na l v essel s th ere a re th r ee ot h er mi no r -

long i t udinal trunks ( Fig Two o f th ese are very s ma ll and lie on .
,

FI G . 26 .
—Do rsa l vi ew r t f t h v n tr l r v rd h w i gt h rr n g m n t f
of pa o e e a ne e co -
, s o n e a a e e o

th v l f t h v tr l r g d d i p i m t ; i ub i t t i l v t r l
e es se s o e en a e i o n. s, s se en s . s -
n es na or en a

bl d v oo l h u ub ur l ; p n up r ur l Th ub i t t i l r iv
-
es s e s . .
, s -
ne a s . .
. s a ne-
a . e s -
n es na ec e es o n

e i th r i d th v tr l t r l ( M ) fr m t h p h ri d i f w h i h i t f rm th f
e s e e en o -
a e a s o e ne a, o c o s e c .

f t v
ere n l Th ub
es s e ur l i j d h id b y
e s t i u t i f th
-
ne a s o i ne o n ea c s e a co n n a on o e

d t gum nt ry
o rs o
'
-
e f ff r
e a t br h t th p h ri d i um ( f Fig
o , a e en a nc o e ne c . .

e i th er si de a b o v e t h e n er v e co rd ( p sendi ngfi ne b r a nc h es o ut f r o m
-
.

e ac h g a ng l i o n al o ngt h e l atera l ner v es Th ese a e th e s up ra n eura l t r un k s . r -

) Th th ir d l gi t di a l v es s e l ( s ub n ura l) l i es b el o w th e ner v e co r d
( a n e o n u n e
-
-
. .

( See Fig I t r ecei v es o n ea c h s i de th e t er mi na t i o n o f th e do rs o t eg

. u -

ment ar y v es s el Fi g 2 6) wh i c h in its c o ur e is co nnec t ed wi th t h e

. s

c a pi ll a ry net wo r k s o f t h e b ody wa ll a nd t h e d iss epi ment a nd g ives off a -

larg e b ra nc h to th e nep h ri di um ( c f Fi g . .
 THE B I OL OG Y OF A N A N I MA L
58
.

i
B es des th e
l a
t er a l v e s sel s f r om t h e s u b n e u r a l r r l a p a i r -
an d s up a neu a
-

m e fr o m
of
v en
t ro
l a
t e
r a
l (
-
v l Fi g s 26 a nd 2 7 ).
a re . g i
,
v e n o ff in e a
.
c h s o i t
i t t i l h ri d i m r b b l r ec ei v i ng f r o m it th e bl ood
th e s u b n e s n
-
a t o t h e n ep u p o a y ,

h i h ri i ll t r th r h b r h f h d rso teg ument ary

g y d u g a n c o t e o -

o a .

w c o n a e n e e

FI G . 27 .
—N ep hr i d i a h w i ngt h r gi of f t h t ub B t h v
L umbricus .
ulA s o e e o ns o e e. e as c ar

up p ly
s I I I I I I th th .
p ri ip l l p
. , . e ree nc a oo s .

A f f u n l ; u i th
. . n rr w t ube m t m i dd l t ub ; w t w i d ; t ub m p m u
e .
. e na o
“
.
, e e . e e : . . s

ul tub
c ar d v i l i di p m
e o r en t Th rr w t ub x t d fr m t a
-
es c e ( s. s se i en . e na o e e en s o a o

di i li t d b t w
an s c d h ta e d fr m
e I t T h m
ee n a a n i d d l ( i l i t d ) t ub
. a c, a n o ( o c . e e c a e e

t d f m gt h t h b i d t ub fr m h t k w h r i t p n i t th m u ul
e x en s ro o e r e e o o e e O e s n o e sc ar

x t l Op e i g
.

p rt ; a at , e ern a n n .

B L tt r
. b f r : dt d r
e e s as t gum t ry v l br i gi gbl d f m t h d r l
e o e . . o so -
e en a es s e . n n oo ro e o sa

v l r ivi g t b h fr m t h b d y w l l d i ng ff r t br n h t
es se . ec e n a s a
'
ra n c o e o -
a . se n an a e en a c o

th n ph d iu n e dfi
e ll y mi gt h u b ur l ( ) ; M v t r l t r l v l
ri r , an na ni n e s -
ne a a n . en o -
a e a e s se

r y i gth b l d f m th
ca r n p h r i d i um t
e th
oo ub i t
ro t i n l v nt r l v le ne o e s -
n es a or e a e s se

( t) ; m v t r l
a rv rd ( A ft r B nh m ; t h d i r t i n f th bl d u
c , en a ne e co-
. e e a e ec o o e oo -
c r

t rdi gt B ur )
ren s ac c o n o o ne .

E xcr et or y S y st em I t is the o fli c e of the e xc reto ry sy stem to

'

remove fro m the body pro per the was te mat te rs ulti matel y re
 OR G A N S OF E X R E TI ON C . N E PHR I DI A . 59

sultin
g fro m the b r e a kin g dow n of livi n g ti ssue This doe s not .

m e an t he p assi ng a way of the r efu se of dige stio n through the

an u s ( defaec atio n p fo r such m atters h ave n ever bee n
, .

abso r bed and the r efore h a ve n ever reall y bee n w ithi n the bod y

proper E x c retio n m e ans the re mov a l fro m the bod y of matter

.

w hich has re ally fo rm ed a p a rt of its sub s t anc e but h as bee n ,

use d up a nd is no lo nger a live I n higher an i mal s thi s functio n

.

is pe rfor m ed chiefly by the kid n ey s the l ungs and the ski n t he , , ,

w aste matter s p as si ng o ff in the u ri ne the bre ath and the s we at , , .

I n the eart h w or m it is p ri ncip ally perfor m ed by sma ll organs

ca lled nep hrid ia of which he re are tw o in e ach so m ite except
, ,

ingthe fi r s t th ree o r fo ur ( Fig .

E ach n eph ridiu m ( Fig 2 7 ) co n si st s of a lo ng co nvol uted

.

tube attached to t h e hi nder face of a

,

d issepi me nt and ly i ng in the c oelo ni at

,

the s ide of the ali me ntar y ca nal At .

o n e end the t ube p asse s th r o ugh the

(
bod y w all and O pe ns to the exte ri or by a
-

m in ute po re si tua ted betwee n the oute r

an d i nn e r r o w s of set se ( p The -
.

othe r end of the t ube passe s th rough the

dissepi men t ve ry ne a r to the poi nt
where th is is pe net rated by the n e r ve
cord ( p and ope ns b
.
y a b r o ad ,

funnel l ike exp ansion i nto the cavit y of Fm

-

p h r id i , f u l . ne a nne

the n ext so m ite in fro nt ( f Fig mu h l rg d h w i gth

, .
c en a e . s o n e

w t h b gi i g f t h
c
c a’ e e nn n o e
The ma rgi ns of the fu n nel and the i nner i l i t d l ( ) d th c a e c a na c , an e

s u rface of the upper p a rt of the t ube a re ut r h th o e s ea

de nsel y co v ered w ith po werf ul cilia ( Fig w ho s e actio n te n d s .

to prod uce a c urre nt setti ng fro m the c oelo m i nto the funnel and
thro ugh the neph ridiu m to the exterior .

Th e co i l s of th e nep h ri dium are di s po sed in th ree pri nci pal l oops ( I ,

II ,

III in Fig . Th e t ube i ts el f co mprises fi ve very di s t i nc t regio ns ,

as

fo ll o ws
1 . The f u nnel or n ep h ros to me fl a tt ened f rom abo v e do wnwa rds
much ,

wi th th e openi ngreduced to a h o riz o nt a l ch i nk I t is composed o f beau .

t i f ul ci l i at ed cell s set l ik e f a n ra ys aro und its edge I t l eads i nt o

-

2 Th e
. n a rrow tube ( n t ) a v ery del i ca te th i n wa ll ed co nt o rted t ube
,
-

ex t endi ngf rom the neph ros to me th ro ugh th e fi rs t l oo p and a pa rt o f the

s eco nd I n certai n part s o f its co urse ( a te 6 a t c and f rom d to c) th i s
.
, ,
60 THE B I O L OG Y OF A N ANI MA L .

b
t u e conta ns i c i l i a w h i c h a r e a rr a n g e d in t w o l o n g i t u d i n a l bands o n th e

i nner s ur face A t 9 it pass es i nto t h e

.

3
.
“
M iddl e t u b ( m t ) (g )
e
”
t o h e x t e n d i n,g s t r a i g h t th r o u gh th e sec o nd

l oop o f grea t er di amet er ci l i at ed th rough o ut and wi th pi gmented walls

,
, ,
.

A t I t it ol ens i nt o th e
4
. I l
/ id s t u b e
”
T h i s is o f s t i ll g r ea t e r c a l ib r e W i th g ra nu lar ,

l
ga ndu a l r w a ll s a n d w i th o u t c i l i a I t e x t.e n d s th r o u g h t h e s ec o nd l oo p

m h t i I I ) i n t n d th r o u g h t h e fi r s t f r o m i t o j a n d fi n a ll y i nto th e
( f ro o ,
o a ,

th ird openi ngat k i nt o th e

,

5 Mus cu la r p a rt o r d uct ( m p ) wh i c h fo r ms th e th ird l o op and opens to

.

h x t ri r t T h i t h w i d t a r t o f t h e e n t ir e n eph ridi um h as
t e e e o a e x s
. e e s , p ,

muscul a r wa ll s a nd fo r ms a k i nd o f sa c o r r es er v oir l i k e a b l adder in ,

wh ic h th e ex c ret ed ma tt er ma y accumul a t e a nd f ro m wh i c h it ma y be
ass ed o ut to th e ex t e ri o r
p .

The v arious par t s o f t h e neph ridi um a re h el d t og eth er b y co nnec t i v e

t i ue ( p a nd a re c o v ered wi th a ric h net wo r k f b l ood v ess e l s th e

o -

ss . ,

a rra ng ement o f wh i c h is s h o wn in Fig 2 7 B The s m ll er v ess el s us uall y

.
, . a

s h o w n umero us po uc hl i k e di l a t a t i o ns wh i c h mus t ser v e to ret a rd t h e fl o w

o f b l o od so me wh a t The v ess el s s uppl yi ngt h e ne ph ri di um a re co nnec t ed

.

( Fig27 B ) o n t he o ne h a nd wi th t he s ub i nt es t i na l v esse l th ro ug h t he
-

.
,

ventro la tera l trunks

-
o n th e th e r h a nd wi th th e s ub n u ra l ( a n ) a nd
o -
e

do rs a l v essel s th ro ug ,
h th e dors o tegmneuta ry -
Th e c o urse o f t h e
b l ood is so mewh a t do ubt f ul A cco rdi ngto th e v ie w h ere ad pt ed ( c f p 5 6 )
. o . .

t he b l ood proc eeds f ro m t h e do rs o t eg ument a ry t r unk to th e neph ri di a a nd

- -

th ence th ro ugh the v ent ro l at eral to the s ub i nt es t i na l a s s h o wn by t he -

arro ws in th e fi g ur e B enh a m ( f rom wh m th e fi g

. oures a re c o pi ed) ado pt s
th e re v erse v i ew The dev el o pme nt o f th e neph ri di um s h o ws t h a t its
.

ci l i at ed a nd g l andul ar po r t io ns a rise fro m a so l i d co rd o f dis k s h a ped cells -

wh i c h a f t erwa rds bec mes t ub ul a r by th e h o ll wi ngo ut o f its a x i a l po rt i o n

o o .

The t ube is th erefo re co mpa rab l e to a drai n pi pe in wh i c h eac h cy l i nder -

represents a cell I ts ca v i t y is no t i nt ercell ul a r ( bet ween t h e cell s l i k e t he

.
,

a l i me nt ar y ca v i t y) b ut i n tra cellu la , ( with in t h e c ell s l i k e a v ac uo l e)

r , .

The m ode of acti o n of the neph ridi a is as y et o nly pa rtiall y

under stoo d tho ugh there is no do ubt rega rdi ng their ge ner al c h a r
,

a c ter I t is c ert ai n th at their pri nc ip al offi c e is to re mo v e fro m

.

the body waste nitr oge no us matte rs re sulti ng fro m the dec o mpo
siti o u o f proteid s
; an d there is re aso n to beli e v e th at the se was te
matters are p assed o ut either as u rea o r as a n ea rl
y
rel ated substance together with a c e rt ai n q uantity of w ater and
,

i norganic salt s .

E x c ret i on in L u mb ic us appears h o wev er to i n v o l v e two q ui te dis t i nct

r
, ,

ac t i o ns o n th e par t o f th e ne h ri di a I n th e fi rs t pl a c e th e gl a ndul a r wall s

p .

o f th e t ube whi c h are ri c hl

, y s uppl i ed wi th b l oo d v essel s e l a bo ra t e certai n -

l iq i d was te s ubs t ances f rom th e bl ood a nd pass th em i nt o t he ca v i t y o f

u
 B B E A THI N G . 61

th e ut be . I n th e l ace th e ci l ia t ed f unnel s are bel i eved to t ak e up

sec o nd p
s o l i d was te pa r t i c l es fl o a t i ngin t h e c oel o mi c fl ui d a nd to pa ss th em on i n t o

t he tube wh enc e th ey a re ult i ma t el y v o i ded t o t h e ex t eri o r t og

,
eth er wi th

t h e li quid pr oduc t s desc ri bed a bo v e I t is nea rl y cer t a i n th a t th ese pa r t i

.

c l es are deri v ed f ro m th e b rea k ingup o f l ymph o id cell s so me of wh i ch

”
,

may h ave been ph ag oc y t es ( p fl o a t i ngin t he coel omi c fl ui d and th a t

.
,

m os t if no t all o f th ese cell s a ri se fr o m c hlo rag og ue cell s set f ree f r o m

t h e sur fac e o f th e b l oo d v essel s a nd o f th e i n t es t i ne

-

R espir ati on . o r b re athi ng is a twofold oper atio n

R e spiratio n , , ,

c o n s i sti ng of the t a ki ng in of f ree ox y ge n a n d the gi v i n g off of

c arbo n dioxide by ga seou s diffu sion th rough the surface of the

bod y S t rictly spe aki ng thi s free ox y ge n mu st be rega rded as
.
,

food while carbo n dioxide is to be rega rded as o ne of the ex ore

,

tio ns He nce re spi ration is t ri b uta ry both to ali me nt atio n and to

.

exc retio n ; but s i nce ma ny ani mal s po sse ss s peci al m echani sms to
ca rry o n re s pi ratio n it is co nve nie nt and cu stoma ry to t reat of
,

it as a di sti nct proce ss .

R e s pi ra tio n is e ss e nti all y an excha nge of ga se s bet wee n the

blood and the air c ar ried o n th rough a delic ate m e mbr ane l y i ng
,

bet wee n the m The e a rth w or m repre se nt s the si m ple st co ndi

.

tio ns po ss ible si nce the exch ange t ake s place all over the bod y
, ,

p reci sel y as in a pl ant I ts m oi st and delic ate w all s are every

.

w here tr a ve r sed by a fi ne n etwork of blood ve ssel s l y i ng j ust -

be neath the surface The oxyge n of the air either in the

.
,

a t m o sphere or di s solved in w ater re adil y diffuse s i nto the blo o d

,

a t all poi nt s an d carbo n dioxide m ake s its exit in the rever s e

,

dire c tio n Freed of ca rbo n dioxide and e n riched w ith oxy gen
.
,

the blood is the n car ri ed a way by the circul atio n to the i nner
p a rt s where it give s up its ox y ge n to the ti ss ue s and beco mes
,

o nce m ore l ade n w ith carbo n dioxide .

I n h igher anim al s it h as bee n proved th at the r ed colo ri ng

m atter (h ae moglobin) is the e s peci al vehicle for the a bso rptio n
a n d c a r ri age of the ox ge n of the blood e nteri ng i nto a loo s e
y ,

che m ic al unio n with it an d re adily setti ng it free agai n under the

a pprop riate co nditio ns This is doubtle ss true in the ea rth w o rm
.

al so .

I t is i nt eres t i ngto s t udy the v a ri o us dev ices b y wh i c h th i s f unc t io n is

p er fo r med in di ff er ent a ni ma l s I n t h e ea r th wo r m th e wh o l e o u t er s ur fa ce
.

is r es pira t o r y a nd no s peci a l r es pira t o ry o rg

,
a ns ex i s t I n o th er a ni mal s
.

s uc h o rg a ns a ri s e s i mpl y b y th e di ff er ent i a t i o n o f ce r ta i n r eg i ons o f th e

 THE B I OL O G Y OF A N ANI MA L .

62
i h th rr x h g f th w h ol e
eral sur f ace w h c e n ca y on t h e g a s e o u s e c a n e o r e
g e n ,
i l h r i b r fil m t r fl at
o rg ani sm I n m a n y a q u a t i c a n m a s s uc e g o n s e a a e n s o

k w h i h a re b a th ed
.

r w n as g i l ls o r b r a n c h i c e c
r f ea th er y p
n
p
l a t es o o ce s se s o ,

c h ani ma l s
w t r n t a i n i n g d i s so l v ed a i
r, th o u g h i n m a n y s u
by th e a e c o
e a s well I
r espira t io n t a
k e s p
l a c e t o s o m e e x t e n t o v e r t h e g e n e r a l s u r f a c . n

r f i fi d t n rr w t u b e s ( t r a c h e a ) w h i c h
i ns ect s th e esp r ir a t o r y s u a ce s c on n e o a o

ro w i nt o th e b ody f ro m th e s urf ace a n d b r h th r gh v r

a n c o u e e y p a rt
,
b ut
g r r i f l d d r t f t h o ut er s ur fa ce
g o e
v r th l b d ed a s a n n o e p a .

m us t ne e e e ss e e a

th r i b r th i n g v e r t e b r a t e s t h e r e s p ir a t o r y s u r f ac e is
I n man a d
n o e a r a-
e

ma y
i n
l c o n
fi n e d t o th e l u n g s w h,
i c h a re s i m p l y l oc a l iz e d i n f o l d i n g s o f th e

o u t e r s u r f a c e s p e c
i a ll y a d t
ap ed to e ff e c t a r a p i d e x c h a n g e o f g as es be t wee n

t h e b l oo d and th e a ir .

i ge
I t is e a s y to s ee w h y s p e ci a l r eg i o n s o f t h e o u t e r s u r f a ce h a v e i n h h r
i l b i d f r ir t i o n I t is e ss e n t i a l t o r a p i d d i ff u si on
an m n s e t a s e or es p a .

a s e e
r ir t r r f h u l d b c o v er e d w i t h a t h i n m o i s t m emb r a ne
th at th e esp a o y su ace s o e , ,

l t i l th t m n a n i m a l s s h o u l d b e pro v i ded wi th a
a nd it is no ess esse
n a a a y
t r v ri r t t i g i t m h i l i nj ur y o r des ic ca
fi rm o u e c o e ngas a p o e c o n a a n s e c a n c a

t i o n Hence th e o ut er sur face becomes mo re l ess di s ti nc tl y di ff erent ia t ed

.

i nt o two par t s v iz a pro t ect i ngpa rt t he genera l i nt egument ; a nd a

,
.
, ,

respirat ory par t wh ich is usua ll y preser v ed f ro m i nj ury by bei ngf ol ded
,

i nt o th e i nt erio r as in th e case o f l ungs or t rach eae o r b y bei ngco v ered ,

i th f l k i i ll fi h l b t r T h i co v eri ng
w o ds o f s n as in th e g s o f s e s ,
o s e ,s e t
. c s or

t ur ni ngin o f th e respira t o ry s ur faces b ri ngs wi th it the need o f mec h a ni cal

rr t i t r i t r ir t o ry c h a mber a nd
a ng emen s f o p m p g i w t h
r u n a r o r a e n o e e s p a
a
th us arise many compl ica t ed a ccesso ry res pira t o ry mech a nis ms .

B . OR G A N
S or R E L A TI ON .
( For A see p .

Mot or The move me nts of the b ody have a twofold

Sy st em .

purpo se I n the first pl ace they e nable the animal to alte r its
.

rel ati on to the enviro nme nt to move about (lo como tio n) to seiz e ,

m
,

an d swa llo w fo o d and to pe rfor va i

r ou s adap t i v e a c tio n s in
,

re spon se to ch ange s in the en viro nme nt I n the s eco nd plac e .

,

the m ovem ent s m ay alter the rel atio n of the v ario us pa rt s o f th e

body one to another ( v iscera l m ov emen ts and the like) s uch as ,

the move me nt s which propel the blood d ri ve the food alo ng the ,

ali me n t ar can al and roll it abo ut ( p those w hi c h expel

y .

waste matter s from the neph ridi a di s ch a rge the reprod uc t ive ,

prod uct s etc ,

.

Mo st of the se m ovem en ts are pe rfo r m ed by st ruct u re s kn own

as m uscles whi c h co nsi st of elo nga ted cell s ( fib res) e ndo wed in a
,

high degree with the po wer of con trac tility —i e of sho rte ning
'

, ,

“ ”
or drawi ng together (cf p O rdi na ry muscles a re in
. .
 M US OL E S . 63

th e fo rm of long bands or sheets of pa rallel fib r es such as tho se ,

that fo rm the bod y wall that m ove the set ae and dil ate the -

, ,

pha ry n x O ther mu sc ular st ructure s however do not form dis

.
, ,
”
t inct mu scle s but co nsi st of m u sc ular fibre s mo r e o r le ss
,

i rreg ula rl y ar ra nged and ofte n i nter m i ngled with other ki nd s of

ti ssue O f thi s cha racte r are the m usc ul a r wall s of the co ntrae
.

tile vessels and of the m u scul ar po rtio ns of the neph ridi a and
,

dissep ime nts I t is cle a r fro m the above th at the mu sc ul a r sy s

.

t em is n ot is olated bu t is i nti matel y i nvolved in m a ny o r gans

, .

Th e musc bo dy wall a re a rranged in two concent ri c l ayers

l es of t he -

bel o w th e s k i n I n t he o ut er l a yer t he musc l es run a ro und th e body a nd

.
,

a re th ere fo re ca ll ed c ircu la r mus c l es Th os e o f t h e i nner l a yer s h ave a

—
.

lo ng i tud i na l co urs e p a r a ll el wi th t h,e l o ng a x i s o f t h e b o dy a nd ,

are a rra ng ed in a numbe r o f di ff e re nt b a nds The mos t i mpo r t a nt o f th ese .

1 . Th e do rs a l ba nds ( Fig o ne o n ei th er s i de a bo v e in co nt ac t a t
.
,

th e medi an do rsa l l i ne and ex t endi ngdo wn o n ei th er s ide a s f a r as th e

,

oute r ro w o f set ae .

2 Th e ven tra l ba nds o n ei th er s i de th e middl e ven t ra l l i ne a nd oc cupy

.
,

ingth e space bet ween th e t wo i nner ( l o wer) ro ws o f setee .

3 Th e la ter a l ba n ds occ upyi ngt h e space o n ei th er s i de b et ween th e

.
,

t wo r o ws o f ac t ai .

A ll th es e va ry g rea tl y in diff erent regio ns o f th e b ody and in so me pa r t s ,

become mo re or l ess bro k en up i n t o s ubsi di a ry ba nds Th ere is a l so a .

na rr o w b a nd t r a v er s i ngt h e s pa ce be t ween t h e t wo s e t ae o f eac h g ro up .

Th e s eta; wh i c h may be r ec k o ned a s pa r t o f t h e mo t o r s ys t em a re pro

, ,

d uced b y g l a ndul a r cell s co v eri ngth eir i n ner ends a nd t h ey gro w co n ,

s ta n tly f ro m th i s po i nt so me wh a t a s h airs g ro w f ro m the ro o t A f t er

,
.

bei ngf ull y f or med and a f ter a cer t ai n a mo unt o f use t h e s et ae a re ca st

, ,

o ff a nd r epl a ced b y new o nes wh i c h h a v e mea nwh i l e b een fo r mi ng In .

ea c h g r oup we fi nd th erefo re setae o f di fferent s iz es A t th eir i nner ends

, , .

th ey a re co v ered by a co mmo n i n v es t ment o f gl andul a r c ell s wh i c h appea rs

as a s l ig h t ro unded pro mi nence wh en v ie wed f ro m wi th i n Th ese pro m .

inences a re c all ed t h e s etig ero us g la n d s Wh en a wo r m is l a id o pen f ro m .

a bo v e t h e g
,
l a nds a re seen in fo ur pa rall el ro ws t wo o f wh i c h lie on eith er ,

s i de o f t h e n er v e c o r d ( see Fig-
.

E ac h g roup o f setae is pro v i ded wi th speci a l retractor o r p rotra ctor

musc l es a nd a na rro w musc ul ar b a nd passes f ro m t he upper t o th e l ower
,

group o n eac h side i nt er na l to th e bo dy wall -

.

Cilia A s eco nd set o f mo t o r o rgans a re ci lia ( t h eir mode o f a ct i o n h as

.

be en re f e rr ed to o n p wh i c h a re o f th e ut mo s t i mpo r t a nc e in t h e
.

l i fe f th e ea rth worm Th ey c v er th e i nner sur face o f the s t o mac h i nt es

o . o -
»

t i ne ( wh ere th ey do ub tl ess as si s t in th e mo v ement s o f t h e f ood) pl ay t h e

i mp r t a nt par t in ex cret i on al ready descri bed co ll ec t a nd h el p to di sc h a rge
o ,
64 THE B I OL OG Y OF A N A N I MA L .

ments ( p i
a nd, as s s t i th f r t i l i t i o n o f th e eg g
t h e reproduct i v l
e e e .
n e e za

(p .
T h e ir ac t i o n l i
,
k e th a t o f t h e m u s c l e fi b-
res is do ub tl ess d ue to t h e
,

r r t f t t i l i t t h r t l a s m a lt e r n a t el y co nt rac t i ngo n oppos i t e

p o pe y o c o n ra c y ,
e p o o p
i
s des o f t h e c i l i u m a n d th u s c a u s i n g i ts w h i p l i k e a c ti o n .

m C l L m h ll P h ag oc y tes
Wh ite B l d p u s cl e s A oe bo id e l s y p ce s -

o o-
co r .
. . .

i m l ll d i l i t d l ls th e r e is a th ird v a ri e t y wh i c h dis pla y

B es des u sc -

e c e s a n c a e ce

t r t i l i t a n d m o v e m e n t T h e s e a re t h e coel o mi c c o rpuscl es ref erred to

c on ac y .

a bo v e ( p . U n t i l r e ce n tl y th e ir f u n c t i o n w as w h o ll y unk no wn b ut it ,

r ll b l i v th t th h a v e ng e rs o f th e body de v o ur
is no w g e n e a y e e e d a e y a re t e sc ,

ingt he dead t iss ues or fo reig n bodi es wh i c h i nv a de t h e o rg a ni s m Wh eth er .

h l tt k v r l i v i r i t h r g
t ey a s o a a c a n d d e o u n gp a as e s s u c as G a rin a a nd B a cteria
e

is not yet f ull y det ermi ned Th ey mo v e th eir p r t s much as A moebae do

.
a ,

e ng ul fi ngpar t i c l es a bo ut th em by a k i nd o f fl ux
. .

N er v ou s Sy st em . Org
ans of Coor dinati on .

The ge neral office of the ne rvous sy ste m of

'

I ntrod uctio n .

o rgans is to regulate an d c o ii rdinate the actio ns of all the othe r

p art s in such wi se th at the se actio ns shall fo r m an ha rmo nious

a nd orderl y whole Through nervo us organs the wo rm receives
.

fro m the e n v iro nment i m pre ss io ns which pass inward s th rough

the nerve s as sen sory or afferent i mp ul se s t o the nervo us ce nt res ; ,

a nd thr o ugh o ther ner v o us o rgans i m p ul se s ( e f eren t o r m o tor

j )
pass o ut ward s fro m the ce ntre s to the v arious pa rts so as to
a ro use m odif or su s pe nd their activitie s Thu s the ani mal is
, y , .

e nabled to c all f o rth m ove me n t s r e s ulti ng in the t wo ki nd s of

a dj ust me nt s referred to o n p 6 2 v iz ( a ) adjus t me nt s of the

.
,
.
,

b ody as a whole to cha nge s in the e n v iro nm e nt ( e g the wi th . .

,

drawal of the e a rth wor m i nto its b urro w at the appr o a c h of day ) ;
a nd ( l adj ust me nt s bet wee n the p a rt s of the b o d
) y it s e lf so th at ,

a ch ange in o ne p art ma c all fo rth n w e r i g ch ge in other

y a s n a n s

part s the i ncre ased suppl y of blood to the ali me nta ry ca nal
d uri ng dige stio n or vigoro us move m e nt s of the fo re end of the
,

b ody w he n the hi nd end is irritated ) .

The se functio ns a re al way s pe rfo rm ed b v o ne or m ore n erve

c ells whi c h give off lo ng sle nder b ra nc he s k n o wn as n erv e
,
fi bres -

usuall
y gathered together in bu ndle s the n er ves exte ndi ng i nto , ,

a ll p a rt s of the bod I n all higher a nima ls the m ai n b ulk of

y .

the ner v e cell s are aggregated in defi nite bodie s k nown as

-

ga ng lia o ut of whi ch i nto which or th ro ugh whi c h the nerve s

, , , ,

proceed ; and as a matter of co nve nie nc e it is cu sto m ary to desig

n ate the mo st i mpo rt ant of these ga ngli
a c o llec tiv elv as the cen
 N E R VE S A N D GA N G L I A .

tral nerv ous The r em ain i ng po rtio n which con sists

sy s tem .
,

mai nly of ne rve fi bres though it may al so con tai n m any ne rve
-

cells an d small spo radic ga nglia is k n o wn as the p erip heral ,

n er vo us sys tem .

Genera l A n a tomy of the N ervous S ys tem the e a rth . I n

wo r m the ce nt r al sy ste m co nsi sts o f a lo ng serie s of double ga ng li a ,

m eta mericall y r epeated and co nn ected by n e r ve cord s k now n as

,
-

com m is s ures The mo st a nte r ior pai r of ga nglia k n own as th e

.
,

s up ra ( es op hag
-
ea l or ce r eb r al ga ngli a , lie o n the do r sa l as pect of

the pha ryn x a sho r t di st ance beh ind the an terio r ext r e m ity
,

( Figs 2
. 4 F,
r o m each of the m a sle nde r co r d the si re um ,
-

(es op ha g ea l com mis s ure pass es do wn at the s ide of the p har n x

, y
to end in the su b ( es op ha g eal o r fir s t v en tra l
-

g g
a n lio n o n th e

lower side for mi ngwith its fellow a co mplete r i ng o r p ha ryn

,

g ea l co lla r a r ou n d the ali me nta r y c anal F r o m the sub oeso h a

p .
-

g eal ga n gli o n a lo ng double v en tra l nerv e cord p r oceed s bac k w a rd s -

in the m iddle ve ntr al lin e The ve ntral co r d co nsi sts of a series

.

of double ga ngli a one to each so mite co nn ected by co mm i ssures

, ,

and givi ng off l ate r al n erv es

*
.

I nter nall y the ce r ebral ga nglia and the ve nt r a l co r d ( c o m

miss ures as well as ga ngli a) co nsi st of both n e r ve cells and ne r ve -

fi bres as de scribed o n p 9 4 . .

P erip /zera l N ervous S ys tem To an d f r o m the ce ntr al sys .

tem ju st de sc r ibed run the n erves which co n stitute the periphe r al

s ste m The se a r e as follo w s
y
1 A pai r of n e r ve s r unni ng out o n eithe r s ide of each v en
.

tral ga nglio n and lo st to view a mo ng the mu scles of the body

w all.

2 A si ngle n er ve p r oceedi ng f r o m the ve n tr al co mm i ss u r es

.

o n each side i mm ediatel

y be h i n d the di ssepi m e n t to wh i ch it is
ma inly di stributed .

3 A pair of n erves fro m the sub (E sophageal ganglio n

.
-

4 A n erve fro m each half of the ph ary nge al coll ar j ust

.

bey ond its di ve rgence fr o m its fellow ( O r igi n i nc orrectl y .

sho wn ) .

5 .Two lar ge ce r eb r al ne r ves which run fo r war ds fr o m th e ,

So ly l th
c o se are e
‘

h l v es f th e v nt r l
tw o a o e a co rd uni t ed th at its do u ble
na t ure ca n s carc l y b e e made ut w i th o u t sec t i ons
o .
 THE B I OL O G Y OF A N A N I MA L .

66

FI G —A n t ri r p rt i f th rth w rm l id p f m b v w i th th lim n
. 29 . e o o on o e ea o a O en ro a o e, e a e

t ry d i ul t ry y t m d i t d w y
a an c rc i um m ph m l m
a o s s e s s s ec e a a . c. c .
, c rc -
so a oo

mi u ; ssg r b r l g gli a d i p i m t f t u
re c . .
, ce e l f p hr i d i um ; up
a an a , s, ss e en : , nn e o ne
nep h i d i um ; rv ry 0d vi du t ; p h p h y x p p r t m i m m m i l
o, o a , o c , ar n a, os o u ; 3 . se na

r p t l ; d p r m d u t f p rm tu u l ; L l t l min l v i l ;
ec e ac e 8 . .
, s e -
c ; s
. .
, s e -
n e a v. , a era se a es c e

t t ti ; c des s v tr l v
v . . d
, an v n c
. . .
, en a ne r e c o r -
.
 N E R VE JMP UL S E S 67
’
.

ce r eb r al ganglia b r eak up i nto m any b r anches

, ,
and ar e dis
tributed to the an terio r p ar t of the body .

i
B es des t h e ma i n gangl ia o f th e cent r al s ys tem th ere a re many s ma ll er ,

g g
a n l i a in v a ri o us pa r t s o f t h e bo dy O f th es e t h e mos t i mp o r t a n t a re t h e
—
.

p h a r y n gea l g a n glia 3 t o 5 in n um b e r — w h i c h lie o n t h e wa ll o f th e

p h a r y n x o n e a c h s i de j us t wi th i n th e p h a r y ng ea l co ll a r T h ey are co n .

nec ted wi t h th e l a tt er by fi ne b r a nc h es a nd send mi nut e ner v es o ut upo n ,

th e wall s o f th e ph ary nx Th i s series o f g

. a ng l i a is o f t en inappropria t el y
c a ll ed t h e s y mp a th etic s ys t em .

P hysiolog S ys tem i mp u lses

I

y f
o the N ervo vs erve . .

Wh at is the origi n and natu r e of a n e r ve i m pul se ? U nde r no r

-

mal condition s the i m pulse is set u p as the r e sult of so me dis

t urbanc e tech n icall y c alled a s ti mulus acti ng upo n the end of
, ,

the fib r e A tou ch o r pre ssu r e upo n the ski n fo r exa mple acts
.
, ,

as a sti mulu s to the ne r ve fi bres e ndi ng n ea r the po in t touched

-

th at is it cau se s nerve i mpulse s to travel i nwa r d s alo ng the fibres

,
-

to war ds the ce ntr al sy ste m The n erve s may be sti mulated by

.

a g r ea t variety of age nt s by m echa nical di stu rba n ce as in the

-

case ju st cited by heat electricit y che mic al action and in

, , , ,

speci al cases by wave s of light or of sou nd and u pon this p r op ,

e rty of the n erve s depe nd s the po w e r of the w or m to r eceive as

a ffere n t i m p u ls e s i m pre ss io ns f r o m the oute r w orld B ut be side s .

,

t hi s n erve fi bres m ay al s o be s ti mulated by ph y siological c hanges

,
-

t aking place withi n the ner ve cell s w hich may thus se n d out -

e ff e r ent impulse s to the various organs and so co nt r ol their ao

t io n .

R ega d ngth e p
r i reci se na t ure o f th e ner v e i mpul se we are igno rant but -

it is pro ba b l y a c h emica l o r mo l ec ul ar c h a nge in t h e pr o to pl as m t ra v ell i ng ,

r a th er rapi dl y al o ngth e fi bre l i k e a wa v e We k no w th a t t he na t ure

, .
*

o f th e i mpul s e is no t in a ny way depende nt upo n t h e c h a ra c t er o f th e s timu

lus Th e s t i mul us ca n o nl y th r o w th e ner v e i nt o a c t i on a nd th i s ac t io n

is a l wa ys th e sa me wh a t e v er be th e s t i mul us —as th e ac t i o n o f a c l oc k
.

remai ns th e sa me wh eth er it be dri v en b y a weig h t o r by a s pri ng .

Oo -
ordina tion
The activitie s of the variou s organs are c o
.

o r dinated by a chai n of eve nt s which in its si mple s t for m is k n ow n

as a refl ex a ction and w hich lie s at the botto m of m o st of

,

t he m o r e co m plica te d fo r ms of n e r vou s a ctio n I ts na ture is .

I n th f r gt h n r v u i m p ul
e o e tr v l t th r t f b ut 28 m tr p r
e o s s es a e a e a e o a o e es e

nd
s ec o in m n i t i n i d r b ly m r r p id
a s co s e a o e a .
68 THE B I OL OG Y OF A N ANI MA L .

ill ustr ated by the di agr am ( Fig o r di nation be . (J

o -

tween S and M (two organs) is no t effec t ed by a di r ect n e r vou s

conn ectio n but i ndir ectl y ,

through a nerve ce ntre 0 -

, ,

which is a n erve cell o r gr ou p -

of n erve cell s situat ed in o ne -

of the ganglia with which both ,

S and I ll are sep aratel y co n

nec ted by n erve fi bres If S -
.

be th r o wn i nto actio n an af er ,

en t i mpu ls e travels to 0 ex ,

cite s the ne r ve ce ntre and -

FI G Sid —
. Di g r m f i mp l r fl x t i n c ause s an eff eren t i m p ulse to
a a o s e e e ac o .

S k i t w h i h t im ulu i p p li d : f
o ut t O JI W h mh 1 8 the r e
o

th
,

ff r
e a
s n
e en
o

t rv fi b : C rv
c

ne
s

tr ;
e
t-
r avel re
s

,
s a

ne e c en
-
e

e
,

c f ff r
.
t n r v fi b ; M m u l in by thro w n i nto actio n also o r
e e en e e -
re . sc e
,
W h h th
'

10 ff r t fi br d e e e en e en s x

i s m od i fied m r e s pect to acti o ns

.

alre ady goi ng on Th us the actions of S a nd M ar e ( to ordi

.
-

na ted through the age nc y of C ; the whole chain of eve n ts

constit uti ng a r eflex action .

For example let S be the ski n and I l[ a ce r tai n gr ou p o f

,

muscle s I f the ski n be i r ritated aff ere nt i m p ul ses tr avel ih

.
,

w ard s to nerve centres in the ga ngli a ( C which thereupon se n d

-

forth e ff erent i m pul se s to the app r o p r iate mu sc le s Mu scula r .

contr acti o ns r e sult and the w or m draw s b ack f r om the un wel

,

co me irrit ation .

Thi s ch ain of event s i n volve s th ree di sti nct ac tio ns on the

art of the n erv o us s ste m w hich must be carefull di sti ngui sh ed
p y y ,

v iz
( ) the affere nt i m pul se ; (6) acti o n of the ce ntre ; ( 0)
a

the eff erent i mpul se I t must not be su pp o sed that the affere nt
.

impul se p asse s unch anged out of the ce ntre as the e ff ere nt i mp ulse ,

i e is si mpl y “ ”
. .
, r eflected like a b all th r own agai nst a w all as , ,
”
the word r eflex seems to i m pl y The aff e r e nt i m p ulse as su c h .

end s with the n erve cent r e whi c h it th r ows i nto acti vit y The
-

, .

eff erent i mpulse is a new action set up by the age ncy of th e

centre .

There is r eason to believe th at many if not all n e r ve cent r es -

are co nnected with a numbe r of differe n t aff e r e nt and eff ere n t

p ath s and al so with othe r c e nt r es as sho wn in the di agr a m

,
,

Fig 3 1 E ff erent i mp ulses may the r efo r e be se nt out f r o m

. .
 S E N SE S OF THE E A R TH WOR M . 69

th e centre in var ious di r ections and the precise path chose n ,

depen d s on som e u n kn ow n
actio n taki ng pl ac e in the "

ce ntre The action of the

.

ce n tre m oreove r may be

m odified by e ff ere nt i m pulses
a r rivi ng fro m othe r ce n t r e s ,

an d th us we can di ml y pe r

c eiv e h o w reflexe s may be

co nt r olled and guided and ,

how even the m ost co mpli

c ated for m s of n e r vou s ac
F 3l — D g r m r pr t gth
I G n rv e s en i n
t ity m ay be ( 30111 13 011 11 d
. . ia a e r ee e e

nt r d nn t A r w r pr
ce nt es a n co ec io n s . r o s e ese

out of ele m e nts si milar to th p ib l d ir t i n f r v i m p ul e o ss e ec o o ne e -

se s .

f n ff r n t p th ; f ff r
a t p th o e a e e a c o ne e e en a
reflex a ctio ns
.
, ,
.

There is r e aso n to believe th at in the earth w or m each v en

t r al g anglio n pre side s over the s o mite to w hich it belo ngs and ,

is probabl y in the mai n a collectio n of reflex ce nt r e s fr o m who se

actio n the ele m e n t o f co nscio usn e ss is ab s e n t B ut there is al so .

so m e r e ason to believe that the cereb ral ganglia occu py a highe r

po sit io n si nce they probabl y receive the n erve s of sight ta ste
, , ,

and sm ell be sides tho se of to uch w hile th e ve ntral gangli a re

, ,

ceiv e o nl y tho s e of to uch E xp eri m e nt h as sh o w n fu rther that

.

the cerebr al ga ngli a exerci se to a cert ai n li mited exte nt a c on

trol li ng actio n over tho se of the ve ntr al ch ai n by m e an s of im
pul se s se nt b ackward s thrbug h the co mmi ssu r e s tho ugh this ,

action is far le ss co nspicuou s he r e th an in highe r m et am eri c ani

mal s such as the i nsects *
.

Th e S ensitiv e Sy st em ( Or ga ns o f Sense ) The se n s itiv e

. .

sy ste m is di s ti ngui shed from the n ervou s system as a m atter of

co nve nie nce of de scription si n ce m o st of th e higher an i m al s
,

po sse ss defi nite s ense organs w hich r e c eive sti muli and thro w
-

i nto actio n the sensor y n erve s proceedi ng from the m A lthough .

“ ”
the e arth w or m po ss e sse s the s en se s of touch t aste sight , , ,

a nd smell it has no speci al organs fo r the se se nses ap art f r o m

,

the ge n er al i ntegu me n t coveri ng the su rface of the body and ,

Fo r a f ull r di s
e i
c uss o n th e t
s uden t is r f rr
e e ed to s p ec a i l wo rk s on Ph i
-

s i
l gy
o o .
70 THE B I OL OG Y OF A N ANI MA L .

hence can h ardly be said to possess any proper se nsor y sys tem .

“ ”
We do not k now m oreover w hether the so ca lled se nsatio ns
, ,
-

of the earth wor m are re all y st ate s of co nsciousne ss as in o u r selve s

for we do no t even k now w hether e ar thw o r ms po ss ess any for m
of c onsciousness W he n therefo re we speak of the e arth wor m
.
, ,

as po sse ssi ng the

“
se nse
”
of touc h or of sight we m ea n si m ply
th at so me of the nerve s ter mi nati ng in the sk in m ay be stimu
l ated by mech ani cal m e ans or by r ay s of light without necessa ,

r ily i mpl yi ng th at the wor m actu ally fe el s or see s as we feel and

86 8 .

I t h asrecentl y b een s h own th at th e sk i n co nta i ns many c ell s eac h o f

wh ic h g i ves o ff a si ngl e ner ve fi bre th a t may be t raced direc tl y i nt o th e
-

v entral ner v e cord Th ese s enso ry cell s may be rega rded as end
-
“
.
“

o rg a ns th ro ugh wh ic h th e s t i mul i a re co nv eyed to the fi b res I t h as al so

”
.

been s h o wn th a t th ese cell s a re aggrega t ed in mi nute gro ups th ic kl y sca t

t ereu o v er t he s ur face of th e bo dy E a c h o f th ese groups may be regarded
.

as a s i mpl e fo r m o f s ens e o rg
an -
.

The se nse of touch exte nds over the whole su r face of the
body Th at of tas te is probably located in the cavity of the
.

mo uth and pharnyx ; the l o catio n of the se n se of s mell is un

Da r wi n s eXperiments h a ve sho wn th at the earth
’
k no wn .

w or m s feeble se nse of s ig h t is co nfi ned to the a nte r io r end of

’

the b o dy I t is probable th at the ne r ve s of s ig

. ht taste and , ,

s mell e n ter the cerebr al g g

an lia alo ne while those of touch run ,

to other ga ngli a as w ell .

Sy st ems of ( Org ans of ) S uppor t Connecti on Pr ot ect ion etc , , , .

The struct ure and mode of life of many ani mals are such as to
require so m e solid support to the soft p art s of the body S uch .

supporti ng st r uct u re s a re for i nsta nce the bo ne s of ve r tebr ata

, , ,

the h ard outer shell of the lob ster o r beetle and the cor al ,

which for ms the skeleto n of a pol yp The ea rthwor m has .

,

however nothi ng of the sort and it is obviou s th at a hard sup

, ,

p o rtm
g org a n w
-
o uld be n o t o nl
y u s ele s s b ut eve n det r i m e nt al ,
.

The power of creepi ng and b urro w i ng th r o ugh the e ar th depe nd s

upon great flexib i lity and exte nsibility of the body ; and with
thi s the pre se nce of a skelet o n m ight be i nco mpatible .

The connectingsy ste m c o ns i st s s i mpl y of v ar io us ti ssues by

whi ch the di ff ere nt o rgans are b ound fi rmly tog
-

ethe r These -

can onl
y be see n u po n m i c ro scopica l ex am i nation The most .

impo r ta nt of the m is k no w n as con nec tiv e tiss ue .

 D E FE N CE S OF THE E A R TH WOR M
.

A s to p rotectiv e st r uctu re s the ea r th w or m is prob abl y one of

,

the m o s t defe ncele ss of ani mals N e v erthele ss there are ce rt ai n

.

s t r uctu re s w hi ch arecle a rl y for this p u rpo se The c uticle which

.

c over s the su rface is a thi n but tough m e m b r an e w hich protects

the delicate s ki n fr o m direct co ntact with h ard o bject s I t .

pas se s i nto the m outh and l ine s the alime nta ry ca nal as fa r do wn
a s the beginni ng of the sto mach i nte sti ne I n the gi z z ard
-

w he r e food is gro un d up the cuticle is p r odigiou sl y thick and

,

t o ugh and m u st for m a ver y e ff ective protectio n for the soft

,

ti ss ue s bene ath it The main defe nce of the ani mal lie s how
.
,

e ver n o t in an s pecial a r m o r but in those i nsti ncts which l ead

, y ,

it to lie hidde n in the earth du r i ng the day and to ve ntu r e fo r t h

o nly in the co mpa r ative safety of dar kn ess .
 C H A PT E R V .

THE B I OLO G Y O F A N A N I MA L ( Continued) .

Th e E a r t h w orm .

R E PR O D UCTI O N . E MB R Y O L OG Y .

R epr oducti on.

The l ife of ever y orga nic specie s ru ns in
r egul arly r e cur ri ng cy cles fo r every i ndi v idu al life has its l imit
,
.

I n yout h the co nstr uc tive proce sse s prepo nderate ove r the de
s tr uctive and the orga ni sm gro w s The nor mal adult attains a
.

s t ate of app a re nt ph y siological bal an ce in w hich the p r ocesse s of

w aste and repair are approxi matel y eq ual S oo ne r o r l ate r

.
,

how e v er thi s b al ance is di stu rbed E ve n though the o r gan is m

,
.

e sc ape s every i nju r y or spe c i al disease the c o ns tr uctive process

fall s behi n d the de stru ctive old age e nsue s and the individual
, ,

die s fro m sheer i nability to li v e W hy the vi t al machi ne sho uld

.

th us w e ar out is a mystery but th at it h as a defi nite ca use and

,

meani ng is i ndi c ated by t he famili a r fac t th at the sp a n of natu ral

life v aries w ith the specie s ; m an live s l o nge r th an the dog the ,

eleph ant lo nger th an m an .

I t is a w o nderful fact th at livi ng thi ngs h ave the powe r to

detach fro m the mselve s portio ns or f r ag me nt s of their own
bodie s e ndowed w ith fre sh p o wer s of gro wth and de v elop men t
an d c ap able o f r unn i ng through the sa m e c cle as the pare nt
y .

There is therefore an unbroke n materi al ( protopl as mic ) co nti nuity

fro m one ge ner atio n to another th at for ms the ph y sical b as i s of
,

i nheritance and upo n w hi c h the i ntegr ity of the s pecie s depe n ds

, .

A s f ar as k no wn li v ing thi ngs never ari se save th rough thi s

,

proce ss ; in other words every mass of exi sting protopl as m is

the las t link in an unbroke n ch ai n that exte nd s backwar d in the
past to the fir st origi n of life .

The detached portio ns of the pare nt that a r e to give r ise to

off sp ri ng are so meti me s masse s of cell s as in the sepa ratio n of
,

br anches or bud s amo ng plant s but mo r e co mmo nl y they ar e s i ngl e

,

72
 R E P R OD UOTI ON . 73

c ell s k nown as g
,
erm cells like the eggs of a nimal s and the
-

s pores of fe r ns an d m o ss e s O nly the g erm cells

( which may
.
-

co n v e nie ntl y be di sti ngu i shed fro m tho se for ming the re st of the
body o r the som a tic cells) esc ape death and that o nl y u nde r
, , ,

c e r t ai n co n ditio n s .

A ll fo r ms of r ep r oductio n fall u n der o n e o r the othe r of tw o

heads v iz A g ,
amog enes i s ( a s exu a l reproductio n) or Ga mog
.
,
enesi s

( sens u a l r eproductio n
) I n the for m er ca se the det ached portio n
.

( w hi c h m a
y be either a s i ngle ce ll or a gro up of cell s h as th e
)
po we r to develop i nto a new i ndi vidual without the i nfl ue nce of
o ther livi ng m atter I n the latter the detach ed po r tio n in thi s
.
, ,

c ase alw a s a si ngle cel l ( ovu m oo sphe r e is acted upo n

y , ,

by a seco n d po r tio n of livi ng m atte r l ikew ise a si ngle cell w hich , ,

i n m o st c as e s h as bee n det ac hed fro m the body of an other in

divid ual The ger m is c alled the f ema le g
. erm cell the cell act -

ingupo n it the m a le g erm cell and in the sexual proce ss the -

t w o fu se together (f ertiliz a tion i mp reg n a tion to for m a s i ngl e

) ,

n e w cell e ndo w ed w ith the power of develop ing i nto a n ew in

d ivid ual I n so m e orga ni sm s

. the y e as t pl an t an d bacteri a) -

o nl
y aga m oge ne si s has bee n ob ser v ed ; in other s ( e
g vertebr ate s
) . .
,

o nl
y ga m oge n es i s ; in other s s till both proce sse s take place as in

m any higher pl a nt s .

The ea rth w or m is not k now n to multipl y by any natura l

proce ss of agam oge ne si s I t po ssesse s in a high degr ee how ever .
, ,

the closel y r elated powe r of reg en era tio n f o r if a w or m be cut

t ransver se l y i nto tw o piece s the a nterio r piece w ill u sually make ,

good or reg enera te the m i ssi ng po r tio n w hile the po sterio r piece ,

m ay rege n er ate the an te rior regio n Thu s the w or m can to a .

c erta i n li m ited exte n t be artificiall y p r op ag ated like a pl ant by , ,

c utti ngs a proce ss clo sel rel a ted to t r u e a ga m oge n e sis I ts

*
, y .

u s ual an d n or mal m ode of r eprod uctio n is by ga mogen e si s that ,

is by the for matio n of m ale ger m cell s ( sp erma toz oa ) an d fe male

,
-

ger m cell s (ov a ) I n higher ani mal s the tw o ki n d s of germ

-
.

c ell s a r e produced by di ffere n t i ndivid ual s of oppo site sex The .

e a rth w or m o n the co ntr a r y is herm ap hrodite or bis exua l ever

; y

s" Man r
r l y r l t d t L umbri us t h gn u D r nd th r
y wo ms n ea e a e o c e e s e o, a o e

N ei d — p nt n u l y d ivi d th m l v
s s o a i nt t w p rt
eo s h f whi h b m e e se es o o a s ea c o c eco es

a p rf t n i m l
e ecT h i s proc
a i tru gm gn i th ugh bvi u l y c l l y
a . ess s e a a o e es s , o o o s o se

rel t d t r gnerat i n
a e o e e o .
74 THE B I OL OG Y OF A N A N I MA L .

i ndividual is both ma le a nd f ema k produci ng both eggs and ,

per m ato z o a The o v a ari s e in s peci a l o r g a n s the ov ar ies th e

s . , ,

sper mato z oa in sp erma ries or tes tes .

The ripe ovu m ( Fig 3 3 B ) is a r el atively l ar ge spherical .

,

cell agreei ng clo sely with the egg of the st ar fi sh ( Fig

,
but -

havi ng a thi nner and m ore delic ate m e mbrane I t is stil l cus .

to mary to appl y to ova the old te r m i n ology cal li ng the cell ,

sub stance v itella s the m e mb r ane v ite lline m e mbr ane the n ucleus
-

, ,

ermi na l v esicle an d the nucleol us g erm in al sp o t

g ,
.

The ripe spe r matoz oon ( Fig 3 3 0) is an ext r e mel y mi nu te .

,

elongated cell or fila ment thicke ning to wa r d s o ne end to fo rm

the head (n) which co ntai ns the nucle us of the cell e nveloped by a
,

thi n l ayer of p r otopl as m This is follo wed by a sho r t middl e .

”
piece (m) to which is attached a lo ng vib rato ry flagellu m o r tail
The tail s virtuall y a lo ng ciliu m ( p hi ch by vigo r ou s
()t . i w .

lashi ng drive s the whole cell alo ng head fo r e most ve ry much as -

a tadpole is drive n by its tail .

S i nce the ova r ies and s per ma r ie s give r ise to the ge r m cells -

they are called the essen tial org an s of

r eprod uctio n B esides these L um bricus .

, ,

like mo st a nimal s has a ccessory org a ns o f ,

r eproduction which act as r e se r voi r s or

carrier s of the ger ms assi st in secu ri ng ,

c r oss fertili z atio n and m i nister to the -

w ant s of the y ou ng wor ms .

E ssentia l R epr odu cti v e Or g a ns The .

ov a ries are two in num be r and lie o ne on

either side in the 1 3 th so mite attached to

the binder face of the anterior di ssepi me nt
( v Fig
o They are abo ut 2 ,
“m
in.
'

le ngth di sti nctl y pea r sh aped and at ,

-

tac h ed b v the broader end ( Fig Th e .

n a rro w extre m it co n t a i n s a s i ngl e ro w of

y
Fm . v ry much o v a and is called the e
o a ,
g g s tri n
g ( es
) I n -

en al rgd b th b l p t ; thi s the ova are ripe or ne arl y so ; behi nd

e .
, e a sa ar
a, b d y f th v r y
o o
thee o co n
sh ade off i n t o tho se m ore and m or e
a

t i i gi mm t ur
a n n
; a y e ov a 68 '
e gg t i g: n i p
-
s r n vu m i mmatu re till the se are lo st in a mass of
o , r e o
d y m f lmfi
,
rea a
n earl d iff ere nti ated cell s ( p r im itiv e
'

y u n
ov a
) co nsti tuti ng the gr ea t bulk of the ov a r
,
y E ach of these .
,
 R E P R OD UGTI VE OR G A N S .

however is su r rou nded w ith still smaller cell s co nstit ut ing its
,

nu trie nt e n velope or f ollicle A s the o v a m at u re the follicle s

.

still per s i st ,
an d the
y m a
y be detected eve n in the eg g g
strin .

Wh e n fully ripe the ovu m b ur st s the fol licle and is shed fro m
the end of the egg stri ng i nto the body cavity I t is ulti matel y
- -
.

take n i nto the oviduct a nd c a rried to the exterior .

The develop me nt of the ov a ry sho ws it to be mo r phologic ally

a thicke ni ng of the pe rito neal epitheli um The eggs therefor e .

are origin all epithelial cells

y .

The sp erma ries or tes tes ( t t Fig 2 9 ) are fou r in nu mber and, ,
.

in outwa rd a ppe a r an ce a re so me w hat s i mil ar to the ov a rie s .

They are small fl atte ned bo die s with so mewh at i r regular or lobed
border s l y i ng o ne o n either side the n erve cord in a po sitio n
,
-

corre spo ndi ng with that of the ov a rie s but in so m ite s 1 0a nd 1 1 , .

L ike the ov ary the te sti s is a s olid m ass of c ell s w hich a re shed ,

i n to the body cavity and are fi nall y c a rried to the exterior

-

The sper m cell s le ave the te sti s ho w ever at a ve ry e arl y period

-

, ,

and un dergo the l ater s t age s of m at ur atio n withi n the c a vitie s of

the se m i nal v e sicle s de scribed below .

A ccessor y R epr oductiv e Org a ns The m o st i m port ant of the .

ac ce ssor y orga ns are the ge ni tal d ucts by w hi c h the ger m cell s ,

-

are p assed o ut to the exterior B oth the fe male d uct s ( ov idu cts)
.

an d the m ale (sp erm duc ts) a re t ub ul a r org a n s ope ni ng a t o ne

-

en d to the o ut s ide through the body w all an d at the other end

,
-

i nto the coelo m by m e ans of a cili ated fu nnel so m e wh at si mil ar

to a n ephridi al f unnel but much l a rger B y m e ans of the se
,
.

cili ate d funnel s the ge r m cell s after their di scharge fro m the
-

ov a r y or te sti s are t ake n up and passed to the exterior .

The ov idu cts (0d Fig 2 9 Fig 2 3 ) are two short t r u mpet
,
.
,
.

sh ape d t ube s l y i ng i mm ediatel y po sterior to the ov a rie s an d p as s

ingth rough the di ssepi m e nt betwee n the 1 3 th and 1 4 th s o mite s .

The i nner end ope ns freel y i nto the cavity of the 1 3 th s o mite ,

by m e an s of a wide and m uch folded cili ated fu nnel fro m th e -

ce n t r e of which a sle nde r tube p ass es backward th r o ugh the

dissepim e nt t u rn s r ather sharply towar d s the o ute r side and
, ,

p ass i ng thro ugh the body w all ope ns to the outside on the 1 4 th
-

som ite (see p . I mm edi atel y behi n d the di ssepi me n t th e

ovid uct give s off at its dorsal and outer side a s m all po uch ,

rich l su lied with blood ves sel s I n thi s the rece ta c ulu m
y pp p
-

,
.
76 THE B I OL OG Y OF A N A N I MA L .

the ova t ke b the f el are te m po r a ril y sto r ed

o v orum n u u nn
,
a p y
before p assing o ut to the exterior .

prob ble th t the egg eve r flo t freel in the c oelom

I t is a a s n a y ,

b drop t of the ov a r at m a t u rit y d ir ectl y i nto the m outh of

u t o u y
the funnel They pass the nce i nto the recep tac ulum w he r e they
. ,

may re mai n for a co nsider able period .

The sp erm ducts ( vas a def erentia ) (sd Fig 2 9 ) a r e very

-

,
.

long sle nder tubes ope n like the oviducts at both e nds The,
.

outer openi ng is a co nspicuou s sli t su r ro unded by fles hy lip s

Fig on the ve ntr al s ide of the l oth so mi te Fro m this
( .
.

poi nt the duct r uns str aight for wa rd s to the 1 2 th so mite whe re ,

it br anche s like a Y the two b ra nches passi ng forwa r ds to te r,

mi nate one in the 1 1 th so mite the other in the l 0th

, ,
N ea r its .

end e ach bra nch is t wi s ted i nto a pe c uli a r k not and fi nall y te r
“
minates in an i mme nse cili ated funnel ( the so called ciliated -

the border s of whi c h are folded in so co mplicated a

manner that they fo r m a l aby r i nthi ne b o d y the t ru e natu r e of ,

w hich c an o nl y be m ade out in micro sc o pic s e c ti o ns .

The two pair s of sper m fun nel s ( Fig 2 9 ) lie in the 1 0th -
.

and 1 1 th so mite s i m medi atel y poste rio r to the re s pe c tive te s tes

, ,

i e they h ave e sse nti all y the sa me r el ati o n to the te s tes as that
. .
,
.

of the o v id uct fu nnel s to the ov arie s -

.

The t es t es and s per m f unnel s c a n be readi l y made o ut o nl y in young

-

s peci mens I n ma t ure wo r ms th ey a re co mpl et el y en v el o ped by t he se mi

.

nal v es ic l es desc ri bed bel o w .

S em ina l ves icles The se the mo st co ns picuou s p ar t of the

.
,

reprod uc tive appar atu s are v o l umi no u s po uche s in which the , ,

s perm cell s u n dergo their later de v el o p m e n t

-

afte r le avi ng the ,

t e st i s They are l arge whi t e bodie s l y i ng in s omites 9 to 1 2 and

.

usuall y o v erl appi ng t he ( e soph ag us in th a t regio n I n all case s .

there are three pairs of l ateral se mi nal ve sicle s v iz an a nte rior ,

.
,

p air in som ite 9 a middle pair in s omite 1 1 a nd a po ste ri o r p ai r

, ,

in so mite 1 2 I n immat u re S pe c i m e ns the s e six are e ntirel y

.

sep a rate and allo w the te ste s to be easil

, y s ee n i n m atu re .

w orms (as shown in Fig 2 9 ) the poste rior p air of l ate ra l

.

vesi cle s gro w together in the m iddle li ne thu s fo r mi ng a p os ,

terio r media n v es icle l i ng belo w the ali m e ntar ca nal in the

y y
1 1 th so mi te I n like man ner an a n ter io r media n v es icle is
.

fo rmed in the l 0th so mite by the u nio n of the two ante rio r pairs
 E G G L A YI N G
-
. 77

of l ateral vesicles The two m edian vesicle s thu s fo rm ed e nvelop

.

the tes te s and spe r m f unnels of the ir r e spective so m ite s and hide
-

t he m fro m view .

The s per m cell s l eave th e t es t i s at a very ear l y pe ri od and fl oa t f reel y

-

i n t h e ca v i ties o f t he semi nal v es i cl es wh ere ma ny s ta g es o f th eir dev el o

p ,

ment ma y eas i l y be o bs er v ed Th ey a re dev el oped in b a ll s k no wn as .

s p erma tosp heres eac h o f wh i c h co ns is t s o f a cent r a l so l i d mas s o f r t

, p o o

p l as m s urr o un de d b y a s i ng l e l ayer o f s per m cell s Wh en ma t ure the -

.

s pe r ma t o z o a s epa r a t e f ro m t h e ce nt r a l mas s a nd are dra wn i nt o t h e f un

n el s o f th e spe r m d uc ts The ma nner in wh i c h t h is ac t i on is co nt ro ll ed is

-
.

n o t unders t ood .

The semina l recep ta cles a re accessory o rgans of r ep r oductio n

in the sh ape of sm all ro unded sac s o r pouche s ope n to the o ut ,

s ide o nl at abo ut the level of the u pper ro w of set ae The

y, y .

lie betwee n the 9 th and 1 0th and l oth an d 1 1 th so mite s (s r , .

,

Figs 2 4 and
. where their ope n i ngs may be sought for ( Fig .

Thei r fu nctio n is explai ned u nde r the he ad of cop ul atio n .

A ccessory g la n ds B e side s all the s t ruct ure s so far de sc ribed

.

there are many gla nd s w hich pl ay a p a rt in the r eprod uctive

functio ns The set igerous glan ds fr o m about the 7 th to about
.

the 1 9 th somite ( so meti me s fewer so m eti me s n o n e at all) a re ,

often greatly e nlarged and fo r m the gl and ul a r pro mi ne n ce s men ,

ti oned at p 4 6 . They see m to be u sed as organs of adhe s i o n

.

du ring cop ul atio n The clitell um is filled w ith gl an d cell s which

.
-

prob ably serve in p art to secrete a no uri shin g fl uid for the y ou n g
worms and in pa rt to provi de a tough p r otecti ng m e m b rane to
,

cove r them .

Copul ati on Eg g y g
. la i n I nas m uch as
-
e ach i ndi vid ual earth
.

w o rm produce s both ova and sper m ato z o a it m ig ht be suppo sed ,

th at copul ation o r the sexu al u nio n of tw o di ffere nt i ndivid ual s

,
-

would n ot be n e ce ssary Thi s ho wever is not the c as e The .

, , .

o v a of o n e in dividual are i n variabl y fe rtili z ed by the s per ma to z oa

of an other i ndividual after a proce ss of cop ul atio n and exch ange

of sper mato z o a as follows : Duri ng the n ight ti me and u sually
,
-

in the sp ri ng the wor ms leave their bu rr o ws and p air pl aci n g

, ,

the m selve s so th at their he ad s poi n t in oppo site directio ns and

holdi ng fi rm ly together by the e nl a rged setigero us gl a nd s and the
thi c ke n ed lo w e r l ateral ma rgi ns of the clitell um Duri ng th is .

ac t the se m i n al r eceptacle s of e ach w or m are filled with spe rm a

t o z oa from the sper m duct s of the other a fter w hich the w or m s -

,
78 THE B I OL OG Y OF A N A N I MA L .

s eparate [The s per mato z oa thu s r eceived are si mply sto red up
. ,

a nd do not pe rf orm th eir f un ctio n u n til the ti m e of egg lay i ng]

'
-

When the wor m is ready to lay its eggs the gla nd s of th e

clitell um become ve ry active pouri ng o ut a thick gl airy fluid ,

which soon h arde ns i nto a tough m e mb ra ne and for ms a gir dle

a ro und the body B e s ide s th is a l arge qua ntity of a thick jell y
.

like nutrien t fl uid is poured out and ret ai ned in the space be
t ween the girdle and the body of the wor m The gir dle is .

thereupo n grad uall y worked forward t o wa rd the head of the

w or m by co ntr actio ns of the body A s it p ass e s the 1 4 th so m ite .

a n u mber of ov a are received from the ovid ucts and bet ween ,

the 9 th and 1 1 th so m ite s a qu antity of sper mato z o a are added

from the se mi nal receptacle s whe re they h ave bee n sto red si nc e
the ti me of copul atio n w he n they were obt ai ned fro m anothe r ,

wo rm The girdle is next st ripped fo r wa rd s ove r the ante rio r

.

e nd a nd is fi nall thro w n
y
co m pletely off A s it .

pas s es off its ope n e nds

i mmediatel y c o n t r a c t
tightl y together and the ,

gi r dle bec o mes a cl osed

g capsule ( Fig 3 3 ) co ntai n .

ingboth o v a a n d s pe rma
B o
— to z oa fi o a tingin a n ut ri
F I G33 A.
gg p u l . l rgd 5 d i m t r
, e -
ca s e en a e a e e s
( f w g
a g n e l r gd t t h m
e s, o l tl v e fi lll d O I l nl lk
, en a e The o e sa e sc a e a re
’
.

mu h l rg d ; C p m t
c en a rm u l y
e
m e m br a ne 8 0011 a ss u me s a
. a e e a o z o o n, e n o o s
m gnifi d ; n h d ; m m i d d l p i
a e
; t t il , light y ellO W lS ll O l LI O WI )
ea , e ec e , a .
’ '

color bec o me s hard a nd to ugh and serve s to pr o tect the de

, ,
'

p ge mbry o s
v elo in
The c ap sule s may be fo und in May or Ju ne .

in e arth u nder logs or sto ne s or e s pe c i all in he ap s o f manure “ '

y
'

, .

Withi n the capsule s the fertili zatio n and develop me nt of the o v a

t ake pl ace .

Ferti li zati on and E mb r ol o i cal Dev el o ment

y g p The s pe r ma .

t o zoa sw nn activel ab o ut in the nutrie nt fl uid of the cap ule

y s
,
a ppro ach an ovu m a nd attach the mselve
s to its su r face b the ir
y
,

he d s S everal of the spe rmato z oa the n e nte r the v itellus (c f

a .

p .
but it h as bee n proved th at o nly o ne of the se is c o n
cer ned in fertili zatio n the othe rs dy i ng an d beco mi ng absorbed ,

by the ovu m .
 FE R TI L I ZA TI ON OF THE E GG . 79

p r ob able th at the t ail pl ays n o p a rt in the ac tual fe rtili

I t is
z ation but is m erel y a loco m o tor app a r at us for the head (nucle us
, )
and m iddle piece -
.

Withi n the ovu m the he ad of the spermato z o o n per si st s as

the sp erm nu cleus ( or m a le p ro n u cleus) while the protopl asm in
- -

its n eighborhood as su m e s a peculi a r a nd ch a r acte ri stic r adi at e

a rr ange me n t l ike a star probabl y through the i nflue nce of the ,

m iddle piece -
.

A fte r the e nt r a nce of the sper m ato z oo n the egg segme nts off

D
‘

9
FI G . 34 .
—Fer t i l i z a t i o n o f r m t z n ( i th th e o v um . A , en tr a nc e of th e s pe a o oo n e s ea

ur h i c f t r F l)
n, a B th ur h i
e gg f t r tr o f th p rm t z o ;
.
, e s ea -
c n e a e en a nc e o e s e a o o n

w i th i d t t h l ft i th
n an gg u l u ; b v i t h p rm u l u w i th
o e e s e e -
n c e s a o e s e s e -
n c e s, a c en

t mro so r i t ( m d i fi d f r m H r t w i g) C d i gr m f t h v um f t r t ru i n
e n ea o e o e .
, a a o e o a e ex s o

o f th p l r e ll o a d u i f th tw
ce p u l i ts f rm t h gm t an n on o e o ro n -
c e o o e se en a

ti u l u
on n - Th m ll r
c e d d rk r p rt i
s . f t h l tt r i d r iv d fr m t h
e s a e an a e o on o e a e s e e o e

s p rm e u l u Tw n t c e r h p l m ph r s h w r th u l u
o a s ers o r a c o as s e es a re s o n n ea e n c e s
- -
. .

Th ri b y th d ivi i f i gl t r d r iv d f m t h m i dd l p i
es e a se e f th
s on o a s n e as e e e ro e e -
ec e o e

s p rm t z o
e D tw
a o ll d t g f th
o n rth w rm f t r t h fir t fi i f
. , o ce-
e s a e o e ea o , a e e s s s on o

th v um ( A f t r V j d k y )
e o . e e ov s .

at o neide two small cell s o ne after the other k n ow n as th e

s , ,

ola r cells or p ola r bodies The s e take n o p a rt in the for m atio n

p .

of the embr y o and their for m ation probably se r ve s in so m e w ay , ,

n o t y et wholl y cle a r to prep are the egg fo r the l ast act o f

‘

fertili zatio n A fter the for matio n of the pol a r cell s the egg
.

n ucl e us ( no w ofte n c alled the f ema le p ro n u cleu s) and the s per m -

n ucle us appro ach o ne a n othe r and fi nall y beco m e i n ti matel y

80 THE B I OL OG Y OF A N ANI MA L .

a vag
a ss ociated to for m the s e
g m e n ta t io n o r c le e n u cleus b -

y th i s
a c t fertili z atio n is co m pleted .

Th e proc ess of fer t i l iz a t i o n a p pea rs to be essent i a ll y th e sa me among

a ll h i g h er a ni m a ls
,
a n d in a b r o a de r se ns e t o b e i de n t i c a l wi th t h e se xua l

r o
c
e s s a mo nga ll h ig h er a nd ma ny l o wer pl a n t s ( co mpa re t h e f er n p
p , .

b ut its precise na t ure is s t i ll i n dis put e I t is cer t a i n th a t o ne esse nt i a l

.

a r t o f it is th e un io n o f t wo nuc l ei deri v ed f ro m t h e t wo r es pec t i v e pa ren ts

p
.

T h is h as led to t he view no w h el d by ma ny i nv es t ig
,
a t o rs th a t i nh eri t a nce ,

h as its sea t in th e nuc l eus a nd th a t c h ro ma t i n ( p

,
is its ph ys i ca l
.

bas i s L a t er researc h es h a v e s h o wn th a t a noth er el ement k no wn as the

.

a rc h o pl a s m or a tt ra c t i n sp h er e is c o ncer ned in fer t i l iza t i o n a nd th is is

o -

a ppa rentl y a l wa ys de ri v ed f ro m t he mi ddl e pi ec e I t is no t yet c e rt a i n

-
.

wh eth er t he arc h o pl as m is to be reg a rded as a nu c l ea r o r a c yto pl as mi c

s t r uc t ur e a nd it is eq ua ll y do ub t f ul wh eth e r it pl a ys a n essent ia l o r merel y

,

a s ubs idiary m i l e in ferti l i za t i o n a nd i nh eri t a nce ( c f p . .

Cleava g e of th e Ferti li zed Ov um . S oo n afte r fertili z atio n the

o v u m begi ns the re ma rk able proce ss of segme ntatio n which
h as alre ady bee n briefly sketched o n p 2 5 . The seg men.

tation nucle us divide s i nto t w o p a rt s and thi s is followed by

-

a divi s io n o f the vi tellu s each h alf of the o r igi nal nucleus becom
,

ingthe nucle us of o ne of the h alve s of the vitellu s ; th a t is the ,

o rigi nal cell divides i nto t wo s maller but si milar cell s ( see Fig .

These di v ide in t urn i nto fou r and the se i nto eight and , ,

so o n but et re mai n clo s el co n n ected i n n m I n the

, y y o e ass .

c a se of the e a rth w o r m the cell s do no t m ultiply in regul a r

,

geometri cal progre ss io n but sho w ma ny i rr egul a ritie s ; and m o r e

,

o ver the be c o m e un eq ual in s i z e a t a n ea rl pe riod

y y .

The bl ast ul a ( pp 2 5 .
,
sho w s sca rcel
y any differe ntiatio n

o f p art s tho ugh the cell s of o ne he mi sphere are s o m e w h a t sma lle r

,

th an the oth e r s . From thi s ti me for wa rd s t he w hole co urse o f

de v el o p me nt is a proce ss of differe nti atio n both of the c ell s and of ,

t h e o rgans i nto which the so o n a r ran ge the mselve s O n of

y e .

the fi r st step s in thi s proce ss is a fl atte ni ng of the e mbryo at the

lower p o le the h alf co nsi sti ng of l a rge r cell s ( Fig 3 5 D ) .
,
.

The l arge cell s are the n folded i nto the segme ntatio n cavity so -

a s to for m a po uch O pe ni ng to the exterior at the sa m e ti m e

;
the e mbr yo beco mes so me w h at el o ngated ( Fig 3 5 E F ) .
, ,
.

Thi s pr o ce ss is k now n as g as tru la tio n a nd at its co m pletio n

,

t h e e mbr o is c alled the

y g a s tr ula . The i nfolded po uch ( ca ll ed
the a rchen teron ) is the futu re ali me nta r y canal ; i ts ope ni ng ( now
know n as the bla s top ore) w ill beco me the mou t h ; and the l ay e r
 THE GE R M L A Y E R S -

. 81

of small cells ove r the out side wi ll fo r m the ski n o r outer lay e r
of the body w all -
.

The e mb ry o ve ry soo n begins to swallow through the blasto ,

po re the m ilklike fluid in which it floats and to digest it with

, ,

in the cavity o f t h e a rche n tero n .

I t is obv io us that the e mbr y o already shows a disti nct diffe r

FI G . 35 .
—Di a gra m s o f th e ea rl y s t a g s
t i th rth w r m A u e of de vl e op m en n e ea o .
, acc

r t dr wi g f t h bl t u l ur u d d b y th vi t ll i m m br n ( ft r V j
a e a n o e as a, s ro n e e e ne e a e a e e

d k y ) ; B bl t ul i
ov s p ti l t i h w i gt h l rg gm t t i n vi t y
, as a n o ca s ec on s o n e a e se en a o -
ca

d th p r t ll f t h m bl t
an e a C l t r b l t u l h w i gf rm
en -
ce o e eso as , a e as a, s o n o a

t i n f m bl t ll : D fl tt i g f t h bl t u l pr p r t ry t i v gi t i n ;
o o e so as -
ce s , a en n o e as a e a a o o n a na o

E th , g trul i ei d i w ; t h i f ld i g t k p l t h t w m bl t
as a n s e V e as e n o n a es ac e e o e so as

b d an l ft t th
s a re i d f t h b d y i th p i t i h w b y th d tt d l i ;
e a e s es o e o , n e os on s o n e o e n es

F t i n f E l gth l i n
. sec o o h w i gth m bl t b d d p l ll
a on e e 8 8, s
-
o n e e so as -
an s an o e ce -
s .

entiatio n of p a rt s which perfor m u nlike fu nctio ns I n fa c t w e .

m ay r egard the gast r ul a as co m posed of two tissue s still nearl y

s i m ilar in s tr ucture though u nlike in f un cti o n O ne of these /
.

consi st s of the lay e r of cell s w h ich fo r ms the o uter co v eri ng ;

thi s ti ssue is k nown as the ecto blas t ( cc Fig The secon d ,
.

ti ssue is the lay e r of c ell s form i ng the w all of the arche nteron ;
it is called the en to bla st (en ) The ectoblast and e ntoblast to .

gether are k nown as the p r ima ry g erm la ers

y
-
.

M eanwhile change s are taki ng place which r esult in the fo r

m atio n of a third ger m l ay er l ying in th e segm e ntation c avity - -

betwee n the ectobl as t and e ntobl ast and therefore called th e

m es obla s t (m Figs 3 5 I n so m e ani mal s the m esobl as t
, ,
.
,

does not ari se u ntil afte r the co mpletio n of ga st r ulatio n I n .

 THE B I OL O G Y OF A N A N I MA L
82
.

howeve r it goe s on d r u in g g a t r ul atio n an d beg i n s s

L umbmc
’

u s , ,

gast r ul tio E ve in the bl as tula s t a ge two l ar ge

evenbefore a n n .

s gi ve r is e to the
.

cell s may be disti ngui shed which afterward .

m w w bl s tw c lls
a e c lled the an e
s
mesobl t as a n d r h en c e a p r i a r y o .

They soon bud fo r th s m a ller cell s i to the

n s
e g m e n ta h o cav it
y
-

l fl tte n s they
the m s elve s s i n k below the sur
a nd as the bl tu
as a a

the m e oblast fo r m s tw o ba n ds
face A t thi s pe r iod the r efore
.
s
, ,

each ter m i ti g behi n d in the la rge

o f cell s ( m es obl a s t ba n d s
) n a n -

other cell or l ell

Througho u t the late r s tage s the pole
m p o e c
-
-
.

c
ell s
con
ti nu e to b u d fo r th s m alle r cell s which are added to the

b inder end s of the me obla t s s ba n d s ( Figs 3 5 -

.
,

FI G . 36 .
—D ia gra m s o f l a t er embry n i c s ta ges
l t t g in l n gi t ud i n l t i n o . A , a e s a e o a sec o ,

s h wi ngth p p r
o f th vi t i f th m i t ; B th m in
e a ea a n c e o e ca es o e so es , e sa e c ro ss sec -

t i n ; E d i gr m f y u gw rm in l ngi t ud i n l t i n f t r t h f m t l f
o , a a o a o n o o a sec o a e e or a on o

th t m d um p t dmum d u : C th m i r
e s o o ae , roc ti h w in g
o , an an s , e sa e n c o ss se c -
o n, s o

th b g i n i g f t h n rv u y t m ; D r
e e n n o t i n f l t r t g w i th th
e e o s s s e , c o s s sec-
o o a e s a e e

r v u y t m mp l t l y t b l i h d l l i m t y
ne o s s s e co l: h nt r
e e es a s e . a , a en a r c a na a r, ar c e e on

a n u ; w
, l m;an t bl t
s c n t bl
, t ; m p ri m ry m bl t i ll
c oe o cc, ec o as : en , e o as ’
, a eso as c ce s

m m bl t ; mh m uth ;
'
, eso rv u y t m ;
as vi ty f m i t ; m m t i
, o 71 , n e o s s s e 8, c a o so e a , so a c

l y r f th m
a ebl t w h i h w i th th t b l t f r m th m t p l ur ; p un
o e eso as , c e ec o as o s e so a o e e s .

pl n hn i l y r f t h
s a m c bl t wh i h w i th t h t bl t f m t h p l n h
c a e o e eso as , c e en o as or s e s a c

nop leure .

A fte r
each divi sio n the pole cells i nc r ease in s i z e so th at up -

to a late stage in developme nt they may be d isti nguished fro m

 CE L L -
DI VI S I ON . K A R Y OK I N E S I S . 83

I lh e t w o m as se s of me sobl astic
r
the c ells to which they give r ise .

c ells gradu ally i ncre as e in siz e andxfi nally fill the segmentatio n
s
i nter nal ph enomena of cell di v i sio n are of grea t compl ex i ty and
Th e -

c a n h er e b e g i v en o nl y in o utl ine Th e o rdi na ry t ype o f cell divis io n a s .

,

s h o wn in th e s eg men t a t i o n o f th e ov um a nd in th e mult i pl i ca t ion o f most

ti ss ue cell s i n v o l v es a co mpl i ca t ed seri es o f c h a ng
,
es i n th e nuc l eus known

as km yok ineszs o r mi tos is Th es e c h ang es wh i c h a ppea r to be o f ess en

.
,

t ially t h e s a me c h a r ac t er in nea r l y a ll k i nds o f cell s a nd bo th in pl a nts and ,

-
i n ani mals a re i ll us t ra ted by th e f o ll o wi ngdi ag
,
rams

C D
‘

FI G . 37 .
—Di a gra m s o f i n d i rec t c ell d iv isio n o k a ry o k i n es i s
~ r .

A . C ll j u t p i r t
e sivi i r o h w i gnu l u ( n) w i th it h m t i t i u l um nd
o d s on, s o n c e s s c ro a c re c a

th tt r t i p h r d t r m ( )
e a ac on s -
e e an c en o so e c .

B Fi t p h
. rs ; th tt r t i n p h r h d ivi d d i n t t w w h i h h v m v d
a se e a ac o -
s e e as e o o, c a e o e
1 80 p r t ; th r t i ul um h b r lv d in t fi h r m m ( b l k ) h
°
a a e e c as een e so e o ve c o o so es ac , eac
f wh i h h
o p lit l
c gt h w i
as s en se .

0 S. d ph
ec o n ; f ull y d v l p d k r y k i
ase t i fi gur ( mp hia t ) w i th p i n d l
e e o e a o ne c e a s er , s e
an d t r ; th h m m h l v
as e s e c m vi g p r t
ro o so e -
a e s a re o n a a .

D Fi l ph na : th ll b d y i d i i d i n g th
a se e ce p i dl d i
-
o pp ri ng th d ught r
s v e s n e sa ea e a e
f rmd
.
, ,

nu l i b u t t b
c e a o o e o e .

I n its r es t i ngs t a t e th e nucl eus conta i ns a netwo r k or reticulum of

c h roma ti n ( Fig 3 7 A ) A s t h e cell pr epar es f o r di vi si on a s ma ll b ody ( 0)
.
, .
84 THE B I OL OG Y OF A N ANI MA L .

mak es its appearance near t he l k nown as th e a ttraction sp hwe o r

nuc eus , -

a rch op la s m mas s a nd in its i nt eri o r th ere is o f t en a s ma lle r bo dy

-

,
th e ,

centroso me Th e fi rs t s tep in c ell di v is io n is t h e fi ss io n o f th e a rc h o plas m

.
-

mas s i nt o t wo eac h cont a i ni nga c ent roso me ( derived by fi ss i on o f t h e

,

o rig inal cent rosome) ; a f t er th is the t wo mas ses mo v e apar t to o ppo s i t e

o l es o f t h e nucl eus ( Fig 3 7 B ) The ret ic ul um no w bec o mes in mos t
p ,
.
,
.

cases resol v ed i nt o a th rea d co i l ed i nt o a s k ei n ( no t s h o wn in t h e fig

,
ure) ,

wh ic h fi nall y b reak s up i nto a number o f bodies k no wn as ch ro m so mes o .

Th eir f or m (g ranul a r rodl ik e l oop s h aped) and number ( t wo ei gh t t wel v e

, ,
-

, , ,

s i x t een etc ,
or o f t en muc h h ig
.
,
h er numbers ) a ppear to be c ns t a nt f o r o

eac h s pec i es o f pl a nt a nd a ni ma l Th e seco nd p ri nc i pa l s t ep is t h e l o ng

. i
tudinal spl i tt i ng o f eac h c h ro moso me i nt o h a l v es ( Fig 3 7 B ) a nd t h e .
,

disappea ra nce o f t he nuc l ea r memb ra ne .

I n th e th ird pl a ce s t a rl i k e ra ys ( a s ter) appear in th e pro t o pl as m a ro und

th e arc h opl as m masses a s pi ndl e s h a ped st ruc t ure a ppea rs bet ween th em
-

,
-

( Fig 3 7 C) and th e doub l e c h ro mo so mes a rra ng

.
, ,
e th e ms el v es a r o un d th e

eq ua t o r o f th e spi ndl e The s t ruc t ure th us fo r med is k no wn as th e a mp h i

.

as ter or ka rg e k i net ic fi g u e r .

Fourthl y t h e t wo h al v es o f ea c h c h ro moso me mo v e a par t to wa rds th e

,

respect i ve pol es o f th e spi ndl e a nd th e ent ire cell body t h en di v ides in a -

pl ane passi ngth ro ug h th e eq ua t o r o f the spi ndl e E a c h gro up of da ugh ter .

c h r o moso mes no w g i ves ri se to a ret icul um wh i c h beco mes s urr o unded wi th ,

a memb ra ne a nd f o r ms t h e nuc l e us o f t h e da ug ht er c ell The s pi ndl e dis -

.

appea rs a nd in so me cases t h e a rc h o pl as m mas s wi t h it s s t a r r a v s ( as t er)

,
-

,
-

seems to dis a ppea r a l so I n o th er cas es h o we v er t h e a rc h o pl as m mas s a nd

.
, ,
-

c ent ros o me pers i s t a nd may be fo und in th e res t i ngcell in l e uc oc y t es

a nd co nnec t i v e t i ss ue cell s ) l yi ngnea r th e n uc l us in t h e c yt o pl a s m
-

, e .

I t a ppea rs f ro m th e fo reg o i ng desc ri pt i o n th a t eac h da ug ht er ce ll re -

cei es ex ac tl y h a l f th e s ub s t a nce o f t h e mo th er nuc l eus ( c h r o ma t i n) mo th er

v -

a rch o pl as m a nd mo t h er cen t r oso me

, I n ma ny cases th e c y t o pl as m a l so
-

di v i des eq uall y in o th er cas es uneq ua ll y

, .

I t h as b een pro v ed in a co nsi dera b l e number o f c ases t h a t in th e f er

t iliza t ion o f th e o v um eac h g e r m cell co nt ri b ut es t h e sa me numbe r o f ch ro
-

moso mes and th e wo nder f ul f ac t h as bee n es ta b l is h ed wi th h i g

, h pro ba b i l i t y
th a t the pat er nal and mat er nal c h ro mat ic s ubs t a nces are eq uall y dis t ri but ed
t o th e two cell s fo und at th e fi rs t se ment a t i o n o f th e o v um I t is f urth er
g .

pro b ab l e th a t th i s eq ua l dis t ri b ut ion co n t i nues in a ll the l a t er divis io ns ;

a nd if th i s is t r ue ever cell i n the who le a du lt bod
y , y co nta ins ma te ria l
directly deri ed f ro m both pa r ents a nd hence ma inh eri t ro m both
v
y f , .

Ga strulati on . Germ la y er s -
. Diff er entiati on . Origin of th e
B ody . A l mo s t
fr o m the fi rst the cell s a rrange the mse lves so as
to surrou nd a ce ntral cavity k no wn as the seg men ta tion W -
.

Thi s cavity i nc reases in si z e in late r s tage s so that the e mb ry o ,

fi nall y appea r s as a hollow sphe r e su rr ou nded by a w all co nsist

 D E VELOP M ENT OF THE OR G A N S . 85

ingof a si ngle l ay er of cells Thi s stage is k no wn as the bla s tu la

.

( o r bla s tosp here) ( A B Fig , ,

.

The fo r matio n of the G E R M L A Y E R S is o ne of the m o st im -

po rtan t and sign ifi ca nt p r ocesse s in the w hole cou rse of develop

me nt . G er mlay e rs l ike tho se of L u m bric us an d called by
-

the sa m e name s are fo und in the em br y os of all higher ani

,

m al s ; a nd it w ill hereafter appe ar that thi s fact has a p rofou nd

m ea n i ng .

Dev elopment of th e Org a ns ( g g y

or a n o e n ) The e mbry o grad u . .

all increase s in s i z e an d a t the sa me ti me elo ngate s A s it

y .

le ngt he ns the bl astopore ( in thi s cas e the m o uth) r e mai ns at o ne

,

e nd,
w hich is the r efore to be reg a rded as anterior and the ,

elo ngatio n is b ack w ard s The cell s of all three ger m l ayers
.
-

co nti nually i ncrease in nu m be r by divi sio n new m atter and ,

e nergy bei ng s upplied fro m the food w hich is swallo wed by the ,

e m br yo in such quantitie s as to sw ell up the body like a bl adder .

The ar c he nte ro n e nl a rge s until it co m e s i nto co nt act with the

ectoblast and the segme n tation ca vity is obliter ated -
.

The two pri ma ry m e sobl astic cell s are carried back ward s ,

and al w ay s re m ai n at the extre m e po ste rior en d ( m Fig ,

.

The me sobl as t is in the for m of two b and s l y i ng 011 either side

of the arche ntero n and exte ndi ng forwa r ds fro m the pri mary
,

m e sobl as tic cell s .

Thi s is cle a rly see n in a cro ss section of the e mbry o as in -

Fig 3 6 B G The m e s oblastic b and s are at fi r st so lid but

.
, ,
.
,

after a ti m e a s erie s of p air ed c a vitie s a ppear s in the m con ,

tinually i ncreasin g in n u m ber by the for matio n of new c avitie s

n e a r the hi n der en d of the b an d s as the y i ncre as e in le ngth A .

cro ss sectio n passi ng th ro ugh o ne p ai r of the se c avitie s is shown

-

at B Fig 3 5
,
A s the ban d s le ngthe n they al so exte n d up
. .

w ards an d down w a rds ( 0 Fig u ntil fi nally they meet above

,
.

an d below the arche ntero n The c a vitie s at th e sa m e ti m e .

co nti nue to i ncrease in si z e and fi nall y m eet abov e an d belo w ,

the a rche nte r o n w hich th us beco m e s sur rou n ded by the bod y
,

cavit y o r coelo m ( D ) The cavitie s are sepa r ated by the double

.

p ar tition wall s of m e sobla st These p artitions are the diss epi

-
.

m e nt s and the cavitie s the mselve s co nstit ute the c oelo m

,
The .

outer m e soblastic wall of each cav i ty is kn own as the so ma tic

Myer it u nite s with the ectoblast to co nstitute the bod y
86 THE B I OL O G Y OF A N ANI MAL .

ma top leure) The i er wall o r lcmc hn tc l a

wall ( so nn .
s
p y er ,

m ite with the e tobl t to co n s titute the wa ll of the

(sp l ) u n s
,
n as

alim e tar y
n c an a l (p
s la n ch n cp l e a r e
) A n in g r owth of ectoblast .

s tomod m t ke pl ce i to the bl topore to for m th ph r nx

( c
e u ) a s a n as e a
y ,

s ite extre m it roc todceum

a n d a s i m il a r i n gro w th a t the oppo y ( p )
u n ite s w ith the bli nd en d of the arche n te r o n to fo r m the a n us
and ter mi n al pa rt of the i ntesti n e .

A s to its origi n therefore the ali me nt ary can al co ns ists of , ,

th r ee portions v iz : ( 1 ) the ar che nte r on co nsisting of

,
.
,

FI G . 38 .
—Di a gra m of a i g th r l t i n
r
c o s s sec -
ti on of L u mb ricus , s h ow n e e a o

v ri u rg n t t t h g rm l y r E t b l t i tru t ur h d d w i th fi
a o s o a s, e c .
, o e e -
a e s . c o as c s c es s a e ne

p r ll l l i
a a nt bl
e t i w i th r r p r ll l l i n m b l t i w i th r l i ;
nes, e o as c coa se a a e es , eso as c c oss - nes
al li m
. t ry n l : h hl r g gu l y r ;
c, a en a ca wl m ;
a s m i r ul r m u l
c , c o a o e a e o o c .
-

, c c a sc e s

of b d y w ll ; o m i r ul r m u l f li m t r y w ll ; p l i i g p i th l i um f
-
a c . a, c c a sc e s o a en a a c , n n e e o

al i m t ry l ; d d r l v l ; h y h y p d rm i
en a c a na . v,k i ; L m l gi t ud i n l
o sa ess e , o e s or s n , on a

mu l f b d y w ll ; l m
sc e s o o l gi t ud i n l m u l f li m t ry w ll
-
a . . a,tr l on a sc e s o a en a a : 1 1, c e n a

p rt f
a rv rd ; up nep h ri d i um ;
o ne e co-
h th f rv rd ; p p r i t , l ns, s ea o ne e co- . e, e o n ea

p i th li u m ; r r p r d u t iv
e e rg n ;
, e ub i to ti l v l c e o a s s - n es na es s e .

origi nal entobl ast ; ( 2 ) the sto modseum o r phary ngeal r egio n ,

li ned by e ctobl ast ; and ( 3 ) the proctod aeum or hi nd mo st pa r t ,

a l so li ned b
y ectobl as t The s e three p a rt s a r e c a lled the
.
f or e

g (stomodze um) mid g

.
at u t or m ensentero n ( a rche ntero n) and
,
-

hand g u t ( proctod aeu m d it r e m rk ble f th at the s e

) a n is a a a a c t
-

s am e p art s can be di sti nguis hed in all highe r a ni mal s no t ex ,

ce
ptin
gman .

The body now become s joi nted by the appe ar ance of t rans
v er se fold s oppo s ite the di ss epim e nt s and the m eta me ri s m of th e
,

body bec ome s evident o n the exteri o r The y oung wor m has .

thus re ac hed a stage ( E Fig 3 6) where its re se mbl ance to the , .

 FA TE OF THE GE R M L A Y E R S .

a dult is obvio us I t h as an elo ngated joi nted body tr aver sed

.
, ,

by the ali m e n t a ry c an al w hich ope ns in fro nt by the m o uth and

,

behi n d by the anus The m etameri sm is expre ss ed exter nally

.

b y the joi nted appe a ra nce i n ter nall y by the pre s e nce of p aired ,

c avitie s ( coelo m) separ ated by di ss epi m e n t s B oth the body w all .

-

a nd the ali me n tar y w all co n s i st of two l ay er s : the for m er of

ectoblast w itho ut and so matic m e sobl ast w ithi n ; the latter of
s pl anch n ic m esobla s t w ithout to w a rd s the bod y c avit y) -

a n d either e n tobl as t o r ectobl as t w ithi n accordi ng as we c o n ,

sider the mid g ut o n the o ne h and o r the fore

-

an d hi nd
g u t on ,
-

t h e other This is sho wn in Fig 3 8 which repre se n t s a cro ss

. .
,

sectio n of the e mbr y o th r ough the mid g ut I f this be clearl y -

.

bor ne in m i nd the developme nt of all the othe r organs is e asy to

u n der stand si nce they are for m ed as thicke ni ngs outgro w th s
, , ,

e tc of the part s al read y exi sti ng For i nsta nce th e blood

.
,
.
,

ve ssel s m ake thei r appea rance every where throughout the m e so

b last and the r ep r oductive o r ga ns are at fir st me r e thicke ni ng s
,

o n the so m atic lay er of the m esoblast a fte r w a rd s s ep a ra ti n g ,

m o r e or le ss fro m it so as to lie in the c avity of the ece lo m .

T he n ervou s sy s te m is produced by thicke ni ngs and i ngrowth s

f ro m the ectoblast The origi n of the diffe r e nt parts is show n
.

i n the follo w i ng sche m e

T H E G E R M L A Y E R S A N D T H E I R DE R I V A T I V E S
-
.

The above state m ents as to the o r igi n of the various organs

a cquire g r e at i n tere s t in view of the fact th at the are e s se n
y

h r i di h v be n m i tt d i n th i r p r i e ri gi n i in di put
Th e ne p a a e e o e s ce e ec s o s s e .

I t i rt i n th t th ut r p rt i n f th t u be ( mus ul r p rt ) i n i ngr w th
s ce a a e o e o o o e c a a s a o

f r m th
o t b l t Th l t s t re a rc h s e m t h w th t th nt i r n
e ec o as . e a e se e s e o s o a e e e e

p h id ium h
r t h sa m
as ri gi n th ugh m uth rs d sc ri be th i nn r p rt i n
e e o ,
o so e a o e e e o o

as a ri ingf r m m b l t
s o es o as .
88 THE B I OL OG Y OF A N ANI MA L .

tiall tr ue of all an i mal s above the e a rth w o r m as well as of

y ,

ma yn belo w it— o f all in a word in which the

,
three ger m ,

l ayer s are developed i e all those above the Cwlcn tera ta o r

,
. .
, ,

poly p s jell y fi sh es hy droid s spo nges etc I n m an as in th e

,
-

, , ,
.
,

e arthwor m and all i nter m edi ate for ms the ectobl as t gives rise ,

to the outer ski n (epider mi s) the b ra in and n e r ves fo re and , ,

hi nd g ut ; the e ntobl as t give s ri se to the li ni ng m e m br an e of th e

-

sto mac h i nte sti n e s and other p a r ts pert ai ni ng to the mid g

, ,
ut ; -

w hile the somatic and Splanelmie l ay ers o f the me sobl as t giv e

ri se to the muscle s kid neys repr od uctive organs heart blood
, , , ,

ve ssel s etc I t is no w ge ne r all y held th at the ge r m l aye r s

,
.
-

thr oughout the ani mal ki ngdo m ( w ith the pa r ti al exceptio n of

the Gaeleu tera ta already m e ntio ned ) a r e e sse ntiall y ide ntic al in
origi n and fate Thi s view is k no w n as the G erm la yer Th eory
.
-
.

I t is o ne of the mo st sig nifi c a nt and i mport an t ge nerali zatio ns

w hi c h the st udy of E mbr y ology h as b r o ught to l ight s i nce it ,

re cogni z e s a str uct ural ide ntity of the m ost fundame ntal kin d
a mo ng all the higher a ni m al s .

S oo ner or l ater the y oung earthwor m bu r s ts th rough th e

w all s of the c ap sule and make s its e nt ry i nto the w orld Whe n .

fir st h atc hed it is about an i nc h lo ng and h as no clitellu m .

I t is a cu rious
th a t in cer t a i n speci es o f L u mbric us th e yo ung
f a ct
wo r ms a re a l mos t a l wa ys h a t c h ed a s t wi ns t wo i ndi v i dual s bei ngderi v ed ,

f ro m a si ngl e eggby a process wh ic h is descri bed by K l ei nenbe rgin th e

Qua rterly J u r na l of M icros cop ic a l S c ience Vo l XI X 1 8 7 9
o I t o f ten , . .
, .

h appens th a t the t wi ns a re per ma nentl y uni t ed by a ba nd o f t iss ue as in ,

th e case o f th e well k no wn S ia mese t wi ns

-
.

W e h ave n o w tr ac ed r oughl y the evolution of a comple x

.

many c elled ani m al fr o m a si m ple o ne celled ger m

-

I t is im -

p o rta nt to notice at thi s poi nt a few ge neral p r i nciples which are

true of higher ani mal s in ge ne ral .

1 The e mb ry ological hi stor is a t r ue proce of develop

y ss
.

ment —not a mere grow th or u nfoldi ng of a pre exi sti ng rudi

, -

me nt as the le af is u nfolded fro m the bud N either the ovum .

nor an
y of the earli e r stage s of develop me n t be a rs the sligh te st
re se mblance to an ea rthwo rm The e mbr y o undergoe s a t r a ns .

fo r mation of st ructur e as w ell as an in creas e of s i z e .

2 I t is a p rogr ess fro m a o ne celled to a ma n cell ed con

y
. -
-

dition .
 S UMMA R Y OF DE VE L OPM E N T . 89

3 . I t isa p r ogre ss f r om relative si mplicity to r elative com

lexit Th e ovu m is certain l y vastl y m ore co m plex tha n it
p y .

a ppea rs to the e e but n o one c an do ubt that the full grow n

y
-

wor m is m o r e com plex st ill .

4 . I t is a progre ss fro m a slightl y d iff e r e ntiated to a highl y

d iffere nti ated co n ditio n The life of the ovu m is th at of a
.

single cell The bl astul a is com po sed of a n umber of n e a rly

.

s im ilar c ells which in the gastr ula beco m e d iff e r en ti ated i n to

,

t w o di sti n ct ti ss u e s I n l ate r stage s the cell s beco m e diff ere nti

.

a te d i n to m a ny diffe r e nt ti ss ue s w hich in tu r n bu ild up d iffere n t

,

o r ga ns pe rfo rming u nlike fu nctio ns .

5 . L astl y the develop me n t fo r ms a cy cle begi nn in g with

, ,

the ge rm cell and after m any co m plicated ch ange s re sulti ng in

-

th e prod uctio n of new ger m cell s w hich repeat the proce ss an d

-

give ri se to a n ew g e ner atio n A ll othe r cell s in the body m us t

.

soo ne r o r later die The ge rm cell s alo ne per si s t as the st arti ng

.
-

poi nt to which the c y cle of life co nti n u ally r et u r ns (cf p . .

”
Their p r otoplas m the , g erm
p la s m is the bo n d of co nti nuity
-

t h at links togethe r the succe ss ive ge ne r atio ns .

 C H A P TE R VI .

THE B I OL OG Y O F A N A N I MA L ( Continued) .

Th e E a rt h w o rm .

MI CR O S CO P I C S TR UCTUR E on H I STO L OG Y .

WE have followed the develop me nt of the one celled ge rm -

th rough a stage the bla s tula in which it co ns ists of a mas s of

, ,

nearl y si milar cell s out of which the variou s ti ssu es of the adult
eventu ally a rise The fir st step in thi s directio n is the diff e r e n
.

tiatio n of the g erm la yers o r th r ee pri mitive tissues

-

A s the e mbr y o develop s the cell s of these th r ee tissues beco me ,

dif erentia ted i n s truc t ure to fit the m f o r diffe r e nt duti es in the

phy siologi cal divi sion of l abo r A nd whe n thi s p r ocess of dif .

f erentiation is ac c o mpli shed and the adult s tate is reached w e

fi nd six well mar ked v arietie s of tissue as follow s
-

PR I N C I PA L Tiss ues or L umb icus r .

I . it
E p h el i al. ll s co v eri ngf ree s ur fa ces
L a ye r of ce .

( )
0 P a v em en t E p ith eli u m C ell s th i n a n d fl a t a rr
. a nged l i k e th e ,

s to nes o f a pa v ement .

( b) Columna r E p itheliu m C ell s el o ng a t ed s ta ndi ngs i de b y s i de

.
, ,

pal is ad e l i k e
-
.

( c) Cilia ted E p itheli um C ol umna r o r c ubo id a nd bea ri ngc i l ia

.
,
.

I I Muscular
. C ell s co nt ract il e and el onga t ed to f or m fi bres O f ten
. .

a rra ng ed in pa rall el mass es or bund les .

I I I N erv ous
. C ell s pear s h aped o r irregul a r wi th l arge nucl ei h av
.
-

,
~

ingp roc es ses pro l ong ed i nt o s l ender c o rds or fi b res b undl es o f wh i c h co n ,

s titute th e n er ves .

I V Germina l
. I ncl udi ngth e g
. e rm ce ll s A t fi rs t in th e f o r m o f epi
-

t h elial cell s co v eri ngth e c oel o mic s ur face but a f te r war ds diff erent i a ted ,

i nt o o va and sperma t ozoa .

V B lood
. I sol a t ed cell s or cor puscl es fl oa t i ngin a fl ui d i nte rcell ul a r
.

s ub s t ance th e p la s ma
, .

VI Connect iv e Ti s sue
.
C ell s of diff erent s h apes o f ten b ranch ed b ut
.
,

somet i mes ro unded se a r ated f ro m o ne a no th er b

p , y mo re o r l ess l i f el ess
( i nt ercell ul ar) s ubs t ance in th e f o r m o f th reads o r h o mog eneo us ma t eri a l .

90
 A R R A N GE M E N T OF TI S S UE S . 91

These six ki nd s of ti ssue con stit ute the main bulk of the
e arthwor m as of higher an i mal s ge nerall y ; but there are in ad
,

ditio n other ti ssue s which w ill be treated of hereafter .

A rr a ng ement of th e Ti ssu es The si m ple st and m o st direct .

m ode of di scoveri ng the a rra nge m e nt of the ti ssue s is by the m i

c rosc o pic al study of thi n tra n s ver s e o r lo ngit u di na l s ectio n s A .

FI G . 39 .
—Tra n s v ers e sec t i o n o f
b h i nd th l i t ll um
th e b o dy e e c e . a c, c a
. vi ty f th lio e a

m n t ry
e l ; ut i l ;
a c a na l m ; m i r ul r m u l ;
c, c c e ca
'
, c oe o c . , c c a sc e s c m, c ir ul r v l ;
c a es se

d r l v l ; h y h y p d rm i ; l m l gi t ud i l m u l ;
o sa e ss e , o e s . , on na sc es n c,
. v tr l rv
en a ne e

c h i p p ri t
a n ; .l p i th l i um ;
c, e o n eat ; et i g r gl
e s. s e a se e o us a nd ; s .i . ub i t
v, s -
n es

ti i v l; m mu l
ns es se s .t i gth t w gr up f t
, sc e c o nn ec n e o o s o se a; o n th e sa m i d ; tu
e s e ,

t y p hl l o so e .

tr ansver se section t aken th rough the regio n of the sto mach

i nte stin e is repre se nted in Fig 3 9 I ts co m po s itio n is as . .

follow s
A . B O DY W A L L -
.

Thi s consi s ts of five l ay e r s v iz (begi nn i ng with the out , .

s ide
) ,

1 Cu ticle (c)
. A ve ry thi n t ransp a r e n t m e mb r ane n ot
.
,

compo sed of cell s and pe r fo r ated by fi ne pore s I t is a p roduct .

o r secr etion of the

92 THE B I OL OG Y OF A N AN I MA L .

2 Hyp .
m is ( y) (
h epide rm is
oder o r sk in ) A lay e r of colu m .

m co m po ed of eve r l kin d of elo n gat e d ce ll s , set

nar epitheli u s s a s
,

ve rticall to the sur face of the body S ome of the se , k n ow n as

y
.

g
la n d cells , h ave the po w e r of p r oduci ng w ithi n their substa nce
-

a
gl air y
fl uid ( m uc u s
) w hich ex ude s to ,
the ex te r ior through the
pore s in the c uticle O ther s ( se nsory cell s) give ofi f r o m thei r
.

i nne r e nd s nerve fi bres which may be traced i nwards to the

-

gangli a ( Fig .

C lit ll m i r d u d b n e n o r mo us th i c k e ni ng o f th e h ypoder
T h e e u s p o ce y a

mis ca us ed especi all y by a g

,
rea t dev el opment o f t h e gl a nd cel ls Th ree -
.

f o r ms o f th ese may be dis t i nguis h ed wh i c h pro ba b l y produce different ,

s ec ret i ons Th e t i ss ue is permeated by numero us mi nut e b l ood vessels

.
-

wh i c h r ami fy bet ween th e cell s .

a er of pa r allel m u scle m) A
3 .
(
Circ ula r llf uscle s l y cu

fibre s r unni ng a r o u nd the b o dy O n the up pe r side they ar e .

i nter mi ngled with c onn e ctive ti s sue cell s c o nt a i ni ng a gr anular -

browni sh sub stance ( pigment ) which gives to the do r sal as pect

its da rke r ti nt .

4 L o ng .itudi n a l M uscles A l ay e r of mu scle fi bres -

r unni ng le ngthwi se of the body They are a rr a nged in compli .

c ated b undle s w hi c h in cro ss s ectio ns have a feather y appea r

-

ance I n lo n git udi nal sectio ns they appea r as a s i m ple lay e r and
.
,

r e se mble the circ ul ar fibres as seen in the cross sec tio n -

The cir cul ar mu scle s are ar ranged in so mewhat similar bun

dle s as may be see n in longitudi nal secti o ns
, .

5 Ocelo m ic or P erito nea l E p itheliu m

. A ver y thin
l ay er of fl atte ned cell s next the c oelo mic ca vi ty .

The h ypoder mi s and therefo re al s o the cuticle to which it

,

gives ri se is deri v ed fr o m the ectobl as t The other laye r s ( 3

, .
,

4 5 ) a ri se fro m the s o matic l a e r of the m es oblast

, y .

B . A LI M E N TA R Y CA N A L .

The wall of thi s tube appe ar s in c ro s s secti o n as a r i ng su r

‘
-

r o unded by the c oelo m The t vphlo so le ( ty) is see n to be a deep

.

i nfoldi ng o f its uppe r porti o n I n the m iddle regio n the wall is .

co mposed of five l ayers as follow s s ta rti ng fro m the alim e nta ry ,

c avity (Fig 4 0) .

1 L in ingE p i theliu m e
.
( p ) A l ayer of closel y pac k ed nar .
,

r ow ciliated colu mnar cell s w ith oval nuclei .

2 Vas cula r L a y er v l m
.

( ) N u ero us m i nute blood vesse ls u

-
.
 HI S TOL OG Y OF THE A L I M E N TA R Y CA N A L . 93

3 . Ci rc ula r M uscles ( c . m) A thin laye r of m uscle fi bres -

r u nni ngar ou nd the g

ut .

4 . L ongitu d ina l M u scles A th in l ay e r of muscle

fi bres r u nni ng al o ng the gut .

5 Chlora g
. og ue L a y er ( ch) C o m po s ed of la r ge pol y hed r al .

o r r ounded cell s co n t ai ni ng yell o w i sh gree n granule s The cells -

fill the holl ow of the t y phlo sole and cover the su r face of the ,

d o r sal an d l ate r al blood ve sse ls T h is lay e r r ep r ese nts the -

s plan clmic pa r t of the pe r ito n eal epitheliu m .

Th e sa rrangement ex is t s in all pa r t s o f the al i ment a ry

me g
ene ra l a

c a na l b ut is so me t i mes g
,
rea tl y modi fied Fo r i ns t ance th e gizza rd and .
,

p h a r y n x a re l i ned by a t o ug h t h ic k cut i cl e a nd t h e m usc ul a r l a ers a e

y , , r

e no r mo us l y de v el oped I n a pa r t o f th e g iz za rd th e c hl o ragogue l a yer is

.
-

near l y o r q ui te a bs ent a nd th e t yp hl os o l e di sa ppea rs A f ull er desc ript i on .

o f th ese modi fi c a t i o ns wi ll be fo und in B roo k s s Ha nd bo o k of I nvertebra te

’

Zoolo g y a nd a co mpl et e ac co un t in C laparede Zeitsch rif t f ur wis s en

, ,

Zoolog
“
salsa/ Clic h e ie Vol XI X, .
,
1 86 9 .

The li ni ng epitheliu m is deriv ed f r om the e ntoblast The .

rem ai ni ng l ay e r s a r ise by diffe r e ntiatio n of the spla nch ni c lay e r

o f m eso blast .

FI G . 40 .
—Hi ghl y m agn fi ed i
t i n th r ugh th w ll f th l i m n t ry n l c r o s s s ec -
o o e a o e a e a ca a .

h
c hl r g gu l y r ; m i ul r m u l ; m li i g p i th li um ; l m l gi
, c o a o e a e c .
, c rc a sc es c , n n e e .
, on

t u d in l m u l ; I v ul r l y r
a s c es v . .
, a sc a a e .

B lood v essels appear in the sectio n as r o un ded o r i r regu l a r

-

cavitie s bounded by thi n w all s They co nsi s t of a delicate lin i ng .

epitheli um covered by a thi n l ay er of muscle fi bres I n the -

.

w all s of the sto mach i nte st in e the ve ssel s are ofte n co mpletel y -

in vested by chlo r a gogue cell s which r adi ate f r o m the m with -

,
.
94 THE B I OL OG Y OF A N ANI MA L .

gre at r egul ar ity ( Fig The fi ner branches have no muscu .

lar l ayer co nsi sting of the epitheliu m alo ne

,
.

Di ssepi ment s These ofte n appe ar in cross o r longitudi nal

.

sections They consi st chiefly of mu scle fi bres i rr egul ar ly dis

.
-

posed i nter mi ngled w ith co nnective tissue cell s and fibre s and
,
-

covered o n both side s with the peritone al epitheli um .

N erv ous Sy st em A cro ss sectio n of a ganglion ( Fig 4 1 ) .

-

s how s it to be co mpo sed of two disti nct pa rt s v iz ( ) the

1 ,
.
,

FI G . 41 .
—Hi ghl y m a gn i fi ed c ro s s -
se c ti o n of a v tr l g ngl i n gf gi n t fi b ; Ln
en a a o . ., , a - re s ,

t r l rv ; n
la e a rv ll ; m u
ne e . c, n e e ce-
s s, sc ul r h
a th f t h g n gl i n ;
s ea o ub n u e a o s -
e

ralv l ; n upr n ur l v l
ess e s . . v, s a -
e a esse .

glion proper on the i nside and ( 2) a sheath which e nvel ops it ,

.

The sheath (8 Fig 4 1 ) consi st s of two l ay er s v iz

, .
, .

1 P eri to nea l E p ithelium

. O n the out side . .

2 I ll usc ula r L a yer o r sheath a thick l ay e r of i rr egula r ly

.
, ,

a rr anged mu sc le fi bres i nter m i ngled w ith c o nn ective ti ssu e

-
Im .

bedded in it are the sub ne ur al blood vessel o n the lowe r side - -

an d the supr a n eu ral blood vessel s o n each s ide a bove

-

I n the -
.

m iddle li ne are three ro un ded space s ( 9 f Fig w hich a r e , ,

.

the c ro ss sections of three hollo w fibre s ru nni ng alo ng the e ntire

-

“
le gth of the ve ntr al ne rve chai n They a re called
n gi ant -

fi bres an d po ssibl
, y se rve to suppo r t the soft p a rt s of the nerve
cord .

The Ga ng lion p r ope r is di sti nctl y bilobed and co nsi st s of ,

tw o po r tions v iz , .

1 N erv e cells.
N u m erou s pear sh aped n erve cell s n ear
-
- -

the surfac e w ith their narro w e nd s t ur ned to wa rds the ce ntre

, ,

i nto whi ch eac h se nd s a si ngle b ranch o r ne r ve fi bre They are -

.

co nfi ned chiefl y to the ve nt r al and l ate ral pa r ts of the ganglion .

 HI S TOL O G Y OF THE N E R VO US S Y S TE M . 95

2 . Fibrous P ortion
This occupies the cen t r al part I t .
.

co nsi st s of a c lo se and com plicated n et w ork of n erve fi bres i nter -

m ingled with c o n nective ti ss ue S o m e of the se fib r e s co mm u ni .

cate with b r anche s of the n e r ve c ell s as st ated above ; other s -

run out i n to the l ateral ne r ve s w hile still other s run alo ng th e ,

to co nn ect with fibre s fro m othe r ganglia .

FI G . 42 .
—Two of th e v nt
e ra l ga n gli a l t r l n rv
(I , I I) of L umb ricus wi th th e a e a e es,

s h wi g m f th m t r n r v ll d fi b r ( b l k )
o n so e o e o o e e ce d
-
fi br f s an es ac . a sen s es or

w rd a nd b s ak w d w i th i t h
ac ar r v r d ; b fi br i t
s n e f th
ne d ub l e co -
, a e n o o ne o e o e

n ver it w
es o n i d : 0 nd d fi br th t r t t h rv f th pp i t i d
s o n s e a , es a c o ss o e ne es o e o os e s e .

( A f t r R tz i u )
e e s .

A cco rdi ngto th e l ates t resear c h es ( o f L enh o ss é k a nd R et zi us) mo s t if

n o t all o f th e ner v e cells o f t h e v ent r a l c o r d are mo t o r in f unc t i o n
-
N ear .

th e cent r e o f eac h g a ngl io n ( Fig 4 2 e) in a s i ngl e l a rge mult ipo l a r c ell o f

.
,

do ub t f ul na t ur e A ll t h e o th er cell s are ei th er b i po l a r o uni po l a r in th e

. r ,

l a tt er case sendi ngout a si ngl e b ra nc h wh i c h soo n di v i des i nt o t wo I n .

e v er y cas e o ne o f t h e b r a nc h es b rea k s up i n t o fi ne s ub di v i s i o ns wi th i n th e
-

c rd
o Th e o th er b ra nc h in mos t cases pass es o ut o f t h e co rd th ro ug h one
o f t h e l a t er a l nerves to th e musc l es o r o th er peri p h er a l o rg a ns ei th er ,
96 THE B I OL O G Y OF A N ANI MA L .

r i i th i r h i t i d o f th e body or ma k i ngexit
c oss ng d
w n t h e co t o t e o pp o s e s e

co mmiss ura l
”
o n its o w n s i d e S o m e o f t h e ce ll s h o wever are purel y
. , , ,

nei th er b r a nc h l ea v es th e co r d .

r fi b r t ri f r m ri h r t er mi na t e f reel y ( not in
h
T e s e n s o y es e n e n g o t h e p e p e y
r v ll b r k i i t m erous fi ne b ra nc h es o n th e s ame s ide of
ne e c e s ) -
ea n gp u n o n u
,

t h e co rd ( Fig . .

The ner v es l ea v i ngth e cent ral system are mixed xc t hey contai n both , . .
,

s ensory a nd mo t o r fi b res .

FI G . 43 vr .
-
t i n f v n t r l p r t f th b d y h w i ng t h n rv u n
T r a ns e se sec o o e a a o e o , s o e e o s co

n ec ti n v tr l g ngli n givi ng il l t r l n v t l n ; p p r i t n l
ons . . c, en a a o , o a a e a er e a . . . e
.
, e o ea

ep i th l i um ; L m l gi t ud i l m u l : b y h y p d rm i ;
e t A i gl m t r
on na sc e s , o e s 8, se a . s n e o o

n rv e ll b l k) i h w
e ce-
nd i g fi br i nt th n rv t w d th l f t I n
( ac s s o n se n a e o e e e o ar s e e .

th rv t t h r i ght
e ne ry fi br pr d i gi nw rd f m t h n ry ll
e o e a re s e n so es oc e e n a ro e se so ce s
(b l k ) f th h y p d rm i nd t rm i t i g in br h i g x tr m i t i
ac o e ( A ft
o e s, a e na n a nc n e e es . er
L enh os s é k ) .

t ions th rough th e v ent ral commiss ures are s i mi l ar to th ose th rough

S ec
t he g a ng l ia but th e cent ra l po rt io n , th a t wi th i n th e s h eath ) is s ma ll er ,

i s di v i ded i nt o t wo dis t i nc t pa r t s a nd th e ner v e c ell s a re l ess a b unda nt ,

-
.

S ec t i o ns th ro ug h th e ner v es s h o w th em to co ns is t o nl y o f pa rall el fi b res

s urro unded b y a s h eath wh i c h
graduall y fades a way as t h e ner v es g ro w
mall er and fi nall y di sappea rs th e mus c ul a r l ay er fi rs t disa ppeari ng and
, , ,

t h en t h e epi th el i a l c o v eri ng .

With thi s brief sketch of the hi stological st ructu r e of the

e a rth w or m w e c onclude ou r m orphologica l stud of the a ni m al
y .

Tho se wh o de sire fulle r i nfor matio n o n the histolog will fi nd a

y
g e n e ral tre at me nt of it in the work of C la are
p d e al r ead
y ci t ed ,

at p 93 Many l ater wo r ks have bee n publis hed o n the de

. .

t ailed hi stology .
 C HA PTE R VI I .

THE B I OL O G Y OF A N A N I MA L ( Con ti nued)

Ph y s i o lo gy of th e E a r t h w o rm .

I N the p recedi ng pages brief de sc r iptio ns of many special

ph ysiological phe no me na have bee n gi ve n i n co nnectio n with th e
detai led desc r iptio ns of the p r i mar y functions and syste ms I t .

n ow r e m ains to co nsider the m ore ge n e r a l proble ms of the life of

the animal and e speciall y its r elatio ns to the e nviro nm e nt and

, ,

the t r ansfo r mati o ns of ma tte r and e ne r gy w hich it effect s .

Th e E art h wor m and its E nvi r onment The ea r thwor m is an

.

o r ganiz ed mass of livi ng m atte r occupying a defi n ite po sitio n in

s p ace and ti m e an d exi sti ng am id cert a i n de fi n ite an d ch a r acter
,
”
istic ph y sica l s urrou n di ngs which co nstit ute its e nviro nm e nt .

A s o r dinaril y u n de r stood the ter m e n viro nm e n t applie s o nl y

to the i mm edi ate su r roundi n gs of the an i mal — to the e a rth
t hr ough whi ch it bu r ro ws the ai r an d moist ure th at b athe its
,

su r face and the like S tri c tl y speaki ng howeve r the e n viro n

,
.
, ,

m e nt i ncl ude s eve ry t hing th at m ay in any manne r act u po n th e

o r ganis m—th at is the whole u n iver se out side the w or m Fo r
,
.

the ani mal is di rectl y and profoun dl y aff ected by r ay s of ligh t

an d h eat that tr avel to it fro m the s un ; it is extre m el y s e nsiti v e

to the alter natio n s of day an d n ight and the season s of the y e ar ;

,

it is acted o n by gr avit y ; and to all the se as w ell as to m or e ,

imm ediate i n fl ue n ce s the an i mal m ake s defi n ite respon ses

, .

We h ave see n th at the bod y of the earth w o r m is a co m pli

cated piec e of m e c h ani sm co n structed to perfor m cert ai n defi n it e

actio ns B ut ever y o ne of these actions is in o ne way or an

.

othe r depe n de nt upon the env ir o nm ent and di r ectl y or i ndirectly

r el ate s to it A t eve ry m om ent of its existence the organi sm is
.

acted on by its e n vi r o nm en t ; at eve r y m om en t it r e act s u pon

the e n vironm e nt m ai n tai n ing with it a constantl y shifti ng st at e
,

of equ ilibri um w hich fi nally gives way onl y whe n the life of th e
ani mal draw s to a clo se .

A daptati on of th e Or g ani sm to its E nvi r onment I n its r ela .

tions to the en viro nm e nt the ea rthwo rm embodies a fu n dam e nt a l

97
biologic al law , v iz .
,
th at the liv in g g m m u s t be adap ted to
or a n is

i ts en v iron men t, or , in othe r word s, th at a ce rtai n [ta rmo ny

betwee n organi sm and envir onme nt is esse nti al to the co nti nu
ce of life d i fl e ce w hich te n d s to di stu r b or de stro
a n an a n
, y n u n y
thi s har mony tends to d ist urb or de stroy life The adaptation .

m ay be either p ass ive ( str u c tu r a l ) or ac tive ( f u n ctio nal

) S t rue .

t ur al ad apt ation is w ell ill ustr ated fo r i nstance by the ge ner al , ,

s h ape of the bod y so we l l ad

,
a pted fo r bu r rowi n g thro u gh the
e a rth . A gai n the delicate i ntegum e nt gives to the b od y the
,

flexibility dem anded by the pec uli a r m ode of loco m otio n ; it

a fford s at the sa me ti me a highl y fa vor able r e s pi r ato ry su r face

a m atter of no small i mpo r t ance to the w o r m in its badly venti -

l ated burro w ; and y et thi s delic ate i ntegu m e nt doe s no t lead to

d e sic c atio n bec ause the ani mal live s alwa v s in co n tact w ith m o ist
,

e a rth .
The ali me ntar y canal lo ng and co m plica ted is m os t
, ,

perfectl y fitted f o r worki ng over and ext rac ti ng nut r i me nt from

t h e e a rth y diet The r eprodu c tive organs are a re mar k able ln
.

s ta n ce o f co mplex str uc t u r al ad a ptati o n in an a ni mal whi c h o n

t h e whole is of c o m p ar ativel y si mple structure .

F unctional ad aptation is perh ap s bes t sh ow n in the i nsti nctive

“ ”
a ctio ns or h abit s of the wo r m I ts noct u r nal m ode of life .

( f u n c tio nal ad apt atio n to light

) a n d its ti m id i t y p r o tect it

from he at de siccatio n from bird s and othe r e ne m ies I n win

, , .

t er or in seaso ns of drought it b urro ws deep i n to the ea rth .

A striki ng i n st anc e of adapt atio n is s how n in the c ar e whi c h

i s t aken to i nsure the welfa re of the e m br y o wor ms M in ute .
,

d elic ate and h elple ss as they are they devel o p in safet y i ns ide
, ,

t h e tough le athe ry cap sule ( p

, fl o ati ng in a mi lklike
.

liquid w hich is at o nce their cr adle and thei r fo o d .

Ori g in of A dapt ati ons The develop me nt of th e e art h w o rm

.

s how s th a t its whole c o m plex bodil m ech a ni sm t ake s o rigi n in a

y
s in gle cell
(p 7 and th at all the re ma rk able ad a ptatio ns ex
.

pre ssed in its stru c ture and actio n are brought abo ut by a g ra du al

p r oce ss in the life hi sto r of e ch i ndivid u l wor m The r e is

y a a
-

reas on to believe th at thi s is ty pi c al o f the a nce str al history (de

)
s ce nt of the spe c ies as a w h o le an d th at ad a pt atio n h as bee n
,

g r ad uall
y acquired in the p as t We k no w th at e nv ir o nme nts
.

ch ange and th at to a certai n exte nt organi sms ch a nge corre

,

s pondi ngl
y through f unctio nal ad aptatio n provided the ch ange of ,
 N UTR I TI ON OF THE ANI MA L . 9

en vi r onm ent be not too sudde n or ext r e m e I n other word s .

the organism po sse sse s a cert ai n p la s ticity w hich e nable s it to

ad apt it self to grad uall y changin g co nditio ns of the en viro nme nt
-

N o w there is good reaso n to believe that as e nviro nm e n t

has gr adu all y u n dergo n e cha nge s in the p ast o rganis ms have ,

gradually un de r go ne cor re spo nding ch anges of str ucture Tho se .

whi ch h ave beco m e in any way so m odifi ed as to be m o st per

f ectly ad apted to the ch anged e nviro n m e n t have te nded to sur
v ive an d le a ve s i m ilarl ad apted de sce n d an t s Tho e which
y s
-
.

h ave bee n le ss pe r fectl y adapted h av e te nded to di e out t h r ough

lac k of fit ne ss fo r the e nv ir o nme nt ; and by thi s proce ss—called
“ ” “
by Da r w i n N atu ral S electio n and b
y S pe n cer the S urviv a l
”—
o f the Fittest the r e mark able ad aptatio ns eve ry where met
wi th a re believed to h ave bee n grad ually wo r ked out .

It h o ul d be obser v ed th a t N at ura l Sel ect i on do es not r ea ll y ex pl ai n th e

s

orig in of ada pt a t i o ns b ut o nl y th eir per si s t ence a nd acc umul a t i on

, The .

th eo ry o f ev ol ut io n is no t at pres ent s uc h as to ena b l e us to sa y wi th cer

ta i nty wh a t c auses th e fi rs t o ri gi n o f adapt i ve vari a t i o ns .

N ut r iti on . The earth w o r m does work I t wo r ks in travel

.

ling about and in fo r ci ng its way through the soil ; in sei z i ng ,

s wallo wi ng dige sti ng and ab so r bi ng food ; in p um pi ng the

, ,

blood ; in mai nt ai ni ng the ac tio n of cilia ; in receivi ng and se n d

'

ingout ne r ve i mp ul se s ; in g r o wi ng ; in reproduci ng it self— ia

-

s ho r t in c a rr y i ng o n any an d ever y for m of vital actio n

,
To .

live is to w ork N o w w ork i nvolve s the expendit ure of e n ergy

.
,

a n d the an i mal bod y l ike any other m achi n e

,
w hile life c o n ,

tinues r equi r es a co n ti nual suppl y of e nergy

,
I t is cle ar fro m
.

wh at has b ee n said o n p 3 2 that the i mmediate sou rce of the

.

e ne rgy expe n ded in vit al actio n is the worki ng protopla sm it self ,

w hich u n dergoe s a de st r uctive che m ic a l ch ange (k a taboli sm o r

d e str uctive m et aboli sm) h avi ng the na t ur e of an oxid atio n Fro m .

t h is it follow s o n the o ne h an l th at the w a ste product s of thi s

a ctio n must be ulti matel y p assed o ut of the bod y as exc r etio ns ,

and o n the othe r h an d that the lo ss m u st ultim atel y be m ade

good by fre sh supplie s e nteri ng the ani mal in the for m of food .

I t is further evide nt that the i nco m e m ust equal the outgo if the
a n i m al is m erel to hold its own and m us t exceed it if the ani
y ,

m al i s to g r ow .
1 00 THE B I OL OG Y OF A N A N I MA L .

Thu s it come s about th at there is a mo re o r le ss stead y fl ow

of matter and of e nergy through the li v i ng orga ni sm w hich is ,

it self a centre of ac tivity like a whirlpool ( p

,
The che m ical .

phenome na acco mp anyi ng the flow of m atter and e nergy th r ough

the organism are those of n utrition in the w ide s t se nse Thi s .

te r m is more often rest ricted e spe cially to the phe no me na aeco m

p anying the in co me w hile those pe r t ai ning to the ou tgo are
,

r egarded as belonging to ea cretion The i nte rmediate p r ocesses

'
.

directly connected wi th the l ife of p rotopl as m a re put togethe r

unde r the he ad of meta bolis m ; they i ncl ude bo t h the co ns t rue
tive processe s by whi c h protopl as m is built up ( a i mbolism) and
the de structive pro c e sse s by which it is bro ke n dow n (k a ta bolis m)
in the libe ratio n of e nergy .

I ncome
. I t is diffi c ul t to dete r m i ne the exac t i nc o m e o f
L umbricus but it may be set dow n app r oxim atel y as follow s
,

IN COM E O F L UMB R I C US .

The food stuff s are conve rted by the ani mal into the sub
-

st ance of its own bod y (protopl as m an d all its de riv atives ) and ,

they must therefore be the ulti mate so urce of e ne r gy I t fol .

lows th at the ani mal takes in e nergy o nl y in the po te ntial fo rm

in the che m ical pote nti al bet w ee n the oxidi z able p ro teid s
,

carbohydrate s and fats and free oxyge n) I t is tr ue that the

, .
 DI G E S TI ON A N D A B S OR P TI ON . 1 01

animal may un de r ce r tai n ci r c umstance s abso r b ki netic e ne r gy in

the fo r m of he at but thi s is avail able on ly as a condition not as
, ,

a ca us e of p r otopl as mic ac tio n I n thi s in ability to use kin etic .

e nergy the earthwor m is ty pical of anim als as a whole .

O f the orga nic po rt io n of the food proteid s are a s ine g ua

non and in this re s pect again the wor m is a ty pe of an i m al life

,

in ge n er al E ithe r the fats or the c a rboh y d rate s may be o m itted

.

n im al p r obabl y th rives be st upo n a m ixed diet in

( though the a
which both a r e pre se n t) but without proteids no anim al as f ar , ,

as is kn ow n can long ex ist,

.

Gener al Hi st ory of th e Food Di g esti on and A b sor pti on . .

L u mbri cus take s dail y i nto its ali m e ntar y c a nal a certain am o un t
of n ecessary food stuffs but the se are not re ally i nside the body
-

so lo ng as the y re mai n in the ali m e n ta ry ca nal ; fo r thi s is show n

by its develop m e n t to be o nl y a p a rt of the oute r su rface folded
in to aff ord a sa fe r ecept acle w i t hi n which the food may be
wo r ked ove r B efo r e the food c an be ac t ually take n i nto the
.

bod y o r absorbed it mu st un dergo ce r ta in che mic a l ch ange s col

, ,

lec tiv ely called d ig es ti on ( of p A ver y i mp o rtant p a rt . .

of thi s p r ocess co nsi st s in r e nde r i ng non diffu sible sub st ance s dif -

fus ible in order th at they may p ass through the wall s of the
,

a lim e n ta r ca nal i nto the blood P r oteid s fo r ex am ple hav e

y .
, ,

bee n shown to be no n difi u sible (C hap I I I ) I n dige stio n they

-
. .

are cha nged b the fl uid s of the ali me n tar y ca nal i n to p ep to n es

y
—sub sta nces much like proteid s but r eadily diff usible I n , .

like manne r the n o n diffu s ible s t a rch is ch anged in to diffusible

-

s ugar and beco m e s cap able of ab s orptio n I t is highl y pr obable .

th at all ca r boh y d r ate s are thu s t u r ned i nto sugar The fat s ar e .

p r ob abl y co nve rted in par t in to soluble and d ffusible so ap s w hich

i

are readil
y absorbed but are m ain l
y e m,
u l s ified an d di r e c tl
y p ass ed
i nto the cel ls of the ali me nta r y tract in a fi nelv di vided state .

N othing however is k n own of thi s sa ve by a nalogy with highe r

, ,

a ni mals I n all cases dige stion take s pl ace ou ts ide tbc body an d
.
,

is o nl y p r elimi na ry to the real e nt r an ce of food into the phy sio

logica l or tr ue inte r io r
, ,
.

Metab oli sm A fte r ab so r ption into the body prope r the

. .

i nco m i ng matte rs are dist r ibuted by the circulation to the ulti

m ate livi ng u n its o r cell s and a r e finally taken up by the m an d ,

b uilt in to thei r substance There is r e aso n to believe that each .

102 THE B I OL OG Y OF A N ANI MA L .

cell takes fr o m the co mmon ca rrie r the blood only such ma

, ,

terials as it n ee d s
,
lead i ng a so m e w hat in depe n de n t life as to its

ut r itio I t pe rates with othe r cell s u n de r the direc

o wn n n c o 0
-

tion of the ne r vou s sys te m ( co o r dinatingmech anis m) but to a ,

gr eat degree is i ndepende nt 1n its choice of food— ju st as a sol

di er i n a well fed a r m y obe y s o r de r s fo r the co mm o n good but,

et t a kes o n l y what he choose s f r o m the d a il y r a tio n su pplied to

y
all .

Wh at take s place withi n the cell upo n the e nt rance of the

food is al most wholly u nk nown but so meho w the food matt e rs
,
-

rich in pote nti al e nergy are buil t u p i nto the li vingsub stance
,

prob abl y by a serie s of const r uctive p r ocesse s cul mina ti ng in pro

toplas m . A longs ide the se co nst ructive p r ocesses ( anabolis m) a
contin ual dest r uctive ac tio n goes o n ( k atabol is m) ; for living mat
ter is decom posed and energy set fr ee in eve ry vital actio n and ,

vit ality or life is a continuo us p r ocess I t mu st not be su pposed

.
,

howeve r th at eithe r the synthetic o r the de st ru ctive p r ocess is a

,

si ngle ac t . B oth p r ob ably in volve lo ng and co m plica ted che m i

cal tr ansfor mations but the prec ise nat ur e of these chan ge s is at
p r e sent al mos t wh olly u nk no wn I t is ce r ta i n that the d es t r u o
.

tive action is in a ge neral way a p r ocess of oxidatio n e ff ected by

aid of the f r ee ox y ge n take n in in r e spir atio n . We may be
s ur e how ever th a t it is no t a case of s i m ple co m bus ti o n
, ,
the
p r otopl asm is not I t is more probably anal o gou s to
an explo sive actio n ,
the oxyge n fi r st e nter ing i nto a loose
ciation wi t h co m plex o rga n ic sub st an ce s in the p r oto p las m and ,

then suddenly co mbi ni ng with the m un de r the app r op riate s tim

ulu s to form si mpler and m ore highl y o xidiz ed p r oducts O f
-
.

the precise nat ur e of the proc e ss w e ar e quite igno r ant .

Out g o .Just as the i nco me of the an i mal r eprese nts o nl y the

fir st term in a se ries of co nst r uctive processes so the outgo is ,

t h e l as t te rm of a se r ie s of de structive ac tio ns of which we r eall

y
kn ow very little save th rough thei r r esults The outgo is shown .

in the acc ompany ing table .

B oth energy and matt e r leave the ce lls and fi nally l eave the
bod y—the for mer as heat work done o r e ne rgy still pote ntial
,

, ,

( in urea and other organic mat te r s) ; the latt e r as exc retions ,

w hich diffuse freely outwa rds through the skin and n eph r idial
s u rfaces .
 THE A N I MA L A N D I TS E N VI R ON M E N T . 03

Of the d aily outgo the water c arbo n dioxide and salt s are
, ,

d evoid of e n ergy but the u rea co nt ai ns a s mall am ou nt w hich is

,

a shee r lo ss to the a n i m a l .Were the e a rth w or m a pe rfect ma

c hi n e it co uld use thi s r e s id ue of e n ergy by deco m po si ng the u re a

i n to si m pler co mpo und s [v iz a m m o n i a ( N .

,
c arbo n dioxide
an d w ate r but it l ack s this po w er though there ,

a re ce rta in organ i s ms ( B a cte r ia ) which a r e able to utili z e the last

t races of e nerg y in u re a ( p . To the d a il

y o u tgo m u s t be

a dded the occasio n al lo ss both of m a t ter and of e n ergy suffe r ed

in giv ing r ise to ova a nd s per mato z o a an d in pro v idi ng a ce rt ain

,

a m ou n t of food an d p rotectio n for the n ext ge n er atio n .

I nt er acti on of th e A nimal and th e E nvi r onment The actio n .

o f the e nviro nm e n t upo n the a nim al has al read y bee n su fficie n tly

s tated ( p . I t r e m a i ns to poi nt o ut the ch ange s w orked by

t h e animal o n the e nviro nm e nt These ch ange s are of two
.

kinds m ech anical (or ph y sical ) and che m ical The mo st im por
,
.

t a nt of the form er is the co nti nual t ra nsfo r m atio n of the soi l

which the wor ms e ff ect as D ar win sho w ed by b r i ngi ng the
, ,

d eepe r l ay er s to the surf ace w he r e they are expo sed to the at

,

m o sph ere and also by dr aggi ng supe rfici a l object s i nto the b ur
,

ro w s. The che mi cal ch anges are still m o r e s ig ni fica nt The .

1 04 THE B I OL O G Y OF A N A N I MA L .

ge neral eff ect of the metaboli sm of the an i mal is the de st ru ction

by oxid ation of orga nic m atte r ; th at is m at te r origi nally take n
,

fro m the e nvironme nt in the for m of com plex p r oteid s fats and , ,

c arbohydr ate s is r et urned to it in the fo rm of sim ple r and mo re

highl y oxidi z ed sub stance s of which the mo st impo rtant are ca r
,

b on di oxi de and w ater ( both i norgan ic sub s ta nce s) This action

.

further more is accompanied by a di ssipatio n of e ne r gy—that is ,

a co nversion of pote nti al i nto k inetic e ne r gy .

O n the whole therefore the actio n of the a n imal u po n the

, ,

e nvironm ent is th at of an oxidi z i ng age nt a r educe r of comple

,

c o mpou nd s to si mple r o ne s and a di ss ipato r of e ne rg

, y .

he rei n it is typical of animals in ge ne ral .

 C HA P T E R VI I I .

THE B I O L OGY OF A PLA N T .

Th e C o m m o n B r ak e o r Fe r n .

( Pteris aq uili na L i nn us )
, ae .

FO R the stud y of a r ep r e se ntative vegetal o r ganis m so me

plant should be chose n which may be r eadil y procu red and is
neithe r v e ry h igh no r ver y low in the scale of o r ga ni z atio n .

S uch a pla nt is a co mm o n fe r n
“
.

Fer ns g r ow ge ne r all y in da mp an d shad y place s though ,

n o m ean s c o nfi n ed to s uch loc alitie s S o m e of the .

m ore ha r dy s pecies p r efer dry r ocks or eve n bold cliffs in the ,

c r evices of which the y fi nd su pport ; other s live in O pe n field s

o r fo r e s ts an d s ti ll othe r s o n sa nd y hills ide s

,
I n the n orther n .

U nited S tates there ar e altogethe r s o me fifty specie s of wild

fer ns but those which are co m mo n in any p articul ar loc ality are
,

seldo m mo re tha n a sco r e in n u m ber Th roughout the whole

.

wo r ld so m e fou r tho usa n d specie s of fer ns are k no wn but by ,

fa r the greate r numbe r are found o nl y in t r opic al r egion s w here ,

the cl imate is best s uited to the ir w ant s A t an e a rlier period .

o f the e a rth s hi s t o r y fer ns attai ned a gre at s i z e a nd for m ed a

’
,

c o nspicuo us an d i mpo r ta n t fe at ure of the veget atio n At .

pre se nt how eve r they are fo r the m o st part o nly a few feet in
, ,

height N earl y all are pe r e nn i al ; th at is they may l ive for an

.
,

indefi n ite n u mber of y e a r s M o st of the m h a ve creepi ng o r

.

subterranea n ste m s ; but s o me of the tropic al specie s h ave erect ,

a eri al s te m s s o m eti m e s ri s i ng to a height of fift feet or m or e

, y
a nd for m i ng a tru nk whi c h is c li ndric al of equ a l di am ete r
y ,

thro ughout and b ea r s leave s o nl y at the su mmit li ke a p al m

, ,

Of the fer ns perh ap s the co mm on e s t and m os t widely

all
”
di st ributed is the ”
br ake or e agle fe rn w hi c h is k no w n to
-

botani st s as P teris aguilina L i nn ae us o r P teridiu m a g

, ,
u ilin u m ,

1 05
1 06 THE B I OL OG Y OF A PL A N T .

K uh n.
This pla n t is n o t o nl y co m m o n but of co m pa rativel
y ,

i ple t r ucture it i of co ve ie t i ze an d h as bee n m u c h

s m s a n n n s
s ; ,

s tudied . I t m a
y therefore be t a ke n both as a repre s e n tativ e

fer n and as a repre se ntative of all highe r vegetal o r ga nisms .

Habitat N ame etc The b r ake o cc u r s w idely di stribu ted in

, , .

the U nited S tate s u nder a g r eat va r ie ty of co n ditio ns ; e g in

,
. .
,

loose pi ne grove s e s pe c iall y in sand y r egio ns ; in ope n wood

,

l ands amo ngst the o t he r underg ro wth ; ou hill side pastu r es and
in thi c ket s — i ndeed almost eve ry whe r e except in ve ry wet or ,

ve ry dry pl aces I t appe ar s to be e q ually co mm on els e where ;

.

for accordi ng to S ir W J H oo ker P teris a g

,
.u ilin a gr o w s
.
,
- .

“
all round the w orld both withi n the t r opi c s and in the no rth
,

and south te mperate z o ne s I n L apl an d it ju st passe s

.

w ithi n the A rctic c i rcle asce ndi ng in S c otla nd to 2 000 feet , ,

in the C ameroo n M ou nt ai ns to 7 000 feet in A b y ss inia to 8 000 ,

”
or 9 000 feet in the Hi mal ayas to about 8 000 feet
,
S
( y pn o sis .

Fi li cu m ) .

Pteri s ( fl ré p zs the comm o n G reek nam e fo r f ern ) signify

, ,

ingw i ng or fe ather well accord s with the appea ran ce of P teris

,

g
a uilin a the m o s t co m mo n a nd m o s t ge ne rall y di st ributed of
,

E uropea n fe rns I t is poss ible th at th is fer n m ay r ank as th e

.

mo st u n ive rsall y di st ributed o f all veget able prod uc t io ns exte n d ,

ingits do min io n fro m w e st t o e as t over co n t i ne nt s and i sla n ds in

a z one r e achi ng from N o rther n E ur ope a nd S iber ia to N ew
Zeal and w here it is r ep re se nted by and pe r h ap s ide ntica l with
, , ,

the well k nown P teris es c ulen ta The r hi z o m e of o ur pl ant

-

like th at of the latte r is edible and tho ugh n o t e mpl oy ed in'

G re at B rita i n as food po w dered and m ixed w ith a sm all q ua n

,

tity of barley me al it is made i nto a ki nd of gruel ca lled g

-

oj io ,

in use amo ng the poore r i nh abita nts of the C ana r I sl and s

y .

( owerb y )
S .

The spe c ific name a g g

u ilin a ( a u ila e agle
) a n d a pop ul a r ,
“
na m e , e agle fer n ”
in G er many
-

et c h ave c o me f ro m a
, ,
.
,

fanciful like ne ss of the d ark ti ss ue see n in a tra ns ve rse section

of the le af stalk to the figure of an out spread eagle The sam e .

fig u re h as however bee n co m pa red t o an o a k t ree and has also

, , ,

g i ve n ri se to the nam e of dev i l s foo t fer n fro m its alleged ’

-

s m
r e e bl ce to the impre ssio n of the deil s foot etc etc
a n ” ’
,
.
,
.

“
The popul r de ig tio of thi s pl ant as the b rake te s ti
a s na n
 THE P L A N T B OD Y . 1 07

fies to its gr eat ab un d ance ; for a b r ake is a dense thicket o r

—
u nde r gro w th as for ex am ple a cane br ake ”
.

Whe n fully gro w n ( Fig 4 4 ) the co mm o n br ake h as a le afy

.

top supp orted by a poli shed d a rk colored erect stem which in ,

-

, ,

N ew E ngla n d ri se s to a height of fro m o ne to fou r feet abo v e

the gr o und I n t h is c li mate ho weve r it appe ar s to be so m e
.
, ,

what under sized for it grows to a height of fou rtee n feet in

,

the A nde s and in A ustr ali a attai ns to t wice the height of a

*
,

man for mi ng a de nse u n derg rowth be neath tree fer ns 4 0—1 00

,
-

feet h ig hq L
I n G re a t B rit ain it is f r o m six i n che s to n i ne feet
g
hi h ( S o werby ) o r eve n large r in exceptio nal c ase s
,
“
I n dry .

gra v el it is usually pre se nt b ut of small si z e ; w hile in thick ,

shad w ood s havi ng a m oi s t and r i c h so il it a tt ai ns a n e n or m o us

y
si z e
,
and m ay ofte n be s ee n c li m bi ng u p as it were am o ng the , ,

lo wer b ra nche s a nd unde r w ood re sti ng its de lic ate pi nnule s ,

”
on the li ttle t w igs and h angi ng g racefully over the m
, .

( N e w m an
)
G E N E R A L MO R P HO L OG Y O F T HE B O DY .

The body of the fer n like th at of the ea rth wor m consi sts , ,

of cell s grou ped to for m ti ssue s and orga ns Thei r arr ange
, .

m e n t ho w eve r diff er s w idel y fro m th at in the an i mal for t he

, , ,

pla n t bod y is a n e a rly so lid m as s and there are n o exte nded

-

i nter nal cavitie s e nclo s i ng i nter nal orga ns The organs of the .

pl ant are for the m o st par t exte rn al and ari se by loc al mo difi ca ,

tion s of the ge ne ra l m as s L ike ma ny higher pl ant s the body

.

of the fe r n co nsi st s of an a x is or ste m be ari ng br an che s fro m -

w hich a rise le a ve s The fer n diff ers for m ordi nary tree s h o w
.
,

ever ln the fac t th at the stem w ith its br an che s lie s hori z o n tal
, , ,

be neath the su rface o f the gro und O nl y the le ave s ( fro nd s) .

rise i nto the air ( Fig . I t is co n ve nie n t to de scribe th e

.

body of the br ake accordi ngl y as co nsi sti ng of two ver y dif
f ere nt pa rt s —o ne gree n and leafl ik e w hi c h ri se s abo v e the
, ,

grou n d ; the other bl ack an d rootlike l y i ng b uried in the soil , .

The se w ill he n cefo rth be spoke n of as the a erial and the un der
g d p art s
ro u n .

The u nderg ro u nd p a r t lie s at a depth of an i n ch to a foot

Hoo k er, l . 0
.

f K ro ne , B ota n . Ja h r esbericht 1 8 7 6 , 346 .

FI G .
—Th
r k ( P t i q utli n ) h w i ng p rt f t h und rgr und t m ( h)
e B a e er s a a , s o a o e e o s e r.
an d tw l v n (B )
o eaf th e s,pr o t y r in f ull d v l p m n t ; th th r
e , o e es en ea , e e o e e o e

( P ) f th p t y
, o r d d nd w i th r d d p i l b ud t t h x t m i t y f
e as ea , ea a e e . a ), a ca a e e re o a
br n h whi h b
a c th t u m p f l v f p r d i g y r nd n um r u
c ear s e s s o ea es o ec e n ea s a e o s

r t ; 0 m tu
oo s t iv l f 1 d d l f f p r d i gy r ; l m l m i n f l f
, a re ac e ea 7
, ea ea o ec e n ea . , a a o ea

p pi , ; h p rt i
n na f m i n rh i z m ;
ra , o on y u gr pi
o whi h i
a h wn u o e r, o n e nn a , c s s o e
l gd t Bar e Th i p i n i
a . r l y i m i l r t t h p i n ul f l d r p i n ( x
s n a s n ea s a o e n es o o e n a .
 A E RI AL AND UN DE R GR O UN D P A R TS . 09

below the su rface and b r anche s widel y in va riou s di r ectio ns

, .

I t may ofte n be followed for a lo ng dist ance an d in such case s ,

r eveal s a su rpri si ngly co mplicated sy ste m of u ndergr ou n d

b ranche s A t fi rs t sight the u n derg r ou nd po rtio n of the fer n
.
,

a ppea r s to be the root but a close r ex am i natio n shows it to be

,

r eall y the s tem or axis of the plant which differ s fro m ordi nary ,

ste ms chiefly in the fact that it lie s ho ri z o nt ally u n der the

grou nd in stead of ri si ng ve rtically above it The ae ri al portio n .
,

w hi ch is ofte n take n fo r s te m and leaf is reall y le af o n l y The , .

t r ue r oot s are the fi ne fib re s which s p r i ng in g r e at ab un da n ce

from the u nderg r ou nd ste m U ndergro un d ste ms m ore or le ss .

like that of P teri s a r e not u nco mm o n—o ccu r ri ng for i nst an ce , ,

in the potato the S olo mo n s se al the o nio n etc

,
’
I n P teris
-

, ,
.
,

a n d in ce rta in othe r case s the u n dergrou n d ste m is tech n icall

, y
called the roots to ck o r rhiz ome and in thi s pl ant it co nstitute s ,

the l arger and m ore pers i ste n t pa r t of the organi sm I n the .

s peci m e n show n in Fig 4 5 the r hi z o me w as about eight feet

.

long and bo re tw o lea ve s I t w as dugo ut of sa ndy s oil o n the

.

edge of a woodla nd and lay fro m o ne to six i nche s below the

,

s u rface. I t w as c r o ss ed and r ecro ssed in all directio ns both ,

above and below by the rhi z o me s of its neighbors the w hole

, ,

co nstituti ng a co a r se net wo rk of u ndergrou nd ste ms loo sel y fi ll

ingthe uppe r lay er of the s oil .

The aerial p ortion ( the f ron d o r leaf ) is like w i se divi sible

i nto a n umber of p ar t s c o m p ri si ng in the fi r st pl ace the le af
,

s talk or s ti e a nd the le af p r ope r or la m in a The latter is s ub di

p ,
.

v ided like a fe a ther i n a tely i nto a n umber of lobe s ( p i n n a

(p n ) ,

Fig . whi c h v a ry in fo r m accordi ng to the state of de

v elo ment o f the le a f I n l a rge le a ve s the t w o lo w er pi nn ae are
p .

ofte n l a rger th an the other s so th at the leaf appe a r s to consi st

,

of th r ee pri ncip al divi sio ns and is sa id to be terna te or t rip

,

ly divided ( Fig 4 4 A ) . E ach pi n na is in t u r n pi nnatel y sub

,
.

di vided i nto p in n u les ( p in n u les) or le aflet s ( Fig 4 4 B ) e ach of .

, ,

w hich is t r a versed do wn the m iddle by a thicke n ed ridge o r

r od the m idri b The le aflet s so meti me s h ave sm ooth o utli ne s
, .
,

b ut are u su all y lobed alo ng the edge s as in Fig 4 4 B In ,

.
, .

thi s case their for m is said to be p in n a tifid E ach lobe is like .

fu r n i shed with a m idr ib The s tip e e nl ar ge s som e wh at .

j ust belo w the s urface of the grou n d then gro w s smaller and ,
11 0 THE B I OL OG Y OF A P LA N T .

joi ns the r hiz o me The e nlarge .

me n t is of co ns ide rable intere st ,

fo r it occu rs at preci sel y the

poi nt of greate st str ai n whe n the
leaf is be nt by the wi nd or othe r
wi se and m u s t serve to s trength,

en the s tipc .

I t w ill appear fr o m the fol

low i ng de sc riptio n th at the pl ant
bod y exhibits in so me m easu re
certa i n ge ne ral fo r ms of sy m
m etry and diffe r enti a tio n whic h
in a b r o ad se nse may be r ega r ded
a a nal o g o us to tho se occ u r ri ng in
s

the ani mal The rhiz o me gr ows .

onl y at o ne end and in its st r ue ,

tu re sugge s t s the an te ro poste rio r -

differe nti ati o n of the a ni mal I t .

al so sho w s a slig ht diffe re ntiatio n

betwee n the up pe r and lowe r
sur fac es which ap pea rs bo th in ,

the ext er nal fo rm and in the a r

ran ge me nt o f the i nte rnal li nes .

I t is fu r ther m ore dis ti nctl y bil a t

e ral a ve rtica l p l ane dividi ng it
,

i nto clo sely s i mil a r h al ve s These .

featu res are howe v er f ar le ss , ,

p r o mi ne nt in the fe rn than in
the earthwo rm and in plan ts ,

they n eve r atta i n a h igh degr ee

of devel o p me nt w hile in the ,

highe r ani mal s they a r e a mo ng

the mo s t co ns pic uo us and im
po r t ant feat ur es o f the body .

F m 45 _ A n 0mm O f m ore ge n er al im po rta nce in

, ,
,
pl an t f Pt fi o O c s ne
l v i the f ern I S the m petl tlo n Of
.

O f th e ea es s
y u g
o n d m ll an s a
f s i m i l ar pa rt s ( b ranc he s r oot
,

an d mp i
a co a r so n o s
th fi g ur wi th Fig
, ,
e e
“ will h w m f leave s alo ng th e w li iC h
.

s o so e o a xi s
th d i ff r
e b e e n c es e
t w l v f d if sugg
ee n ea e
es o sts pe r haps a ce rta i n an
f erent a ges .
, ,
 A X I S A N D A PP E N D A G E S . I ll

alogy to animal m et a me ri sm though no t usuall y recog ni z ed

,

o r de signated by the sa me te rm A ll of the se c o nditio ns of .

differe nti atio n and symmet ry are m ore e as il y made out by an

exami natio n of the ae rial p o r tio n .

The pl an t as a whole m ay be r egarded as co nsi sti ng of

,

an a xis ( the r hi z o m e a nd its b ra n che s) w hich be a r s a nu m be r

of app end a g es in the for m of root s an d le a ve s The axi s forms .

the cent ral b od y or tru nk of the pla nt and in it mo st of its mat ,

te r and en ergy a r e s tored ; the appe nd es are orga ns for t aki ng . 0r

in food fo r ex c retio n fo r r e spi ratio n for reprod uctio n etc

, , , , .

Th e Under g r ound St em or R hiz ome a nd its B r anch es The

, , .

rhizo me is a ha r d bl ack elo ngated and b ra nchi ng ste m ge ner

, , ,

ally flatte ned so me what iii the ve r tic al di r ecti o n as it lie s in the
e arth and expa nded slightl y o n either s ide to fo r m w ell marked -

late r al fold s —the la tera l ri dg

,

es I ts thick ne ss is seldo m m or e
.

tha n h alf an i nch and u su ally co ns iderabl y le ss I n tr ansver s e

, .

s ecti o n it h as the outli ne s how n in Fig 4 8 and the ma rgi n al .

,

pa rt o nly is black The br anche s r e pe at in all r e spect s the for m

.

a nd structu re of the m ai n a xi s B o th the ma i n axi s an d th e

.

b r a nches end eithe r in conical poi nted and fle shy str uctu r e s
, ,
'

a bout t wo i nche s lo ng or in blu nt y ello wi sh k nobs pl ai n ly de

, , ,

p r essed in the c e nt r e A t the s e e nd s the r hi z o me gro ws ; he nc e

.

they are call ed the gro wi ng poi nt s or ap ica l buds ( Figs 4 4 .

,

B eside s the a pica l b ud s the rhi z o m e be a rs ne a rl y a lw ay s on e

o r m o re dead decay i ng tip s The se a ri se in the follo wi ng m an
,
.

ner : A fte r atta i ni ng a certa i n le ngth both the rhi z o m e and its

b r anche s grad u all y die aw ay behi nd Death of the hi nder p ar t .

follo ws at abou t the sa me r ate w ith w hi c h gro w th advan ce s at

the apica l bud s ; so that the total le ngth m ay no t ch ange m ate
riall fro m ea r to e ar I t is obvi o us th at thi s proce ss mu st
y y y .

r e sult in the gradual and succe ssive det ach me nt of the br an che s
f ro m the mai n axis E ach br anch no w beco m e an in depend
.
,

en t rhi z o me r epe at s the process ; a nd in thi s m a nn er a s i n gl e

,

o r igi nal rhi z o me m ay give ri se to l a rge n umber s of di sti nc t

pl an ts all of which ha v e bee n at so m e ti me in mate ri al co nn ec
,

tio n w ith an ance st ral stock Thi s proce ss is evide ntl y a ki nd o f

.

re rod uc tio n tho ugh it is n o t the m o st i m porta n t or m o s t obv i

p (
o us m e ans fo r the propagatio n of the pl a nt and in thi s w a
) y a ,
.

l a rge a rea may be occ upied by d isti nct though r elated plant s , ,
1 12 THE B I OL O G Y OF A PL A N T .

who se b r anchi ng rhi zo me s c r o ss and rec r oss m aki ng the subter ,

r an ean n et w ork al r eady de s c ribed p 1 09 , . .

Or i gi n of L eav es upon th e R h izome and its B r anch es The .

o ung pl an t of P te ris put s up a nu m ber of leaves ( 7 1 2 ) y ea r l y

-

y ,

b ut the ad ult ge ner all y develop s o ne o nl y wh ich grows ve r y ,

sl owly requiri ng two y e ar s befo r e it u nfold s Towards th e end

,
.

o f the fir s t y ear it is recog ni z able o nl y as a m i n ute kn ob at the

bottom of a depre ssio n near the growi ng poi nt A t the be gin .

ni ng of the s eco nd y ear it is perhaps an i n ch high the stal k ,

ft r S h )—D v l p i ngl f t f Pt i A d f br n h h w
(A e ac s . e e o ea , e c .
, o er s . . en o a a c s o
i g th
n pi
e a l b ud nd th rud i m n t f l f ; B r ud i m t ry l f ; G
ca a e e o a ea , a en a ea , a
si m i l r l f i l gi t ud i n l t i
a ea n on h w i n gth i f l d d l m i ( I ) th tt h
a se c o n, s o e n o e a na . e a ac
m t t t h rh i z m
en o e nd th pr l
o g ti e, af th t i u f t h l t t r i t th
e o on a on o e ss es o e a e n o e

l f ; D l m i f v ry y u gl f ; E h r i z t l t i thr u gh g Wi S
ea , a na o a e o n ea , o on a se c on o a rO D
p i t wh i h h
o n cj u t f rk d t f rm t w p i l b ud
as s o b e pi l b ud ; p pi
o o o a ca s . a . a ca o . o a

d mier s d u d ly i g
an l r t i p r h y m ; L b fi b v ul r bun d l ; l
n er n sc e o c a en c a , ro -
a sc a es .

l mi
a na
; r t;r, l r t i pr
oo hy m ; d v n t i t i u b u d t th
sc e o c b se
o senc a r, a n a
z e o s a e a
of th l e f ea .

on l y h avi ng appea red A t the end of the seco nd y ea r the lami na .

is devel o ped and h angs do wn as sho wn in Fig 4 6 0 E a rl in

,
y .
,
.

t h e spri ng o f the third e a r it bre aks through the gr ou nd an d

y ,

gro ws r apidl y to the f ull y matu r ed state -

.
 L E A VE S A N D R HI ZOME . 1 13

The leaves usuall y ar ise n ea r the apical bud s of the main

axis o r of the b r anche s B ehi n d each m atu re leaf re mnant s of .

the leaves of p recedi ng y e ar s are ofte n to be fo un d alter nati ng ,

o n the side s of the r hi z o m e in r egul ar succe ssio n and showi n

g ,

variou s s tage s of decay The fi r st of the se (which is o n th e .

opposite s ide of the r hi z o m e fro m the livi ng leaf) was alive th e

p r eviou s y e a r ; the n ext (o n the sa me s ide w ith the livi ng le af) is

the le af of the y ear befo r e th a t ; and so 011 Fig 4 7 sho ws an . .

“
example of this so r t The leaf of the present y ear l is fully .
, ,

FI G . f t r S h )—B
47 . (A e ac s . ra no h o f a rh i m
zo e of P t eris , s h ow n i gth e ap cai l b ud ( b ) a ,

th t um p f um b r
e s s o a n e o f s ucc es s iv l v
e ea es l “ l’
( , , ,
’
l a nd a pa r t f th m i n
o e a

rh z m ( h i r t
i o e r . o o .

devel oped ; and the relics of the leave s of the precedi ng y ea r s

l is the r udi me nt of n ext y e a r s le af
’ ’
are i ndicated a t l l etc
‘ '
.
, ,
.

I nt er na l S t r u ct ure of th e R hi zome The rhi z o me is a n e a rl y .

solid mas s co nsi sti ng of m a ny differe nt ki nd s of cell s u nited

, ,

into diffe r e nt ti ss ues and h a vi ng a ver y co m plic ated arr ange,

me nt I ts st ud y is s o m e w h at difficult
. N everthele ss the ar .

range me nt of the cell s is defi nite and c o nst ant and m e rit s c a reful , .

atte ntio n si nce it h as many featu re s which are cha racteri stic of
,

the cellular st r uctu r e of the ste ms of highe r pl ant s We sh all .

fi r st exami n e its m ore obvious anatomy as di spl ay ed in t rans ve r se

and lo ngitudi n al sectio ns afte rwa r d s maki ng a c ar eful m icro ,

sco ical stud of the cell d ti ue

P y s a n ss s .

S ee n w ith a ha n d le ns or the n aked ey e a t ra ns v e rs e section

-

of the r hi z o me ( Fig 4 8) p r ese nts a white or y ellowis h back .

1 14 THE B I OL O G Y OF A PL A N T .

bou ded b bl ck m rgi n ( the id r m is ) a nd m a rked

g
e e
rou nd n y a a a p
vario colored or p ale s pot s a n d ba n d s ; the l atte r a r e diff er
by us

e nt ti ss ue s or s y ste,
m s of ti ss u e The s e diffe r e n t s t r
.uctu r es a re
a rr a nged i n th r ee g r oup s o r sy s tem s of ti ss ue which a r e fo und ,

F ro 48
. .
—
C ross -
sec ti on of th e rh i z o t
m e o f P eris L r, la e a
. t r l ri dg ; f p fun d m nt l
es . ,
a e a

pa r e nc hym a ; s .
p, l r ti
sc e o c pa r h
e nc y m a ; a p o r
sc e o , l r t i pr
c hy m : L b x
o se n c a .
.

fi b ro r bundl
-
v a sc ula es .

a m o ng all highe r pl ant s in e ss e nti all y the sam e fo rm though ,

d ifferi ng w idel y in the m i no r detail s of thei r arra nge me nt .

T he se are

The Fu nd ame nt al Sy ste m of Ti ssue s

I . .

II The E pider mal Sy ste m

. .

III The Fibro vascular S yste m

.
-
.

The Fu nda menta l sys tem co ns ist s in P teris of th ree ti ssue s

n da men ta l m
( )f
a u
p a ren chy a ( Fig 4 8 f p) the s oft w hiti sh .
, ,

m ass for m i ng the pri ncip al sub s ta nce of the rhi z o m e ;

(b) sclerotic p a ren cltym a the brown h a rd ti ssue l y i ng
st below the epide r m i s fro m w hi c h it is sca r cel
j u
y di s ti
,
ngui s h
a ble ;

( ) sclerotic p rosen oby ma (8 p ro) black o r reddi sh dot s and

0 .
,

b and s of ext r e m el y h a rd ti ssue m o st of which is co nt ai n ed in two ,

c o nspi c uous ban ds l i ng o ne on eithe r s ide of a pla ne joi ni n g

y
t he later al r idge s .
 THE GR E A T TI S S (TE S Y S TE MS -

. 1 15

The sclerotic pare nch y m a and the sclerotic p r o sench y ma both

a r i se th rough a t ra nsfor matio n
( h arde n i ng etc ) of portio ns of , .

o rigina ll soft f un d am e n tal p a re nch m a I n mo st pl ant s above

y y
-

t h e fer ns the f und ame ntal sys te m con t a i ns n either of the se ti ss ue s .

The Fibre va scu la r sys tem is com po sed of lo ngitudi nal

-

t h read s o r str and s of ti s sue k now n as the ibro ra sc ula r bun dles
j -

a n d these in o ne fo rm or a n other are cha r acteri stic of all higher

pl ant s They appe ar here and there in the sectio n ( Fig 4 8 f b)

.
.
, .

as i ndi sti nct p ale or silvery a reas of a roundis h ov al or elon

, , ,

ga ted sh ape C lo se ly exa m i ned they show an O pe n text ure eu

.

clos i ng s pace s whi ch are sectio ns of e mpty tube s o r ve ss el s and ,

fi b r e s fro m w hich the bu ndle s t ake their name

, .

The E p iderma l s ys tem co nsi st s of a s i ngle tissue the ep ider ,

m is which cove r s the outside of the rhiz o me

, .

B y a shn ple dissectio n o f the ste m with a k nife the scle rotic
p ro se nch y ma and the fi bro vasc ula r bu ndle s m ay be see n to be -

lo ng str ands or ba nd s cou r si ng th r o ugh the softe r f unda m e nt al

,

ti ssue s .

I t should be clearl y u nde rstood that these th ree sy s te ms are ,

in ge ne r al no t si ngle ti ss ue s b ut g
, rou s of ti ssu e s which are
p ,

c o nst an tl
y associated togethe r fo r the pe rfor mance of ce rtai n
fu nctions *
.

M I CR O S C O P I C A N A TO M Y ( H I S TO L OG Y ) O F TH E R H I ZO M E .

Gener a l A ccou nt M ic ro sCO pic

t d y of thi n sectio ns of the
. s u

r hiz o me sho ws the vario us ti ssu e s to be co mpo sed of i nn um er able

c losel cro w ded cell s w hich d i ffer ver w idel in s tr u ct u re a n d
y y y
-

in functio n I n study ing the se cell s the stude n t should n o t lo se

.

g
s i ht of the fact that the
y are objects havi n g three di m e n s io n s ,

o f w hich o nl t w o are s ee n in s ectio ns A n d he n ce a s i ngle sec

y .

tio n m ay give an i m pe rfect or e ntirely fal se i mpre ssion of the

r eal for m of the cel ls — ju st as the face of a w all of m aso nr y may
,

g ive o n l y a n i m perfect ide a of the block s of which it i s b uilt .

h i l i fi t i n f th t i u i nl y m tt r f nv n i n
T s c ass ca o o nd h s e ss es s o a a e o co e e ce, a a

li ttl
e sc ei nt i fi v l u B y m ny b t ni t i t h b n r j ted lt gth r b ut
c a e . a o a s s as ee e ec a o e e

no a po l g y f
o its u n d be m ad b y th
or wh l k th uth rs h v
se ee e ose o, i e e a o , a e

f und i t u f ul l ng i t i d f nd d by S h ( wh fi r t i nt r du d i t ) nd
o se ,
so o as s e e e ac s o s o ce a

i ts v l u f
a e begi nn rs i n ded by D B ry
or e s co ce e a .
 THE B I OL O G Y OF A PL A N T .

\ /
F or thi s r e ason many of the cells can o nl y be u n de rstood by a
comp ar i son of tr ansver se and longit ud inal s ect io ns and the se ,

sho uld be s t udied togethe r u n til thei r r el a tio n

gy
s are tho r ou hl

mastered .

The follo wi ng t able gives brief defi nitio ns of the leadi ng

veget al ti ssue s and is good no t o nl y fo r P teris but fo r all
plants
PR I N C I PA L AD UL T VE G E TA L T I S S UE S .

These six ti ssue s are not o nl y fou nd in the rhi z o m e but ex ,

tend through o ut the r oo t s and the fr o nd s as w ell M o reover .

,

all the ti ssue s no t o nl

y of the fer n but of all h igher pla n ts are
v arietie s of the m .

Speci al A ccount I t must no t be forgotte n th at the di ffe re nc es

.

bet wee n ti ssue s are o nl y the o utco me of the differe nce s bet wee n
their com po ne nt cell s (p S o th a t the st udy of the h is tology
.

of the rhi z o me eve n if preceded (as it may w ell be) by a dissec

,

tion and a naked ey e ex ami natio n of so m e of the ti ss ue s eve nt

,
-

n all
y re s olve s it se lf i nto the careful m icro scopic s tudy of th e
several ki nd s of cell s co mpo si ng those ti ss ue s .

The mat ure p art s of the rhi z om e c o n tai n at leas t ni ne ve ry

differe nt ki nds of cell s the ch arac te r i sti c s and gr oupi ng of
,

which are sho wn in the follow i ng t able I n the apical bud s .

,

ho wever thi s ar rangement disappea rs and all the cells appea r

, ,

clo sel y si milar

.
 HI S TOL O G Y OF THE R HI ZOME . 117

M I N UTE A N A TO M Y O F T HE R H I ZO M E O F P TE R I S A QUI L I N A .

B esides the above m en tio n ed ti ss ue s the rhi z ome con t ai ns

-

c e rtai n othe r seco nda ry v ar

i etie s w h ich w ill be de scribed fu rthe r

E pi der mal Sy st em E pi der mi s

. I t is the fun ctio n
. of th e
epider m i s (aided in thi s c ase by the u nderl y i ng sclerotic p aren
ch ma to p r otect the i nner ti ssue s fro m co n tact w ith the soil
y )
and to gu a rd aga inst de sicca tio n of the rhi z o m e d u ri ng drought s .

Th e cell s ( Fig 4 9 ) a r e de a d an d e m pty with e nor m ou sl y thick

.
, ,

ha r d w alls perfo rated by num e r ous b ra nchi ng c anal s The outer .

wall is e speci all y thick .

Fu ndamenta l Sy st em The ti ss ues of thi s sy ste m fo rm th e

.

mai n body of the pla nt and in the fe r n have two widel y diffe r
,
 THE B I OL OG Y OF A PL A N T .

FI G . 49 .
—
S ec t i on s h o win g th e ep i d rm i ( p ) nd th und r l y i ng ler t i p
e s e a e e sc o c arcp .
ch y ma of th e rhi zome o f P t i q uili n
er s a C n l m tim a .br n h i ng
a a s, so e es a c , a re
e v ry w h r
e e e s een . Th ese se rv d t k p th n l ivi ng lls
e o ee e o ce -
ce
n ec tion .

Fm 50 —C ro s s s ec t i o n l r t i pr n h y ma f th h i
o f sc e o
me o f P teris (1u
'
. -
c
.
o se c o e r zo
Th e eno m
r u ly th i k n d w ll n i t f th r l y r
o s c e e a s co rf r t
s s o ee a e s, are pe o a ed b y 03 03 18 .
a nd are ligmfi d t u rn d i n t w d
'

e or e o oo .
 HI S TOL OG Y OF THE R HI ZOME . 1 19

e nt fu nctio ns The f un da men ta l p a ren cby ma is a ki nd of sto r e

.

house in which m atter and e ne rgy are stored—mai nl y in the

fo rm of starch — and in which ac tive che m ical ch ange s
,

take place The ce ll s a r e thi n w alled and soft and are rathe r
.
-

loo sel y joi ned together leavi ng nu m ero us i n te r cell ul a r space s ,

Fig The co ntai n protoplas m an d a nu c leu s

( s 5 2 .
y , an d ,

very n u m erous rou n ded grai ns of sta rch Thi s sta rch is sto r ed .

up by the plant du r i ng the summ e r as a re serve suppl y of food

—just as hibe r nati ng ani mal s store u p f at in their bodies for use
duri ng the wi nter A cco r di ngl y sta rch i ncre as e s in qu antit
.

y ,

dur i ng the s u mm e r an d dec r e ase s in the sp r i ng whe n the plan t

r esu me s its g r owth before the le ave s a re unfolded The pare n, .

c hyma probabl y has also the f unctio n of c o nd ucti ng v a rio us s ub

s tance s ( e s peciall y di ss ol ved sug a r th rough the pla nt b di fu io

) y f s n

f r o m cell to cell .

The sclero tic p a rench y ma a nd scleroti c p rosench y ma ( Fig

s 49 5 0) are .
,

d ead, h ence pl ay a pas si ve pa r t in th e

a nd a du lt veget al economy The .

for mer co o pe ra tes wi th th e epidermis ; t he

-

l a tt er pro ba b l y s er ves in par t to s uppo r t th e

s o f t t iss ues a nd to so me ex t en t a ff o rds a
,

c h a nnel f or th e co n v eya nc e o f th e sa p The .

s ap,
h owev er does no t fi o w th ro ugh th e
,

c a v i t i es b ut pass es s l o wl y a l o ngt he s ub
,

s ta nce o f th e po ro us wa ll s The c ell s o f .

bo th th ese scl ero t i c t is sues h av e v ery t h ic k ,

h a rd b ro wn wa ll s per fo ra t ed h ere a nd
, ,

t h ere b y na rr o w ca nal s Th e cell s o f t h e .

p a r ench yma are pri s ma t i c o r po l y h edra l ;

t h o se o f t h e p ro s enc h y ma el o ng a ted a nd ,

p o i n t e d a t th e ir end s I n b o th t h e p r o t o .
,

p l as m a nd nucl ei di sa ppea r wh en th e cell s

a re f ull y f o r med To war ds th e a pi ca l buds
.

bo th fade i nt o ordi nary f undament a l pa ren

chy ma .

Fi br o -
The fib ro
v asc ular Sy st em .

va s ea h a
( /
bun dle? ( p 1 1 5 ) are lo ng F 51 (A f t r S h —
) Vi w f ro e ac s e o

gg g
.

g
. . .

zi iififi fic zff
i
s tr and s o r b and s of ti ss ue which ap
‘ s

o s

pe ar in cro ss sectio n as i sol ated spot s th t w rk f t h up p r fi br

-
e ne o o e e e

v u l r b un d l h l f a sc a es a ea
( Fig The bu n dle s are not
. .

r eall i s ol a ted ho w ever but joi n o ne anothe r here and the r e

y , , ,

formi ng an open n etwork ( Fig which can onl y be seen in a .

1 20 THE B IOL OG Y OF A PL A N T .

late r al view of the rhi z o m e Fro m this n etwork bu n dles are .

given off whi c h extend o n the o ne hand i nto the r oo ts and on

the other i nto the leave s b ra nchi ng in the latte r to fo rm the ,

complicated syste m of vei ns to be desc r ibed he reafte r

Ea ch bu n dle co ns i s t s of a n u m be r of di fi erent t issue s which ,

pe ki g h ve the f un ctio n of co n ducti ng sap f r om

b roadl y s a n a ,

part of the p lant to anothe r .

$ 1 0 52
. .
—Hi hlifi d r
g y m a gn ti n f fi b e c oss sec- o o a re - v l r bundl u und d b y
a sc u a e s rro e
f t l r h y m f p t l rif
t h e un da m e n a p a en c a, . .
, sc a a o rm tr h i d ; b bund l h th ;
ac e s . s, e s -
ea

p hl em h th ; b f h t fi b ; t i v
o -
s ea .
, as -
re s a . s e e-
t ub ; u p p hl em p n h y m ;
es , o -
are c a
m p w d ( xy l m ) p r
. oo hym ; p ir l v
e a en c a a v, s a e sse l .

Th ese t i ssues h av e th e fo ll o wi ngdefi ni t e a rrangement B eg i nningwi th .

th e outs i de o f a b undl e we fi nd ( Fig s 5 2 5 3) , .

,

1 B und le s hea th
.
a si ng -
l e l ayer o f el o nga t ed cell s envel opi ng th e
bundl e pro ba b l y deri v ed f rom and bel o ng
, i ngto th e f undament al sys t em .

2 P h loem s h ea th ; a s i ng
.
-
l e l ayer of l arger pa rench ymat ous cells co n
tai ni ngst a rch in l a rg e q uant i t i es .

po i nt ed cells cont ai ning

3 B as t fi bres ; s o f t th i c k wall ed el on a ted
g
-
-
.
, , ,

pro t o pl asm and l arg e nucl ei .

4 S ieve tubes ; l a rg er so f t th i n walled el ong a ted cell s conta i ning

-
. -
, , ,

pro t opl as m a nd h a v i ngth e wall s ma r k ed b y a reas per forated b y numerous

fi

fi ne po res ( panell ed) Th ey j oi n at th e ends b y o b l i q ue panell ed par t i t ions

.

( sh own in Fig s 5 2 a nd .
 HIS TO L O G Y OF THE R HI ZOME . 21

5 . P h loem p a rench y ma ; o d na y pa enc y ma o us c e

-

h t r i r
ll s fill ed with r
s ta r c h tt r h r
, sca e ed t re e a nd he e among t i ev e t ubes
th e has fi b res a nd s -
-
.

6 Tracheids ( s ca la rif or m) or
. l adder cell s occupy ingmos t of the -

c ent ra l pa r t o f th e b undl e Th eir s t r uc t ure ca ll s f or so me r emar k

.
Th ey .

are empt y o r a ir fi lled f us i fo r m t ubes wh os e h a r d th ic k wa ll s a re in th e

-

, ,

y o u n g t i s s u e sc ul pt ured wi th grea t regul ari t y i nt o a s eries o f t ra ns verse

h o ll o ws o r pi t s wh i c h fi na llv beco me a c t ua l h o l es
, The wall s of th e .

t rach ei d a re th er efo r e c o nti nuo us at th e a ng l es b ut a l ongth eir plane sur ,

FI G . 53 .
—L o n gi t ud i n a l s ec t i o n of a fi b re -
v l r bun d l urr un d d b y th f un
a sc u a e, s o e e
d a m e n ta l p a en c y m a r
I) f , as hfi b re s . . h t -
; b bu d l h th ; f p fu d m t l
. s, n e s -
ea .
, n a en a

p a ren c h m a ; p 4 9 . p hl
o em p a en c y m a -
r h ; p hl em h th ; t i v t ub ; t
o -
s ea a . s e e -
es ,

sc a a o m l r if r tr h i
a c e d s o r la d d e c el s r l -
w p w d p r n hy m
.
, oo -
a e c a .

f aces beco me con v ert ed i nt o a s eri es o f parall el b a rs mak i nga gra t i ngof ,

s i ng ul ar b ea ut y Th e s l i t s be t ween t h e b a rs a re no t rec t a ng
. ul a r passag es

t h ro ug h th e wa ll but a re ra th er l i k e el o nga t ed fl at t ened f unnel s O peni ng

, , ,

o ut wa r ds Th e s ides o f th e f unnel s a re c all ed th e bo rd ers o f t h e p its ; and

.

pi ts o f th is s o r t are ca ll ed bo rd ered s ca la rif orm p i ts ( c f Fig . .

7 Tra ch ece or vess els ( sp i ra l) ; sca tt ered h ere a nd th ere a mo ngth e

.

t rach ei ds and h ardl y di s t i ng

, ui s h ab l e f r o m th em in c ro ss s ec t i on Th ey -

a re co nt i nuo us el o ng a t ed tub es fill ed wi t h a ir a nd s t reng th ened by a b eau ,

t i ful cl ose spira l ri dge ( so met i mes do ub l e) wh i c h runs round th e i nner f ace
o f t h e wa ll ( Fig .

The t rac h ei ds a nd v essel s are of g r eat ph ysi ol ogica l i mpo r tance being ,

p rob ab l y th e mai n c h annel s f or th e fl o w o f s a p S a p is wa t er h ol di ng .

v a ri o us s ub s tances in so l ut i o n Th e wa ter en ters by th e roots fl ows p ri n

.
,

c ip a lly th ro ugh th e wa lls o f th e vessels a n d tra ch eid s , a nd no t th rou g

h
th eir ca vi ties , wh ich a re fi lled with a i r, and is th us co nduc ted th rough th e
r h iz ome and upwar ds i nt o th e l ea v es .

8 Wood p a rench y ma ; cell s l i k e th os e o f

.
-
th e p hl oem parenchyma -

(5 )
s catt ered b et ween th e vessel s a nd t r ac h eids .
1 22 THE B IO L O G Y OF A PL A N T .

r a
B nch es of th e R h i z ome The s e r epeat in all r e s pects tf
st r ucture of the mai n stem They are equivalen t me mbe rs .

the u ndergrou n d p art an d diff e r in no w i se excepti ng in th , ,

or igin from the mai n ste m itself

,
.

B oots The root s may e asily be r ecogn iz ed by thei r sma

.

siz e and tapering fo r m and their lack of the l ate ral ridges of ti ,

FI G . 54 . r —S i v tub fr m th rh i z m f P t i q uiltn h
f r
( A t e De B a y ) . e e -
es o e o e o er s a a, s o
i g A th
n nd f
: m mb r f
, i v t ub B p rt f th i n l ngi tud i l
e e o a e e o a s e e -
e , a o a o na s

t i n Tho t i h pp r xi m t l y h l v d tw i v t ub S d S w h i h
. e sec on as a o a e a e o s e e -
e s. ‘ an ’ c i

so dr w th t t h u i j ur d i d l i
a n b h i d Th br d p t ri r ur f
a e n n f e s e es e n . e oa os e o s ac e o

i v r d wi th i v p l t
s seen c o ne et i ngw i th th r i v t ub S n t
s e e -
a e s co n ec a no e s e e
-
e .
’
, o

t ry bu t b y m th
c o n ra , pl t d
a u f
s up p r n hym t u
a s oo no n -
a e s r ace on a e c a o s cc

wh i h th r ugh i t 0 t i
c ar e se en f w ll b r i ng i v p i t ;
o ti n . 1 , s ec o ns o a s ea s e e
-
s r, sec
z o

a no n p l t d w ll butt i gup n p
-
a e h ym a a n o are n c a .

stem and b ra nches } They a ri se endog eno us ly fro m the ma

stem or its br anche s i e by an outg ro w th of the i nter nal ti ssu e , . .

,

(
and n ot as in the c as e of the false r oot s o r rhizo ids of the p r

thalli um shortl y to be de sc ribed ) by elo ngati o n of supe rfi c i

,

cell s of the epider mi s T r ue root s of which tho se of P teris a .

,

good exa mple s arise al ways as w ell fro m the fu ndame nt al a ]

,

fi bro vascul ar region s and i nclude all the s ste ms fou nd in t ]

y
-

ste m it self Hence cross sectio ns of P teris r oo ts di ff e r b

.
-

slightl
y from those of the ste m o r the b r a nche s and the r oot ,

general is clearl y a me mber of the plant bod y A s in all tr .

'

r oots the free end is covered by a s pecial bori ng tip called t

,
 S TR U T UR E C OF THE A PI CA L B UD S . 1 23

roo t cap but this is apt to be lo st in r e movi ng the speci me n

-

from the e arth .

Th e E mb ry oni c Ti ssue or Mer i st em of th e R hi zome The .

m atu re rhiz o me re mai ns at the tip n earl y u ndiffere nti ated i nto
ti ssue s A t thi s p o i nt the epider mis m a y be di sti ngui shed but
.
,

it re mai ns very delic ate and the un derl y i ng cell s conti nue to ,

gro w and multipl y prod uci ng co nti nued elo ngatio n of the m as s
, .

I n thi s way the apic al bud is fo r med L ater al bud s are give n .

o ff right an d left to co nstit ute the e m br y o s of le a v e s br an che s , ,

or r oots w hich al ways retai n i ng their soft and delic ate tip s are
, , ,

capable of further gro w t h .

”
B ehi n d the se g r o wi ng poi nts the epider mi s an d othe r
ti ssue s g r o w mor e and m ore slowl y and soo n r e ach their maxi ,

m um s i z e whereu po n r apid g r o w th ce ase s

,
The po w er o f .

gro wth is he nc efor wa rd m ai nl y co nfi ned to the apic al b ud s and ,

the g ro
w i ng ti ss ue of w hich the
y a re co m po s ed is k now n as em
'

bry on ic fi 8 8 tt€ or v zer es tezn

The A pi cal Cell of th e R hi zome C lo se exa m i natio n reveal s .

the fact that each apic al bud co ntai ns a re markable cell which is
e s peciall y c o nce rn ed in the fu nctio n of g r o w th v iz the ap ical , .
,

cell w hich lie s in a hollo w at the apex of the b ud

,
I n the .

apical buds of the r h i z o me o r b ranche s thi s cell has so me what the

0 0- .

Fro . t r H fm i t r )—
55 A . A pi
(Af e 1 ll o e s e .
~
ca ce FI G . f t r H fm i t r )
5 33 . (A e o e s e .

of th rh iz m in v r t i l l n gi t ud i l
e o e a e ca o na A pi l ll f th rh i z m i h r i
ca ce o e o e n o

sec ti n opi l ll ; i h i ; m m r i
. a ca ce t, a r , e t l l
zo n a gi t ud i l ti
on na s ec on . a o,.

api l ll
ca ce .

for m of a wedge w ith its base tu r ned for war d s an d its thi n edge
b ackward s the latter pl ac ed at right angle s to a pl an e pas si ng
,

through the l ate r al ridge s I t co nti nuall y i ncrease s in si z e bu t .

,

as it gro ws r epeatedl y divide s so as to cut off cells l ate r ally

1 24 THE B IOL OG Y OF A PL AN T .

lte r tel n its right a n d left s ide s These cell s in tu r n c on

na o
a y
.

t inn e to grow and divide and thu s give rise to two si milar masses ,

o f me r i te
s m which togethe
,
r co n s titute the a pical bud Fro m .

e te m b g r du l though r apid c h ange s th e va r ious tis

t h e m ri s y a a , ,

s ues of the adult rhi z o me are diff ere nti ated ; and lo ngi tudi nal
s
ectio ns p as s i n g th r o ugh the lateral r idge s s how the m atu r e
ti ssue s fadi ng out in a r egio n of i ndi ffere nt me riste m about the
apical cell ( Fig .

h
T e p a i ca l ce ll l i e s a t th e b o t to m o f a f un n e l s h aped depress io n at the
-

i f h t m I t i h a e d a p p r o xi m a t el y l i k e a th i n t wo edg ed wedg e
t p t e s s p
-

o e s . ,

wi th a n a rch ed or c ur v ed bas e t ur ned fo r wa rds t o wa rds t he cen t re o f t h e

f unnel sh aped depressi on The th in edge of t he wedge is direc t ed bac k
- .

r i h i h l r v m e t in a v ertica l
pl a ne a bove
'

w a ds a n d it s s d e s w c a re a s o
,
c u e d e ,
,

and b el ow A l ong .
i t udi nal sect io n t a ken th rough t he plane of the lateral

FI G 56
. . (A e f t r S h )—A v r t i l tr
ac s . e ca a ns vr e se sec t i n th r
o ou gh th e ap ical ecu, w
c .

s h o w in ga b u d r y f h i r nd
o n a o a s a a sec o nd a pi cal ll 1 ce , . i
be lo ngn gto a ea f l .

ri dges th erefore sh ows t he apica l cell in a t ri a ngul ar fo r m as in Fig 5 5 8 . .

A sect i on t a k en at ri g ht a ngl es to th is v ert i c al and l o ngi t udi nal

s h o ws t h e c ell to be a
ppro x i ma t el y rec t a ng ul a r a nd q ua dri l a te ra l ( Fig .

5 5 A ) wh i l e a t rans v erse v ert i ca l s ec t i o n s h o ws it in t he f o r m o f a b i co nv ex

,
-

l ens ( Fig .

The f unnel s h a ped depress i on is c o mpr essed v er t i call y a nd its wa ll s are

-

t hic kly cov ered wi t h erec t b ranc h i ng h airs wh ic h a re c l osel y fas t ened ,

FI G . 57 .
—
0r os s s ec
-
t i on
t i r f r t i l l fl t m m i d ib : v ins ; cp pi
of a n en e e e ea e . . r. r ve e
d m m p h y ll ; p rm ;
p r ng
e
i : i n i nd u i um
is s, es o 8 , s o a a , s .
, .

og
t eth er by a h a rdened muci l age s ec ret ed b y th e api cal bud Th ese h airs .

en
t i rel y cl ose th e mouth of th e f unnel and s h ut o ff th e del i ca te young
 HIS TO L O G Y OF THE L E A F . 1 25

base f ro m th e outer wor l d Protect ed by th ese h airs the

its .
,

of t h e s t em f o rces its wa y th r oug

. h th e t ough es t cl ay wi t h out i nj ury to
t h e del i cate bud b uried in its apex ( Ho f meis t er ) . .

FI G . 58 .
—C ross sec
-
ti o n. st i ll m r
l rg d p i ngth r ugh th m i d ib f l fl t
o e en a e , a ss o e r o a ea e .

I n th nt th
e cei ul fi b v ul r bun d l up p rt d p i ll y b v d
re e c rc ar re -
as c a e, s o e , es ec a a o e an

b l w b y th i k n d p n h y m
e o , c e e O n i th r i d t h p r n h y m t u m
ros e c a e e s e e a e c a o s, es

o p h y ll ll ( h d d ) n d t h in t r ll ul r p
ce s s a e a ( L ) p n i n gb y t m t
e e ce ( t) ; a s a c es s o e s o a a s

o r T HE B R A KE . THE F R ON D on LE AF .

The exte rn al fo rm of th e leaf has bee n de sc r ibed o n p 1 09 .

,

and it no w re main s to co ns ide r its i nte r na l s t ructu r e The .

lami na is to be r ega r ded as a fl atte ned and altered po r tion of the

st ipe made t hin and deli cate in orde r to p r ese nt a l arge su rface
,

to the light and the ai r The stipe in tu rn is a prolongation .

, ,

of the rhi z o me so that the whole plant body is a conti nu ou s

,

mass th r ougho ut which exte nd th e th ree sy ste ms of ti ssue vi r

,

tually u nch anged The t ransve rse and lo ngitudi nal sectio ns of
.

the stipe sho w o nly mi no r poi nt s of diffe r e n ce fro m corre spond

.

ingsectio ns of the r hiz o me I n the l ea f howeve r all th r ee .

, ,
1 26 THE B IOLO G Y OF A PL A N T .

systems u nde r go gr e at changes The epide r mis becom es ve ry

'

thin delicate and transpare nt ; the fi bro vascula r bu ndles b r eak

, ,
-

up i nto an extremely fi ne and co mplex n etwo r k fo rmi ng th e

FI G . 59 .
—
C ross -
sec ti on o f pa rt of a l fl t h wi ngth mi r p i tru tur
ea e s o e c o sc o c s c e .

i d rmi ; t t m t :
ep e s s . s o a a i nt r l lul r p
e ce a sb t w n th m p h y ll
ac es e ee e e so - l
ce l s.
wh i h fi ll d wi th ( h
c a re e s ad e d ) hl r p h y l l b d i
c o o -
l y i ngin th pr t p l m
o es e o o as .

v eins the sclerotic ti ssue s beco me transpa re nt and are fou nd

;
o nly along the vei ns The cell s of the fu ndame ntal pare nch y ma
.

alte r their for m lose their starch and beco me filled with bright
, ,

gr een rounded bodies c alled the chroma top ho res o r chlorop hyll
, ,

bodies which are com posed of a protopl as mic basi s colo r ed b a

,
y s

pigment k nown as chlor ophyll The g r ee n fu ndame ntal pare n .

c h ma of the lea f is so meti mes called the mes o h ll

y p g/ .

A cross section of a leaflet p 1 09 is show n in Fi

( ) g 5 7
-

. .

The fi ner structure of the le aflet is show n in Figs 5 8 and 5 9 . .

O n the out side is the epider mis e withi the m es ph ll

( p); n o y and
midrib —the latter compo sed of thicke ned epide r mal and sclerotic
,

fundamental ti ssue and a l arge fib r o vascula r b undle

,
-

The mesop hyll or lea f pare nch m a consists of i rr egu la r ce lls

y
-

, ,
 HI S TO L O G Y OF THE L E A F . 1 27

which a r e loosel y ar ra nged on th e lowe r side leavi ng ve ry la r ge ,

i nte r ce llular spaces but a r e closel y packed and leave few o r n o

, ,

inte r cellula r spaces o n the u ppe r ( su nn s ide The cell hav e

y) ,s .

ve ry thin wa ll s co ntain p r otoplasm and a la r ge ce nt r al spac e

,

FI G . 60 .
—
E p i d rm i f
e r id f l fl t h
s ro m th e un d e s e o a ea e , s o wi n gw a vy ce ll s l gt
; e o n a ed
( p m n lnfl n t u )
sc c ll v r t h v i n ; nd t m t
a o s ce s o e e e s a s o a a w i th th i gu rd ll t
e r a -
ce s . s ,

t m t
s o d g u d
a a anll ; v i n v r d b y th i k
ar -
ce s v, e s co e e c an d pr hy m t u p i
o se n c a o s e

d m l er ll I t rm d i t
a ce t g b t w n w vy
s. n e e a e s a es e ee a a nd tr i ght ll
s a l ce s a re a so

s h wn ( S urf
o i w )
. ac e v e .

( vac uole) fi lled with sap and nu me r ou s chlo r oph y ll bodi e s im ,

-

bedded in the protopl a m The se are e specially num e r ou s in s .

1 28 THE B IOLO G Y OF A PL A N T .

the uppe r par t of the leaf as might be expected f rom thei r ,

fu nctions in co nnectio n with the actio n of light ( see page

The q uidermis or sk in of the leaf co nsists of tra ns lu ce nt , , ,

reatl fl atte ned cells ha v i ng peculi ar w av y outli n e s and r el a

g y
tiv ely thick w all s ( Figs 5 8 Upo n .

the vei ns they be c ome elo ngated and ,

thei r wall s are co nside r ably thicke n ed ,

e speci ally u po n the mid rib ( Fig 5 8 .

,

They ge ner all y co n tai n large disti nct ,

n uclei and ofte n co ns ide rable p r oto ,

pl as m The wav y epide r mal cel ls .

,

pa rticularly in y oung pla nts co ntain ,

so m e chloroph y ll and s ta rch though ,

in thi s re s pect the fe r n is so mewhat

exceptio nal .

I n the r hi z o m e the epide r m i s fo r ms

a conti nuou s layer ove r the w h o le sur
F I G .61 ( A ft r S
.
h )—E p i fac e I n the leaf howeve r this is no t
e ac s . .
, ,
d rm l ll ! ” ml ” M M
he c as e t h e epide r mi s o n the lo w e r
s 0

I t
e

o a, s
a

h w i gth d v l p m t
o
ce

n
t e e e o en
“

fi t m t s o A v r y y u g s ide bei ng pe rfo rated by holes leading

e o n

g
o a a . ,

fi sxi iliti i é ii fiii

l f i n
nto the i nterior and k no wn as m outh s
‘
a G

c
i a
’

m th ro ll ;
e - ud id i
ce y o r s to ma ta (si ngul a r s to ma ) ( Fig
s s ar .
,
ce l l a gu d ll ; t t m
; o, ar -
ce s s o a
The se hole s do not p ass i nto the cell s
. , .

but are gap s or b r eak s betwee n c ert ai n cells of the epide r mis ,

a nd ope n directl y i nt o the i ntercellul a r s p ace s of w hich the y a re , ,

in fac t the e nd s ,
That porti o n of the i nte rcell ul a r l aby ri nth
.

whi ch direc tl y underlie s the st o ma is so meti me s called the res pi ra

tory ca v ity E ach s to ma is bou nded as in mo st pl ants by t wo
.
, ,

c urvi ng gua rd cells w hi c h are ge ner all y nucle ated and u nl ike
-

, , ,

epider mal cell s ge nerally co nt ai n ab unda nt chlor o phy ll bodi es ,

-

and st arch .

The guard cell s are capable of c h angi ng their fo r m acc o rd

-

ingto the a m ou nt of light the h y gro s copic sta te of the at mo s ,

ph ere and other circu mst ance s an d th us ope n o r clo se the h o le

, ,

o r sto ma bet w ee n the m Thi s actio n is of g r eat i mpo rtance in .

the ph ysiology of the plant (t ra nspi r atio n p , .

I n Pter is cretica and P fl a bella ta th e s t o ma ta devel o as fo ll o ws A

p .

y o ungepider m l cell is di v i ded by a c ur v ed pa rt i t i o n i nto t wo c ell s o ne o f

a
,

wh i c h ( Fig 61 ) is call ed th e i n itia l cell o f t h e st o ma

.
Th is is ag a in
 VE N A TI ON . 1 29

di vi ded by a r v ed pa rt i t i on i nto th e moth er cell of th e s toma (Fig 61

cu -

.
,

m c) a nd a s ubs i dia ry cell ( Fig 6 1

.
.
,

Th e mo t h er cell is th en b isec t ed i n to th e t wo g
-
ua rd cells and th e s t oma -

appea rs as a c h i nk be t wee n t h em ( Fig 6 1 B ) .

, .

The v ei ns a re the fibres o r th r ead s which co nstitute th e

f ra m ework of the leaf E ach co nsi st s esse nti ally of a small
.
, ,

fi bre vascula r bu ndle b ra nchi ng fro m th at of the m idrib ( Figs

-

5 7 58, ,
A bov e and below the m the m esoph y l l and epi
de r mal cell s ar e ge ne rall y thicke ned and p r ose nch ymatou s in thi s ,

way co nt r ib uti ng alike to the fo r m and the fu nctio n of th e

“ ”
vei n .

FI G . 62
. f r
( A te L ue rss e n ).
-
V e na ti on of a l ea fl et of P tcris aq i
uil na .

Th eir rra ngement ( v ei ni ngor vena tio n) is defi ni t e a nd depends on the

a ,

mode o f b ra nc h i ngo f th e fi b re v as c ul a r s t ra nd wh i c h co ns t i t ut es th e pri n

-

c ipal pa r t o f th e mi dri b S eco nda ry s t ra nds ( ne r v es ) p ro c eed f ro m th i s a t

.

a n ac ute a ng l e t h en t ur n so mewh a t a b ruptl y t o wards the edge o f th e

,

l eafl et ( o r l o be) mak i ng a n arc h wh ic h is co nv ex t o wa rds th e dist al ex

,

t remity o f th e midri b ( Fig .

Fro m th is po i n t a f ter b ra nc h i ngonce o r t wi c e the del i ca t e v ei ns run

, ,

par a ll el to eac h o th er to th e edg e o f th e l ea fl et wh ere th ey j o i n o ne a no th er ,

or a na sto mosc Th is f o r m o f vena t i o n is k no wn as N erva t io N europ teri

.

d is a nd is mo re eas i l y seen in th e l ea f o f Os munda reg

,
a lis ( cf L ucrss en .
,

R a benhorst s K ryp tog

a me nFlo ra III
’ -
.
,
s .
 C HA PTE R IX .

THE B I OL OG Y OF A PLA N T ( Continued ) .

R ep ro d u ct io n an d D e v elo p m en t of th e B rak e or F e rn .

R epr oducti on Unl ike the ea rthwo r m the fe r n rep r odu ces
.
,

both by g a mog enes is ( sexuall y) and a g g

a mo enesis
( asexuall y) .

P ierce posse sse s two mode s of asexu al rep r od ucti o n v iz the ,

.
,

detach ment of e nti re branches fro m the rhiz o me and the co n

s e que nt e stabli sh me nt o f i ndepe n de nt pl ant s as alread y men ,
”
tio ned (p . and the for matio n of a dve ntiti o us bud s from
the b ases of the leaf stalk s ( Fig -
B ut besi des thes e the
.

fer n h as a quite diffe r e nt method of r eproductio n in which a ,

process of agamoge ne si s regula rl y alte rna t e s with gamoge nesis

( a lterna tio n of g mwra tzo ns )
’

The follo wi ng brief outli n e of

.

this i mportant proce ss may help to g uide the stude nt th r o ug h

the subseque nt deta iled de sc riptio ns .

Upon so me of the lea ve s are fo rmed o rga ns called sp o ra ng ia

( Figs 5 7 6 3
.
, which p r odu c e nu me ro us rep r oductive cells
,

c alled s a res
p The spore s beco me detac hed fro m the pa re nt and
.

develop i nto i ndepe nde nt pl ant s the ro t/za llia Fig wh ch

p (
,
i .

d iffer e ntirel y in appe a ra nce fro m the fe rn an d ul tim atel p ro

y
d uce male and fe male ger m cell s The fe male cell of the p ro
-

thalli um if fe rt ili z ed by a male cell develops in to an o rdi nary

, ,
”
fer n which aga i n pr o duce s s pore s asex uall y The for ma
, .

tion and develop me nt of the s po re s is evide ntly a process of

g g
a a mo en es is a nd the fer n p r ope r is the r efore n eithe r male nor
,

female it is sexless or as exua l The for ma tio n and de .

v elo ment of the ge rm cell s o n the c o ntra r is a p r ocess of

p -

,y ,

ga mog en es is
; and the proth alli um is a di s ti nct se xual pla nt
,

bei ng b o th male and fe male (hermap hrodite or bis ewua l) I n .

ge eral ter ms thi s is expressed by calli ng the o r di na ry fe rn the

n

s pore be a rer
-

or sp orop hore and the p roth al li um the egg

, ,

bearer or oop hore

, The l ife h is to ry of the fe rn broadly
.
-

1 30
 A L TE R N A TI ON OF G ENER A TI ON S . 131

S peaki ng co nsist s therefo r e in an alter natio n of the sp orop hore

,

( asexu al g ne r atio n
) with the li / ( s exual ge ne r atio n that
c e
p t ore
);
is it co nsists of a n a ltern a tio n of
, .

g en era tio ns A n e ss e n tiall

y s i m. ila r
altern atio n of sporopho r e with o epho re
occu rs in all higher pla nts though in ,

most case s it is so di sguised as to es

cape ordi na ry obse r vatio n .

Th e Spor ang ia and Spores The .

sp ora ng ia of P teris ( Fig s 6 3 64 ) .

,

a rise upo n a lo ngitudi nal thicke ni ng

of ti ss ue sit uated o n the u nde r side of
the le aflet s nea r thei r edge s and in ,

c ludi ng a m a r gin al a nasto mo s i s of the

ve ins Th is s welli ng is kn ow n as
.

the recep ta cle Hai rs are not uncom F 61 ( A f t r S u m i n k i ) —S p0 I G . e s

}
.
.

n gu m f Pt i ul t ra i o er s serr a a
mon u po n the u nde r S i de of the leaf
.

p di l ; 0 p ul , n u l u ; , e ce , ca s e a, a n s

a nd so me a r e fou nd u po n o r n ear the ~S p r 8 o e

receptac le O n the l atter a ri se s t r uct ur e s at fi r st supe rfici all y

.
,

s im ila r to h ai r s which bec o me e nl a rged at the tip and fi nally

, ,

develop i nto the s po r angia Me anwhile the edge of the le aflet .

is be nt do wn a nd u nder so as to make a lo ngitu di nal b an d of

thi n ti ssue co mpo sed of epider m i s k no w n as the outer v eil o r
i ndusi u m ( Fig 64 a c) A s i milar thi n sheet of epider m i s
.
,
.

g row s do w n f ro m the u nde r side of the leaf and p assi ng out ,

ward s to mee t the fo r me r co nstit ute s the i nn er v eil or true ,

in e ln s i um ( Fig 6 4 B i i) .
, ,
. .

I n the V shaped sp ace thu s fo r med the spo r a ngia are de

-

eloped .

A rfi ci al ( epi der mal ) cell enl a rges and becomes divi ded i nt o a
s upe

prox i ma l ( bas al ) cell and a di s t a l ( api c a l ) c ell ( Fig 6 5 a ) Th e fo r mer de .

,
.

v elo ps i nt o th e f ut ur e p ed icel o r s t a l k o f t h e s o ra ngi u m t h e l a tt er g i v es

p
rise to th e h ead or cap s ule wi th i n wh i c h th e S po res are formed ( cf Fig . .

Th e pedi cel a ri ses f ro m t h e o rig i na l pedicel cell by co nt i nued gro wth and -

s ubdi v i s i o n unt i l it c o ns i s ts o f th r ee r o ws o f cell s s o mewh a t elo ng a t ed .

Th e ro unded caps ul e c ell is nex t t ra ns fo r med b y f o ur s uccess i v e o b l i q ue

-

di v is i o ns i nt o f o ur pl a ne co nv ex “ pa ri et a l cell s a nd a t et r ah edra l c ent ra l

-
”

c ell th e a rch esp o riu m enc l os ed b

y t h e o th ers The capsul e cell is th us -

, ,
.

d i v ided b y th r ee pl a nes i nc l i ned a t a b o ut 1 2 0 ( Fig 6 5 b c) A f o urt h

°
.
.
, ,

( Fig 65 d 6 ) passes n ea r l y pa ra ll el to th e top o f t he caps ul e a nd cut s o ff

.
, ,
 THE B I OL O G Y OF A PL A N T
132
.

f rom it the cent ral cell or arc h espo ri um I n th e pari eta l cell s f urth er .

r i l r r f w h i l t h a rc h es po ri um g i v '

t th es
divi s ions fo ll o w pe pen d c u a o e s u a ce e e ,
,

i t r i t l ll r ll l t o t h e o ri g i nal pa rie tal

rise to four n e m e d a e or tap e t a c e s p a a e ,

r r i co ns i s t s o f a cent ra l te t ra h ed ral
g p ou ( F ig 6 5 g ) .
T h e
,
s p o .
a n g u m n o w
r
a c h es p o r i um b o u n de d b y f o u r t a p e t a l ce lls w h i c h i n t u r n a r e ,
en c l os ed b y
ri
th e pa e a ce s
t l ll a t th i s t i,
m e r a p i d l y m ult i p l y i n g b y d iv is i o ns perpen

x ion o f th e p l anes o f
dicular to th e e t eri o r O w i n g t o .
t h e p e cu l i a r p o s i t

FI G . 64 . (F r
m L uerss en a
o ,ft r B ur k )—I nd u i nd r p t l f Pt
e c . s a a ec e ac e o eris aq uiltna

B i r
( d a ga m ma c ) s ee n ti , fr m b l w ; A in th
o e o t i f th d g f
. e s ec on o e e e o a lea fl e t . OJ,

o ut r ( f l
e a s e) i nd u s i um ; u i n r ( t ru ) i du i um ; r r p t l
, n e e n s . ec e ac e ; 8, y o un g
s p r
o gi an a .

d i v ision the wh o l e ca psul e is now somewh at fl att ened and it beco mes s t i ll ,

more so by the fo rma t io n a l o ngth e edg e o f a pec ul i a r s t r uc t ur e ca ll ed th e

ringo r a nnulus wh o s e f unc t i o n is th e r u t uri n l

, p go f th e ca p s u e a n d t h e

l iberat io n of th e s po res The a nnul us is fo r med by a number o f parallel

.

trans v ers e par t i t i o ns ( Fig h i j ) wh i c h s ubdi v ide th e pe ri ph era l

.
, , ,

c ell s o f o ne edg e o f th e capsul e unt i l a cer t a i n numbe r o f cells h a v e been

for med Th ese th en proj ect upon th e capsul e ( Fig 65 j ) and fo r m an in

. .
,

c o mpl et e ri ng( Fi
g 65 k ) .
, .

Meanwh i l e the t apeta l cells somet i mes subdi v ide s o as to f o rm a doubl e

ro w ( Fig 65 h ) and soo n a fter wa rds a re a bs o r bed space bei ngt hus l ef t
.
, , ,
 DE VE L OPME N T OF S P OR A N GI A 13 3
’
.

FI G . tr
65 . (A f —D v l p m t f th
e L u erss e n )p r gi f A p id ium Fili m
. e e o en o e s o an a o s x as,
w hi h i l c ly i m i l r t th t f P t i
s c os e th s y g p r gi um t d i g
a o a o er s . a, e o un s o an s an n
up th
on p i d rm i ll f r m w h i h i t h j t b d ivi d d ; t h p r xi m l
e e e s ce
- o c as us ee n e x, e o a

ce l l ut ff f m th p r gi m t f rm th p d i l d p p r t th p l ;
c o ro e s o an u o o e e ce an su o e ca su e

a, 1 t h fi t p rt i t i
. e i th rsp ul ; b 1
a d 2 th fir t d
on n d p t iti ;
e ca s e , an , e s an se c o n ar o ns

c,l 2 4 th fi t
. , , d d fe rth p r t i t i ; d d
rs , sec o n r ,
ti
an f th ou a o ns an e a re c o s s sec
-
o ns o e
ca p ul h w
s i gth bli q u p i t i f t h p r t i t i
e s o n e o d p i ll y th t f th
e os on o e a o ns, a n es ec a a o e
th i rd ; f l t r t g 0 t h r i gi f th t p t l ll d th f rm t i f th
, a a e s a e ; . e o n o e a e a ce s an e o a on o e
ar h p ri um ; h di i i
c es o f th t p t l ll d th f rm t i
, v s on f tho p r e a e a ce s an e o a on o e s o e
m th r o l l ; 1 f ur Sp r
e -
ce th y ri gi t i t h S p r m th r ll i i k
s , o o es a s e o na e n e o e o e -
ce s : , , ,

th e ul u d ri p
a nn p r gi m i
s an ur f vi w ; 1 p r i p h r l ll
e s o an u , n s ac e e 7, e e a ce s ; a r,
ar h p r i um t t p t l ll ;
c es o ulu , a e a ce s a n, a n n s .
1 34 THE B I OL OG Y OF A PL A N T .

r th enl a rg emen t o f t h e a rc h espo ri um e l a t ter no w

f or th e g o w a n d Th .

d i v i des—fi rs t i nt o 2 th en i nt o 4 8 a nd fi na ll y 1 6 cells th e mo th er cells

, , , ,
-

of th e sp o res Th ese remai n f o r a t i me c l os el y uni t ed b ut e v ent ua l ly

.
,

s epa ra t e and ag ai n s ub di v i de eac h i nt o 4 da ug ht er cell s ( Fig 65 l) The,

-
.
, .

6 4 c el ls th us fo r med a re t h e as ex ua l sp o res I n th eir ma t ure s ta t e th ey .

h a ve a t et rah edral for m a nd cert ai n ex te r nal ma rk i ngs i ndica ted in Fig s , .

63 66,
E ac h s po re acq uires a doub l e memb ra ne v iz an i nner ends
. , .
, ,

sp ori um del i ca t e a nd wh i t e a nd a n o ut er exo sp o riu m yel l o wi s h b ro wn

'

, , , , ,

h ard a nd sc ul pt ured o ver t h e s ur face wi th v ery cl ose a nd fi ne b ut

, ,

irregul ar war t y ex c rescences

,
.

Ger mi nati on of the S pores . Dev elopment of t h e Pr ot h alli um .

I n the br ake the spore s ripe n in Jul y o r A ugust and a re set

free by rupt ure of the s pora ngiu m u nde r
the strai n exe rted by the el astic a nn ulu s as ,

i ndicated in Fig 6 3 G er mi natio n of the . .

s pore s nor mall y occu rs o nly a fte r a c o ns idera

ble pe riod ( pe r h ap s not befo r e the foll o wi ng

)
sp r ing ; it begi ns b
y a ruptu r e of the e xos o
p

F1 0 66 (A ft r
e FI G 67 (A ft re S umin s k L F G e rm in a t Fro 68 (A ft r S um in
— r
. . . . -
. . e
S um ns
i ki ) i ngs po r f M i m l l A in n
es o er t: ser a a a sk i ) V e y y o un gp ro
.
.
.

Si gl pr f r ly t g ; B ft r t h pp r n
.

n e s o eo ea

f n tr n v r
s a e , a e e a ea a ce th l i
a l um of Pte fl s.
Pt w err ul se r a o o
p rt i t i n ;
e a
p rs e se a o 8. s o e h i
s o w n gt h e s po re ( s ).
ta .
pr t n m ; rh i z i d
o o e a r, o .
t wo rh i z i
o d s ( r) , an d

th nl
e e gin g x tr
ar e e m
it y .

rium
which is p rob abl y i mmediately du e to an imbibitio n of
w ater The spore bu rs ts i r regul arly along the bo rde rs of the
.

py r mid al su rface s and fro m the o pe ni ng thus fo rmed the en do

a
s on um
, o

p p r otr ude s as a p apilla filled with p rotoplas m in which

n u merou s chlo r oph ll
y bodie s soo n appear
-

Thi papilla is k no wn as the p rotmze ma o r first po rtion of

s
,
t he proth alliu m Fig
.
( I t develop s ve r q uickl i nto a sto ut
y y
.

c l i ndrical protru sio n divided i nto cell s joi ned end to end
y .

C lose to the spore o ne or more rh

izoi ds are put down f r o m the
 D E VE L OP M E N T O F THE PR O THA L L I UM . 1 35

g r ow ing proto n e m a to se r ve as an chor s a nd root s A t the oppo .

s ite or di st a l en d lo ngit u di nal p a rtitio ns s oo n appear Fig

( 68) .

w hich speedil y co n ve r t thi s po r tio n i nto a bro ad fl at pl ate at

fi rs t o nl y o ne cell thick but eve nt uall y several cells thick along
,
“
the median li ne Thi s thicke i g is the so c alled cu shio n
. n n ” -

see Fig The w hole p r othalliu m is n o w s o mewh at s p at ul ate

( .

( Fig . but by further gro w th a nte r io rl

y by a n a pic al cell o r ,

o ther w ise the wide r end beco m es

,

s till more flatte ned and hea r t

s h a ped or eve n kid ney shaped -
.

N u mero us r hiz oid s (se called b e -

c a use the y are n ot mo rph olog i

c all y tr ue r oots) a r e put dow n ,

a nd the whole s t r uctu r e assu mes

approxi matel y the appe a ra nce in

dicated in Fig 7 0 The spo r e
. .

me mb r anes and p r oto ne ma s oo n

f all away and the p r othalli um
,

e nte rs u po n an i ndepe nde n t exi s t

e n ce ,
bei ng r oote d by its r hi z oids
a nd h avi ng an abu nd ance of
c hloroph y ll I n the b ro ad thi n
.

pl ate of ti ss ue no subdivi s io n i nto

s te m and leaf exist s and the ,

pl ant bod y clo sel y r ese mbles the

“ ”
th allu s of o ne of the lo w est
n
p la ts S i n ce it is the precu r s o r
g
.

g
~

F
of a . ordi nary f em ”
at is s a , as s s s s a a ze s;
c all ed th e
“
p 7 0tfi a llu g
*”
o r p r o y un g th r i di nd o n um r u an e a, a e o s

hl r p hy ll b d c o o o i es
thalli u m
-
.

The cu shion fo rms a p r o mi ne nce on the lowe r s ide ; upon

its po ste r io r pa rt mo st of the r hi z oid s a r e bor n e .

S exual Org ans of th e Pr othallium The prothallia of fe rn s .

a re as a rule bi sexual o r he r m aph r odite ; that is e ach i ndi vid ual ,

posse sse s both male and fe male organ s B ut the l atter appe ar .

so me w h at l ate r than the for me r and poo r ly n our i shed proth allia
,

o fte n bear o nl
y m ale o r ga ns though the
y will f r eque n tl
, y develop
fe male organs al so if pl aced in better ci rcu msta nce s .

The A n theridia o r m ale organs are hem i sphe r ical p r o mi

, ,
1 36 THE B I OL OG Y OF A PL A N T .

occu rr i ng upo n the poste r io r pa r t an d the u nde r side

nences
the p r othalliu m ofte n among the r hiz oids Whe n fu lly f orms
,
.

Fig s 7 0 7 a the r idiu m co sists of a m a s of r ou n ded l

( 1 )
.
a n n
,
n s ee

m id m tt ll ) e n veloped b y a m e mb ra n e one ce
(sp er a to zo a e r ce s -

thickness .

FI G . f r i n k i l i ghtl y m d i fi d )—A dult p th llium f P teria ak

( A t e S um s , s o e . ro a o serr
see n f m b l w h w ingth rh i z i d ( r) t th p t i r nd th d p r
ro e o , s o i n e o s a e os er o e , e e e ss o

th n t r i r nd ; t h
e a eu hi n n
o r th l tt r b ri ng ( in th i
e e c ) f ur s] o ea e a e ea s c ase o a re

g ni o A m ng tha . rh i z i d tho ( p h r i l ) n th i di
e Th hl ph s
o s a re e s e ca a er a . e c o ro

b di o nl y h wn in th ll f th br d p l t f t i u n t i t ut i ng1
es o a re s o e ce s o e oa a e o ss e co s

p r th l li um
o Ju t b v t h
a n t r i r d pr
. i ni
s na p r th ll i um
o f 1
e e a e o e es s o s s ee a o a o
n t ur l i z
a a s e.

E m 71 ° °
ft r S t
(A g )—
e M t ur n fl r er . a e a Fm 72 .
—Di gr m i ll t t h
. a a to u s ra e t e
th er id inm f Pt n th i d i m A v
‘

c ulata P p i p h
O e a s rr er e gin f
o an an er u
g t g B
.
. . .
ra l c e ll : m m th r ll f th p r
s . o e -
ce s o e s e y ou n ld r
s a e: igi , o e ; 0, or i

ep i d rm l ll nl rg d b m th
e a ce e a e , o

cell f th n t i r nth idi um

o e e e a er .
 MA L E GE R JI CE L L S . 137

Th e mode of o rigi n of th e mo th er
ll s difl ers considerabl y in di ff erent
-
ce
’

f erns b ut in a ll cases 1 8 ess en t i a ll y as f o ll o ws : A n o r di na ry cell o n th e

,

lo wer s ide o f t h e p ro th a ll ium s well s a nd f o r ms a h emis ph eri ca l or do me

s h a ped pr o j ec ti o n wh i c h is s o o n sepa r a ted by a pa r t i t i o n f ro m th e o ri i nal
,
g
c ell ( Fig Furth er di v is i o ns th en f o ll o w in th e do me s h aped cell such
“ -
.

t h a t a c en t ra l ce ll is l ef t s urro unded by a
,

la yer o f pe rip h e ral cell s ( Fig By te .

p e a ted d i v i s i o ns t h e c ent ra l cell s pl i ts up

into th e s per mato z o id moth er cells ( Fig -

.

Withi n each m othe r cell the p r oto -

plas m a rra nges its elf in a peculi a r

S pi ral bod y the sp emn a toz oid which
, ,

F 73 ( A ft r H fm i t r ) I G . . e o e s e .

L t r t g i th d v l pm t a e s a e n e e e o en
Whe n the m atu r e anthe ridiu m is f th ri d i um f Pt i o a n an e o er x ser

moi ste n ed the pe r phe ral cells s e ll

i w ul t p p i p h r l ll ; r a a .
, er e a ce 6,
n tr l ll fr m w h i h t h
,
ce a ce o c e

a n d thu s p re ss out the m othe r cell s p rm t z i d

-

m th r ll s e a o o o e ce s

a nd s pe r mato z oid s ( Fig 7 The.

r i a s e.

latte r escape fro m the m othe r cell s and sw i m about ver y activel y -

in the wate r They appe ar as naked si ngle cell s of a peculia r

.
,

c o r ksc rew s ha pe an d bea r u po n the fi n e r s pi ral s n u m ero us ex

,

t re mely active cilia (p by .

which they are drive n s wiftly

th r ough the w ater .

The A rc/zeg on ia o r fe male ,

(A ft r L u
e ers se n . )—
B r ti u s n gof FI G . 75 . (A f t r S t r bur g r )—M tu
e as e . a re

sc a pe of th e a rhgc e on i u m h w i gth o p h r
, s o n e o s e e

s pe rm t z i d
a o o s. a n, a n th r i di um :
e mc
.
. ( ) th
o , e n ec k ( n) , a nd m uc us ( m) is .

s u in gf ro m t h e mo u th of th e c a n a l.
 38 THE B I OL OG Y OF A PL A N T .

o rgans ( Figs 7 0 desc ribed fo r the fir s t ti m e by S uminsk i

.
,

in 1 8 64 like w i s e ar i se fro m si ngle s upe rficial cell s of th e p r o

,

thalli um They are situ ated al mo st excl us ivel y upo n the cus hio n
.

n e ar its a n terior or apical extre mity a nd he n ce at the bot to m of ,

the anterior depre ssion ( Fig S i n c e t hey appea r late r tha n .

the antheridi a they are n o t lik elv to be fe rt iliz ed by s pe r mato

,

z oids de sc e nded fro m the sa me spo r e Thi s phe no me no n of .

matur ation o f o ne se t of se xual or g ans o f a bisex ual i ndivid ual

before the ripe ni ng o f t he o ther set is a co mmo n fea tu r e am o ng
pl ant s and is k no w n as d ichog
,
a my The r e is reaso n to believ e .

th at i mpo rt ant adv ant age s a re gai ned by thus se c u ri ng c r oss fe r -

“
tilizatio n and p r eve nti ng self fertili z ati o n or bre edi ng in and -

”
in .

I n th e dev el opment of th e a rch e g

o n i um th e o rigi na l cell enl a rg es ,

co mes mewh a t do me s h aped,

so -

a nd d i v ides by t ra ns v erse pa rt i t io ns
th ree cells : a pro x i ma l im ,

bedded ih th e t iss ue of th e

pro th all i um, a middl e , a nd a

di s ta l do me s h aped ll ( Fig
-
ce .

Th e f at e o f t h e pro x i ma l
cell is uni mpo r t a nt The dis .

tal cell g i v es ri se by di vis io n

to a c h i mney l i k e s t r uct ure -

the neck ( Fig s 75 wh ic h .

,

FI G . 76 .
—Di a gr
i ll u t t a m to s ra e

th r i gi f n r h g i um
e o n o a a c e on .

A , rl y t g ; B l t r
a n ea s a e . a a e

t g; A
s a e th ri gi n l p i
, a, e o a e
d rm le all l rgd ; B th F 77 ( A ft r S tr burg r )—D v l pi ng r h
ce en a e . a. e I G . . e as e . e e o a c e

b l ll ;
a sa th
ce tr l g i f Pt ie c en
ul t A y u g t g; B
a or on a o er s serr a a .
, o n s a e .

c ana l ll ; t h n k l l
ce c, ld r ; n k ;
e ec n l ;
-
ce o ph . o e 71, ec c, c a a o, o s ere.

encl ose s a ro w o f cel ls ( ca na l cells ) deri v ed f ro m th e o ri i na l mi ddl e ce ll

g -

( Fig s 75 . Th es e a fter wa rds beco me t ra ns fo r m d i nt o a muc i l ag

, i no us e

s ub s t ance fill i n
ga ca na l l eadi ngth ro ugh the neck f ro m t h e o utsi de to the
oosp h r e ( Fi e
g w h i h l ari ses f ro m t h e o rigi nal middl e cell at its
c a
. so “
 FE R TI L I ZA TI ON A N D D E VE L 0P M E N T . 9

pro x i ma l end Th e oos p h ere is th e i mpo r t ant f ema le germ cell to wh i c h

all
-
-
.

t he nec k a nd ca na l cell s
-
a re mer el y access o ry .

Ferti li zati on or I mpr egnat ion . Fe r tili z ation o r the sexual ,

a c t, pe rfo r med as follow s : S pe r

is
m ato z o ids in vast n u m be r s a re at
t rac ted to the mou th s of the a rche
go ni a and the re beco m e e n ta ngled
in the mucilag e ( Fig In .

favo rable case s o ne o r mo r e wo r k

thei r way dow n the mucilag inou
ca nal and at le ngth o ne pe net rate s
,

and f use s wi th the obs ph ere .

I t is k no wn th at o ne spe r ma t o z oi d is
eno ug h to fer t i l i ze t he oo s ph ere a nd F
( A f t r S tr bur g r )
,
78 I G. e as e

pro ba b l y o ne o nl y pe ne t ra tes it i b ut S8 V
. .

M u th f r h g n i um f P l ”

o o an a c e o o e

era l a re o f t e n seen in th e m uc i lag i no us i ml t r w d d w i th p r r -s ser a a. c o e s e

m id t i mgt ff t
c a na l I t h as bee n s h o wn th a t t h e muc i
.
‘ a O ZO s S r v o e ec a n en

u n a c e'
l age co nta i ns a s mall a mo unt ( a bo ut
o f ma l i c ac id wh ic h pro b a b l y ac t s bo t h a s a n a tt ra c t i o n to th e sper mat o
,

zo ids a nd as a s t i mul us to th eir mo v e ment s Pf efi er ha s pro v ed th a t

'

c a pi ll a r y t ubes co nt a i ni nga t rac e o f a ma l a t e in so l ut i o n a re as a tt ra c t i v e

.

to t he s per ma t ozo i ds as is th e muc i l ag e in t h e cent ra l ca na l a nd

ph e ,

no mena o f th i s k i nd ( ch emiota x is ) h ave rece ntlv been s h o wn to be co mmo n

and hi g hl y i mpo rta n t .

The e nt r a nce of the spe r mato z oid i nto the ovu m and its
fusio n with it ma r k an i m porta nt epoch in the life hi sto ry of the -

fe r n The oosphe r e is fro m thi s i nsta nt a new and ve ry diffe r

.

ent thi ng v iz a n embry o a nd is k now n as the oosp ore I t is

'

.
, .
, ,

n o w the fi r st stage of the asexual ge ne ratio n though it is sti ll ,

m ai nt ai ned for so me ti m e at the expe ns e of the sex ual ge neratio n

o r oéq oho re ( p
’

Gr owt h of th e E mbr y o The oespo re or o ne celled e m bry o nic

.
,
-

s poropho r e
( p n o w r apidl y beco.m e s m ulticell ul a r b
y di
y idi ng fir s t i nto he misphe r es the n i nto q uadra nt s etc ( Fig 8 0 ; , ,
. .

co mpar e Fig The fir st plan e of divi sio n is approxi matel y

.

a prolo ngatio n of the lo ng axis of the archego ni um ( Fig .

The seco nd is nea r ly at r ight angles to it so th at the quadr ant s ,

m ay be desc ri bed as a nte rior and poste r io r to the fir st pl an e .

The fate of the q u ad ra nt c ells is of speci al i m porta nce -

The .
 THE B I OL OG Y OF A PL A N T
140 .

lower ante r io r qu a d r a n t as it u n de r goes f u r the r d i vi s io n g r ows

i
o ut n
to the fi rs t r oo t the uppe r a n te r io r q ua d r a n t in li ke m an

n er give s ri s e to the rh iz o m e a n d the fi r s t lea

f The m ass of .

cells de rived from the two poste rio r qu adra nt s r e mains conn ected -

ith the p r oth lliu m as a n orga n fo r the abs o r ptio n of nut ri

w a

m ne t f r o m the latte r a n d is i napp r op r iatel y ca lled the, f oot .

Fro 7 9 Fro 80
ft r H fm i t r ) —D v l
. . . .

FI G . 79 . (A e o e s e . e e op ment o f th e e mb ry o . A wi ngth e
, sec ti on s h o

l
c o s ed n ec k( 71 ) a nd r t ivi i
t h e p la n es of
q uad a n d s o n o f t h e o os po e o r e m ry o ( cm) r b .

Th e f r nd o e e o f t h p r th l i
t e r i ght
o br a l um is to t h e . B a nd C s a g e s o f t h e em
, yo
l t r th A
a e an , s h w i gt h b gi ni g f p i l r th
o f t l f rr t
n e e n n s o a ca g o w ; 1 oo ; 1 ea ; , oo ; , ,

h rh i z m
ft r K i i t z G l fl )—
r , o e .

FI G 80 ( Fr m L u
. . o D v l pm t f th
ers s e n , a m b ry f e en -
er o . e e o en o e e o o

Pt i r ul t Th fi gur
er s se r pti l
a a t i n t k v rt i ll y in th n t r
. e e s a re o ca sec o s a en e ca e a e o

p t r i r xi
os e f th p r th ll i u m p
o a i gth ugh t h l g xi f t h n k f
s o e o a , a ss n ro e on a s o e ec o

th r h g i um x p t C d D wh i h t k t r i ght n gl t t h th r
e a c e on : e ce an , c a re a en a a es o e o e s.

A d p
, th
a, an t ri r nd p t r i r gm t f th o p r ft r th i h
a re e an e o a os e o se en s o e o s o e a e s as

d ivi d d i n t h m i p h r
e Th f r m r ( ) f rm t h
o e t m th e l t t r ( p) th r t
s e es . e o e a o s e s e , a e e oo .

F h w i s l t t g t h d ivi i f t h q u dr nt r g i ngt f r m t h r t
o s n a a e s a e e s on o e a a s, o o o e oo . 8

th t m r h i z m 1 t h l f nd f t h f t ; r I d
e s e or o n t k e. n .

pi l e ea , a e oo . , an s soo a e o a ca

gr w th i d i t d in H nd I
o as n ca e a .

I r r l i ghtl y di fferent Th e l ower dev el opment is

'

n P te i s se rula ta t h e s .

ant erio r cell b ec o mes th e fi rs t l ea f th e uppe r a nte ri o r bec omes the fi rs t

por t io n of th e r h iz o me t h e l o wer po s t eri o r bec o mes th e pri ma ry roo t a nd
, ,

t he upper po st eri o r r emai ns as th e “ f oo t .

”

The seve ral parts no w e nte r u po n r apid gr owt h acco mpanied

by conti nued cell multip licatio n u nti l a s tage is reached r ep ro
-

,
 G R O WTH A N D DI FFE R E N TI A TI ON . 41

se nted in 0 Fig 7 9 A stage so mewhat late r than thi s with

,
. .
,

its attach me nt to the proth alliu m is s ho w n in Fig 8 1 A fte r , . .

this the leaf gro ws upward s i nto the air the r oot dow n wards ,

i nto t he ea rth and the y ou ng fe rn begins to shift for itself

, .

E ve ntuall y it reaches a co nditio n show n in Figs 8 2 and 8 3 . .

The prothalliu m r e mai ns co nnected

wi th the y o ung fe r n for so me ti me ,

an d m a r e adil be fo un d in thi s
y y
conditio n attached to fl o wer pots in -

hot hou se s etc

-

,
B ut s oo ner o r
.

l ate r it falls off and the y ou ng fe rn

,

e nters u po n an e nti rely i ndepe nde nt

exi ste nce The appea ra nce of the
.

plan t and the Shape Of the leaf ( 10 F 81 ( A f t H f m i t )—Y u g I G . . er o e s er . o n

n ot a lways a t fi r s t r e se mble those m br y f P tm m ilin h w i g

‘

e o o s a t a. s o n

it tt h m nt t t h p r th lli um s a a c e o e o a
of the adult fe r n ; gro w th 1 8 al s o h y th f t ; e oo

m o re r ap i d at fi r s t seve ra l lea ve s ,
r t oo .

( 7 — 1 2 ) bei n g developed s ucce ss ivel

y in the fi r st y ear ( p .
.

Diff er entiati on of th e Tiss u es I n the ea rlie st stage s the ti ssu e

.

is n ea r ly o r quite ho moge neo us i e meristemic B ut ve ry ,

. .
,
.

ea r l y in develop me nt as the le af tu r ns u pw ar d s and the r oot

,

FI G 82 ( A e S ac
. . ft r h )—O ld r
s. e em b y f m id h i r f rn (A dv ntum) tt h d t
r o o a en a -
e a a ac e o

th e p o r th i
a ll um . S ee n i n sec ti
on 1 l f ; fi r t r t ; h b gi n i g f th
.
, ea r, s oo T , e n n o e

rh z
i o me ; p , p o r th l li um ;
a rz , rh i z i d ;
o h gm
s a r, a rc e o a .

dow n wa r d s ch ange s ta ke place which lead di rectl y to a di ff er ,

—
,

i
ent ation ni to the
’

three great s s te m s of ti ss ue epide r mal fi b ro

y ,

vascul a r and f un da m e n tal

,
The epider m al and fund am e nt al
.

syste ms take o n alm o st at o nce the peculi aritie s w hich h ave al

1 42 THE B IOL OG Y OF A PL A N T .

re ady been noted in the ad ult p The fibro vascula r system ,

. 117 .
-

of ti ssue s is di ff erenti ated a little l ater Different as the ti ssue s .

of the three syste ms are it is pl ai n fr o m thei r mode of origi n ,

th at all are fundame nt all y of the sam e natu re bec au se of thei r

de scent fr om the same ance stral cell ; he nce ever y cell in the
pl ant partake s more or le ss co mpletel y of the nat ure of ever y
othe r cell The re sembl ance s are pri mary and funda me nt al th e
. ,

differe nc e s sec ond a ry and de rived .

A nd wh at is tr ue of the fe r n in this
r e spect is equall y t ru e of all othe r
ma ny celled o rgan i sm s -
.

C ours e of t h e Fib r o v as cular B undles -

.

C er t ai n fea t ures o f t he dispos i t i o n a nd

c o urs e o f t h e fi b ro v as c ul a r b undl es in th e -

e mb ryo a nd in th e a dult ma y co nv eni en tl y

be s t udi ed a t th i s poi nt Fro m t h e po i nt .

o f j unc t i o n o f t h e b undl es o f t h e fi rs t l ea f

and fi rs t roo t ( Fi g s 79 8 1 8 2) is dev el o ped .

, ,

o ne c e nt r a l bundl e t ra v ers i ng t he young

r h iz me a nd sendi ngb ranch es i nt o th e new o

l eav es a nd roo t s unt i l 7 — 9 l ea v es h a v e been

fo r med A f t er th is t i me t h e r h iz ome .

fork s a nd the co urs e o f th e fi b r v as cul a r ,

e -

b undl es in eac h fo rk is h encefo r wa rds com

F
fig
t
ac
ii h fi n ij zzfx
’

h d t th p t h umm n t
e
n gg g g
o e
A l a t era l depress i o n a ppea rs in
fz the cent ral b undl e o f eac h st em ra idl
p o und

ro
a

a ' , p y
.

l f 1 2 t h fi r t nd
ea : , n d i nc reases in de th
. e s
p a nd
a sec o
soo n di v i des t h e ,
roo s t .

b undl e i nt o two o ne upper a nd o ne l o wer , ,

W h ch a re b es t r ecog
l niz ed in o ld s eci mens ( Fi
p g Wh e n th e fo rk ed .

s h o t s h a v e reac h ed a l en th o f a b o ut t h ree i nc h es th es e b undl es s end o ut

o
g ,

at a s mall a ng l e t o wa rds the peri ph ery th i nner fo r k ed b ra nc h es wh i c h ,

s o o n u ni t e ag ai n to fo r m a net wo r k nea r t h e e i der mi s r m

"

p Th e u p p e o st .

o f th es e b r anch es wh i c h
passes in th e medi an l i ne a b o v e th e a x i l e b undl es
, ,

is us uall y somewh a t mo re f ull de v el o ed a nd a l mos t as b ro ad as t h e lat

y p ,

ter . Th i s s t r uc t ure is g enera ll y r et a i ned in th e ma t ur e r h iz o me ( Fig .

4 8 w) Th e numb er o f peri ph era l b undl es ma be a s rea t as t wel v e in t h e

, .

y g
c ros s s ec t i o n
-

Th ey anas t o mos e in th e v i ci ni t o f th e l ac e o f i nse rt io n o f

y p
.

eac h f ro nd a nd th us f o r m a h o ll o w c l i ndri c a l n t w r k h v i n
,
y e o a gel o nga t ed , ,

mesh es ; but no c o nnec t i ngb ra nch es bet ween th em a nd th e two ax i l e

b undl es a re f ound a nywh ere in the r h izo me The l a tte r f o ll o w a n en .

t i rely iso l a t ed co ur se witb in t h e c ree i n s t em b ra nch es fro m th em

p g
-

S ee h o we vr De B ary Comp A r ga m r ns
Fe
P h a ne p 29 5
. e , na t o s a nd
, . .
, . .

O xf o rd , 1 884 .
 EX CEP TI O NA L M OD E S OF D E VE L GP ME N T . 143

enter th e l ea v es a n d i t is o nl y i ns ide th e l ea f s t a l k th a t th ese r ami fi ca t i o ns

-

a re met b y b ra nc h es f r o m t h e peri pher a l net wo r k Th e b undl es o f th e .

roo t s arise o nl y f ro m th e peri ph er a l b undl es b ut th o se o f l ea v es as al ready

, ,

s ai d
,
rec ei v e b r anc h es f ro m bo t h a x i ll ar y a nd periph era l bundl es Two .

th i ck bro wn pl a tes (s clerotic p rosench y ma ) lie bet ween th e i nner a nd

o ut er s ys t e ms o f b undl es a nd a re o nl y s epa r a t ed f r o m o ne ano th er a t th e
,

s i des b y a na rro w b a nd o f pa renc h y ma Th ey a re o f t en j o i ned o n one si de

.

o r even o n bo th in th e l a tte r c a s e fo r mi nga t ub e wh i c h sepa r a t es t h e

,

t wo s ys t ems o f b undl es ( Ho f mei s t er )

. .

A pog a my I n r a re cases e g in Ptems cretzca th e or di

'

A p os p ory
’

.
. .
, .
, ,

na ry a l te rna t i o n o f g enera t i o ns in t h e l i fe cy c l e o f f er ns is a bb r ev i a t ed b y
-

t h e o mi ss i o n o f t h e sex ua l proces s and t h e i mmedi at e v eg

, et a t i v e o ut growt h
o f t h e s po ro ph o re f ro m th e pr o th a ll i um ( ap og a my) I n o th er cas es th er e .

is a n o mis si o n o f th e spo re s t ag e a nd i mmedi a t e veg

,
et a t i v e dev el o pmen t

o f th e ooph o r e f r o m th e f r o nd ( p os p o ry) a ( c f Farl ow Qua rt Jo urn '

. . .
, .

Mic S cienc e 1 8 7 4 ; De B a r y B ota n Zeitzm g 1 8 7 8 ; Dr uery etc Journ

. , ,
.
, , .
, .

R oya l M ic S oon, 1 88 5
.
, pp 9 9 . and
 C H A PT E R X .

THE BI O L O GY OF A PLA N T ( Continued).

Th e P h y s i o lo gy of t h e Fern .

THE b r ake like the earthwo r m is a li mited po rtio n of o rg

, ,
an .

ized matter occupy i ng a defi nite po sitio n in s pace and time It .

is bou nded o n all side s by m aterial pa rti c le s so m e of whi ch may ,

be li v i ng but m ost of which a r e lifele ss The ae rial po rtio n is

,
.

i mmer sed in and p r e ssed upo n by an i nvi sible fl uid the atm os ,

p h ere
,
while the u nde r groun d por t io n is s u k in a de nse r
n
m edi um the earth w hi c h like wise ac ts u po n it
, ,
A t the sa m e .

t i me the fe r n react s upo n the air and the ea rth ,

duri ng its life an eq uilib r iu m which is di st urbed an d

w av as the life of the pla nt d ra ws to a clo s e .

Th e Fer n and it s E nv i r onment Those po rtio ns o f s pac e

.
,

e a rth and air w hich a re neare st to the b rake co ns tit ute its i m m e
,

di ate e nviro n me nt B ut in a w ide r a nd t r uer s e nse the e n vi r o n

me nt i nclude s the w
.

hole univers e ou t s ide the plant To pe r ceiv e .

the tr uth of thi s it is o nl y ne c e ssa ry to obse rve h o w profo u n dly

a nd di r ectl the pla nt is affe c ted b ra s of l ight which t ra vel to
y y y
it fro m the sun o v er a di st ance of ma ny m ill ions of mile s o r ,

h ow e xtremelv se nsi t ive it is to the alte r natio ns of day a nd n ight

o r of s unnn er an d w i n te r The pl ant is fi tted to make c e rtai n
.

e xch ange s with its e nviro nme nt d raw i ng fro m it ce rtai n for ms
,

o f matter an d e nerg an d ret ur ning to it mat te r and e nerg

y , y 111

o ther form s . I ts w hole life is an unc o nsc iou s struggle to w r e st

fro m the enviro nme nt the me ans of s ub si ste nce ; death and decay
m ark its fi nal and u nc o nditio nal su r re nder .

A dapt ati on of th e Or g ani sm t o it s E nv i r onment We can dis .

t ing uish in P tc r is as c le arl as in L u mbm c ua the adapt a tio n of

’

y
'

the organism to its e n v iro nme nt The ae rial p a rt of P tcms

'

m ust be fitted to m ake exch ange s with and m aint ai n its li fe in , ,

the at mosphere w hile the underground pa rt m u st be si milar ly

,
“ ”
ad apted to the soil in which it live s .
 A DA PTA TI ON TO THE E N VI R ON ME N T . 45

The ae rial pa rt display s ad mirable adaptation in its stalk which ,

r ise s to a poin t of va ntage for procu ri ng ai r and light and in its ,

bro adl y spreadi ng to p w hich is covered by a skin tough and

, ,

i mperviou s to p r eve nt u n due ev apor ation and c onseque nt de sic

,

catio n y et t ransluce n t to allow the s un s ray s to r each th e

, ,
’

s tarch m aki n g tissu e w ithin

-
The rhi z om e al so with its poi nted
.
,

te r m i nal buds its elo n gated root s ar m ed with bori ng tip s and
, , ,

its thick fle sh y pare nch ym a for the sto rage of food is ad m irabl y
, ,

a dapted to its o wn s pecial s ur roun di ng s I n order to r e ali z e

.

thi s we hav e o nl y to i magi ne the fer n to be i n ve r ted the aeri al

, ,

po r tion bei ng pla nted in the earth and the u ndergro und po rtio n
,

lifted i nto the air and expo s ed to the wi nd s and s u nshi n e U nde r .

the s e ci r cu ms ta nce s the w ant of ad apt at io n of the p art s to t hei r

respective e n v iro nme nt s w o uld s peedily beco me app a re nt .

Y et diff ere nt as the se p art s no w a re they h a ve o rigi nall y

,

s p r u n g fro m the sa m e cell M ore rece ntly they w ere b a rely dis
.

ting uish ab le in a m a ss of ti ssue p a rt of w hich tu r n ed u p w a rd s

,

i nto the air w hile another pa rt tu r ned do wn ward s i nto the e art h
, .

B ut as devel o p me nt we nt o n the a eri al an d un dergro un d pa rt s

,

we r e p r ogre ss ivel y differe nti ated th us beco mi ng m ore and m o re

,

pe rfectly ad apted to the peculi ar co nditio ns by w hich e ac h is

s u rr ou n ded .

Th us it appe ars that the h ar mo ny bet wee n e v e r y p art of the

pl ant and its e nviro nme n t is brought abo ut as in the ani mal by a ,
,

g ra d ua l
p roces s in the hi s to r
y of e a ch i ndi v id ual We can he r e .

cle a rl y see al s o the fun ctio nal ad apt atio n of the pl ant to ch ang
ing exter nal co nditio ns The e n v iro nm e nt of P ter zs ch ange s
’

periodica ll y with the r egul a r alter nation of summer and w i nter ,

a nd the pl a nt al so u nde r goe s a corre s pon di ng pe r i odic c h ange o f

s tructu r e in orde r to m a i nt a i n its ad apt atio n to the e nviro nm e n t .

D uri ng the su mmer th e ae r i al pa rt is full y developed and as a , ,

re sult of its activity starch is acc um ul ated in the rhiz om e A t

,
.

the app r oach of wi nte r the ae ri al p art die s and the pl ant is re ,

duced to the un dergro und p a rt safel y b uried in the soil Du ri ng .

the w inter and sp ri ng the starch is gr adually consumed and the ,

aerial part is put fo rth agai n as the ae ri al e n vironme nt becom e s

o nce more favo rable to it The pla nt therefo re like the a ni m al
.
, , ,

posse sses a certa i n p las tici ty which e nable s it to ad apt it s elf to

graduall y cha ngi ng co nditio ns of the e nviro nm e nt .
146 THE B I OL OG Y OF A PL A N T .

l i ttl e
i d r t i n w i ll h w th a t ev ery f unc t i o n o r ac t i o n o f l ivi ng
A e co ns
a o s o

i r r ri b i o th e s ame g r
t h ng y
s ma be e ga d e d as co n t ut ng t ea t e nd v iz b ar ,
.
,

mony wi th t h e env iro n men t ; a nd f r o m t h is po i nt o f v iew l i fe i tsel f h as

been defi ned as th e conti nuous adjus tment of i nterna l rela tio ns to em
“

”
t erna l rela ti ons .

N ut r iti on The fern doe s work

.
I n pu shi ng its stem .

t hrough the s oil in li fti ng its le a

,
ve s i n to the a ir in moving ,

food matte rs fro m po int to poin t in b uildi ng ne w t issue in the

-

, ,

proce ss of reproduc t io n and in a ll oth er fo rms of vital actio n

, ,

the pl ant expe nds e ne r gy Here as in the anim al the imme .

, ,

di ate source of e nergy is the livi ng p rotopl as m wh ic h as it , ,

live s bre ak s do wn i nt o si mpler co m po un ds He nce the need of

,
.

an i nc o me to s uppl y the p o w e r of doi ng w ork .

Th e I ncome The i nc o me of the fer n lik e th at of the ea rth

.
,

wor m is of two ki nd s v iz matte r and e nergy but u nl ike tha t

, ,
.
, ,

a n i n co me b
o
f th e worm i t is n ot chie fly of f oo ds u t on ty f
o the ,

ff M atter e nt e r s the pl a nt in the liquid o r

'

r a w ma teria ls o ood .

g ase o us fo r m by d i f u s iou b o th fro m the s oil th r o ugh the r oots

,

liq id nd fro m the at mo s phere th rough the leav es ( gases )

( u s
) a ,
.

We h ave here the direct absorptio n i nt o the body pro pe r of food

s tuffs p reci sel as the ea rth w or m t a ke s in w a ter a nd ox ge n
y y .

E nergy e nte rs the pl ant t o a s mall exte nt as the pote n ti al e nergy

, ,

o f food stuffs but co m e s in p ri n cip all as the ki neti c e nerg of

y y
-

s unlight ab sorbed in the le a ve s The ta ble c u p 1 4 7 show s the . .

precis e natu re and the mo r e i mpo rtan t sou r ce s of the i nc o m e .

O f the sub st ance s the solid s ( salts etc ) m ust be d issolv ed

, ,
.

in water befo re they ca n be take n in W ate r and d iss ol v ed sa lts .

c onti nually pass by diffusi o n fro m the s oil i nt o the roots w he re ,

together they co nstitute the sa p The sap t ra vel s t hr ougho ut .

the whole pl ant the mai n tho ugh not the o nly cause o f mo ve
,

m e nt bei ng the c onst ant tr a n sp i ra tion ( evapo ratio n of w a te r

) y
v apor fro m the le ave s e speci all y th rough the sto mata Th e , .

gaseous matters ( carbon dioxide oxy ge n nit rog e n) e n te r the , ,

pl ant mai nly by diffu sion fro m the at m os phe re are dissolved by ,

the sap in the leave s and el se where and thus m ay pass to every ,

p o rtion of the pl ant

The Manuf act ur e of Foods—
.

espec i ally S ta rch P teris o w es .

i ts po wer of absorbi ng the e n erg of light to the

y s un chlorop hy ll

S p n r P ri wip l s
e ce ,
f B i lg y l l i p 80 N Y
e o A pp l t n 1 88 1
o o , vo . . . . . .
,
e o , .
 IN C OM E OF TH E PL A N T . 147

bodies or chrmna top bores for pl ant s w hich like fu ngi etc are , , .
,

devoid of c hloroph y ll are unable th us to acqui re e nergy E nter .

ingthe chlo r oph y ll bodie s the ki n etic e n ergy of sunlight is a

-

,
p
p lied to the deco m po sitio n of c a rbo n dioxide
( C O ) an d w a ter ,

A fter pas si ng through m an ifold b ut i mperfectl k now n

y
processes the ele m e nt s of the se s ub sta nce s fi nall y re appe ar as
,

s tarch ofte n in the fo rm of gr anule s i mbedded in the

chlo r oph y ll bodi e s and f ree oxy ge n mo st of which is ret ur ned
-

, ,

IN COM E O F P TE R I S .

to the at m o s phere Th us the leaf of P teris in the light is con

.
.

tinually ab sorbing c arbo n dioxide an d givi ng fo rth free oxy ge n .

C a rbo n dioxide and w ater co nt ai n no pote nti al e nergy s i nce ,

the affi nitie s of their co nstitue nt ele me nt s are com pletel y sat
isfi ed S ta rch however co n tai ns pote n ti al e nergy s i n ce the
.
, , ,

molec ule is rel ativel y u nstable i e ca p able of deco m po sition ,

. .
,

i n to si m ple r stabler m olecule s in w hi c h s tro nger affin itie s a r e

,

I t h as be g r ll y b l i v
en ene a e e ed th t p l nt s
a a un b l to m k
a re ua ef f ree a e se o

a tm os p h r i ni t r gn b ut r
e c o nt e ,
ec e inv t i g t i n h v d i pr v d th i s vi w f r
es a o s a e s o e e o
1 48 THE B IOL O G Y OF A PL A N T .

sati sfied . A t
nd hi s is d u e to the fact that in the m a n ufactu r e
of starc h in the chlo r oph y ll bodies the kinetic e ne rgy of sunlight
-
.

a was
exp e n ded in lift in g the ato m s i n to po s iti o n of va n tage ,

w the m with e n e r g of po s itio n I n this way so me

th us endo i g n y .

of the r ad iant and ki n etic en ergy of the sun co me s to be s tored

pote tial e n e r g in the s t a rch I n sho rt P tc ris like all
-

up as n y .
, ,

green pl ant s is able by co ope r atio n with sunlight to use s im ple

,
-

te r ial c rbo d i oxide w at r ox ge etc ) poo r i eu

r a w m a s ( a n e
y n ,
n , ,
.

or devoid of it d of the m to m a n u a ctu re f ood i e

f
er y
g a n ,
o u t ,
. .
,

co mplex compound s rich in a v ailable pote nti al e ne r gy We .

s h all see hereafter th at thi s po w e r is possessed by gr e

en pl ants

alo ne ; all other o r ganisms bei ng depe nde nt fo r e nergy u po n the

p o te nt i al energy of read y m ade food Thi s mu st in the fi rst
-
.

i nst anc e be p r ovided f o r the m by gree n pl ant s ; and he nce with

o ut chlo r o ph y ll be a ri ng pl an t s a n im al s ( an d colo r le ss p la n ts as
-

well ) app are n tl y c o uld no t l o ng exi st .

The pl ant ab sorb s al s o a small a mou nt of kin etic e ne r gy in ,

depe nde ntly o f the sunlight in the fo rm of hea t ; thi s ho w eve r

, , ,

is prob abl y n o t a source of vit al e nergy but o nl y co nt ribu te s to ,

the mai nte nance of the bod y te mpe ra t ure .

Ci r culati on of Foods I t is c hiefl y in the g ree n ( chlo roph y ll

.

beari ng) par t s o f the pl ant s and in the pre se nce o f sunl igh t th at
, ,

food m anufactu re g
-
o es o n S o m eho w the n the w ate r ab sorbed
.
, ,

by the root s mu s t be tr a nsp o rt ed to the leave s an d the s ta rc h ,

made in the le a v e s m ust be co nvey ed to the s ubte r ra n ea n ti ss u es .

E x actl y h o w the s e tr ansfer s of m ateri al are effe c ted is unc e r ta i n ,

but there i s re as o n to believe th a t they ta ke plac e mai nl y by the

s low proce sse s o f diffus i o n I t is c e rt ai n th a t n o disti nct organ s
.

of cir cul ati on or di strib ution s uch as the blood ve ss el s of the ,

-

e arth w orm exi st in the fer n

,
.

Met ab oli sm S tarch as has just b een see n is fi rst fo rm ed in

.
, ,

the chloroph yll bodie s B ut the for mation of s tar c h all i m po r

-

.
,
-

t ant as it is is af ter al l only the m a n uf a ctu re Of f ood as a pre

,

li mi nary to the re al p r ocesses of n ut r itio n These p r ocesses must .

take pl ace every whe r e in ordin ary p r otopl as m ; fo r it is he re

th at oxid atio ns oc c ur and the need for a re new al of matte r and
e nergy c o nseq ue ntly ari se s ( c f pp 3 2 and . S oo ner o r l a te r
.

the st ar ch gr ain s are ch anged i nto a ki n d of suga r (g luco se ,

which u nlike s tarch dissolve s in the sap and ma y

, , ,
 O UTG O OF THE PL A N T . 149

t hu s be easil y t ra ns po rte d to all pa rts of the pl ant Wherever .

the re is n eed for new proto plas m w hether to repai r previous ,

w as te o r to suppl y mate ri al s fo r growth afte r ab sorptio n in to ,

the cells the eleme nt s of the s tar ch ( or gluco se) are by the liv ,

ingp r oto pl as m in so me u n k now n way com bi n ed with nit roge n

,

a nd s ulphur ( probabl y al so with salt s w a ter ,

to for m p r oteid ,

matter The pa rticl es of this ne w ly for m ed co m po u nd are i ncor

.
-

o rated in to the p r otoplasm ( by

”
p i nt u s s u sceptio n p l )-

and in , .
,

so me way at p r ese nt sh rouded in my stery are e ndo w ed w ith the ,

prope rties of life We do no t k n ow h o w lo ng they m ay r e mai n

.

in the li v ing state but s oo ne r o r l ater they are oxidi z ed and as a

, , ,

r es ult of the oxidatio n that e ne rgy is set f ree w hich e nable s the
,

fe rn to do wo r k and p ro lo ng its e xi ste nce The oxidi z ed prod .

ucts a r e afte r wa r d s eli mi na ted ( excreted ) f ro m the cell s .

I f a l a r ge r qu an ti ty o f s ta r ch is for med in th e chloroph yll

bodi es than is i mm edi atel y needed by the protopl asm for pur
pose s of repair o r grow th it may be re co nver ted i nto st a rch
,
-

afte r journ ey i ng as gluco se through the pl ant and be l aid do wn ,

as r e se r ve sta rch in the pare nch yma o f the r hi z o me or el se ,

where A ppa re ntl y whe n thi s r ese r ve suppl y is fi nall y needed

.
,

at any po int in the pl a nt it is agai n c h a nged to gl uco se and tr ans

,

po rted thithe r I t is prob able th at new le a ve s a nd ne w ti ssues

.

g e nerall
y are al w a s fo r m ed in p a rt fro m thi s re serv e st a rch
, y ,

and no t solel fr o m new lv fo r med s t a r c h

y
-
.

I n deali ng w ith the m et a boli s m of the fer n w e m ay safel y

assu me as we h a ve do ne al re ad y f o r th e e a r th w o r m a co nstr uc ti v e
, ,

phase (a na bolis m) and a de st r uc tive ph ase (k a ta bo lis m) ; but

these te rms repre se nt m e r el y prob a ble eve nt s n o t k no wn fact s ,
.

Th e Out g o .The outgo like the i nc o m e is of t wo ki nd s

, , ,

matt er and e ne r gy but it c a nnot be so re a dil y t abulated

,
.

The plant suffe r s a nnually a gr eat lo ss both of matter and of

pote nti al e nergy in the prod uctio n of spo r e s and in the autumnal
dying d o wn of the fro nd s B ut m atter al so leave s the pl ant
-
.

d aily as ca r bo n di o xide ( in s mall quantitie s) water and ox ygen , , ,

both by diffus io n th rough the epider mi s and by tr anspir atio n

through the stomata S t rictly speaki ng the te r m outgo sho uld
.
,

be re st r icted to the ou tpu t of m atter w hich has at so me ti m e

ac tuall y for med a pa rt of the li v i ng protopl asm ; he n c e it d o e s
no t appl y to the ox ge n which is s im pl y give n o ff in the manu
y ,
15 0 THE B I OL O G Y OF A PL A N T .

facture of sta r ch or to the bulk of the w ater of eva po r atio n

, ,

hich p se t r ight through the pl a n t w ithout u n de rg o i ng any

w as s s a

chemi cal ch ange E nergy lik ewise leave s the pl ant co nt inuou sly
.

both as li eut and in the do ingOf mecha n ica l work both of whi ch ,

a re i nvolved in every vit al ac t .

R espi r ati on . I t has bee n rema rk ed that in the light

he f c uri g tar h P i t ke s in ca r bo n dioxide and
w n m a nu a t n s c
) te r s a

gives off free oxy ge n B ut if the pl ant be dep r ived of light as

.
,

ight the r ever e is t r u e a n d the pl a n t tak es in a s ma ll

at n ,
s ,

amou nt of ox yge n a nd give s o ff a cor r e s po ndi ng am ou nt of ca r

b o n dioxide .
This l a tte r p r oce ss is the t r ue b rea th ing o r r esp i

P TE R I S A QUI L I N A .

(B a l
a nce -
S h t
ee of N u t iti
r on . )

ra tio nof the pl ant and it mus t no t be c o nfou nded with that
,

taki ng in of c arbo n dioxide and givi ng off of oxy ge n whi c h is an

i nc ident in the manufact ure of s tarc h R e s pi ratio n goe s o n in .

t h e light al so pr o b abl y w ith gre ate r e ne r gy th an in d a r k ness

, ,

b ut it is the n l a rgel y ob sc ured by the other an d m o re co nspicu

o us proce ss “W
. e h a v e see n th a t e ne rg
y is se t f ree in livi ng m at

t er by a deco m po siti o n of its o wn sub s ta nce which is rea lly a ,

proce ss of o xid ati o n o r c o mb usti o n whe r e free oxyge n pl ay s

“

a n i mportant p a rt p
( 3 9 C h ap .
,
he nce the a bs o rptio n o f
.

f ree oxy gen in re spir ati o n A m o ng the p r od ucts o f the co mb us

.
~

ti on w ater and carbo n dioxide are the mos t i m po rta nt ; and this
,
 A CTI ON UP ON THE E N VI R ON ME N T . 15 1

is the o rigi n of the ca rbo n dioxide give n off I t will appe a r .

be yo nd th a t p r ec isely the same actio n take s place in the r e spi

ratio n of anima ls a nd that all livi ng thi ngs b r eathe o r re spi r e
, in
esse ntially the sa m e w ay .

l ongt i me bel i ev ed th a t a l eadi ngdi ff erenc e bet ween pl a nt s

I t was f or a

lay in t h e f ac t th a t th e f o r mer g i v e o ff o x ygen a nd absor b

e wh i l e t h e l a tt er g
, i v e o ff ca r b o n dio x i de a nd abso r b o x ygen .

k no wn th a t bo th gi v e o ff ca r bo n dio x ide and bo th req uire

oxyg
en a nd th a t o nl y t h e c hl o r o ph y ll be a ri n
, gpa rt s o f green pl a nt s are en
-

mpos ingc a r bo n di o x i de and wa t er

— a s a r es ult o f wh i c h th e
y do ( b ut in th e l i ght
k i nd o f i ncide nt a l o r by prod uc t -

I N TE R A CTI O N or THE FE R N AN D I TS E N VI R O N M E N T .

The actio ns of the e n vi ro nm e nt upo n the fer n have already

bee n s ufficie ntl y dwelt upo n ( p I t still re mai ns ho w ever
.
, ,

to co ns ide r the actio ns of the fer n u po n the e n viro nm e nt .

These ar e p a rtl y ph y sica l but mai nl y che mica l

, B y p ushi ng .

its fro nd s i nto the a ir a nd s lo wl y th ru s ti ng its rhi z o me r oot s and , ,

the soil the at mo s phere and the e a rth are alike

,

t is by its che mica l acti v it y th at it mo s t pro

n v iro mn ent A bsorbi n fro m the l atter
.
g
io xide and othe r si m ple sub st ance s as w ell
, ,

i n a re markable m et a morpho s i s .

g
raw m ateri al s o r a n i c m a tter in

eve n proteid s The se it give s .

eas ure du ri ng life an d sur ,

B ut the m o st striki ng fact

the fe r n is o n the whole co nstructive and c ap able of pro
and accu m ul ati ng co m pou n d s rich in e n erg I n this
y .

it is u nlike the earth w o r m ( p 1 04 ) and is t yp ical of gree n

.

in ge ne r a l Thu s w hile ani mal s are de stroy er s of e ner

.
,

co m pou n d s gr ee n pl ant s are prod uce r s of the m A ni

,
.

therefo r e in the lo ng run are ab solutel y depe nde n t o n

,

and a ni mals an d colorle ss p lan t s a like upo n gree n pl ant s .

are e nabled to

v i rt ue of the

powe r
15 2 THE B I OL OG Y OF A PL A N T .

i g
Phy s olo f th e Ti ssue Sy st ems The e iderma l tissue
'

y o -

p .

s erve as the s ole m ediu m of excha n ge betwee n the in ne r pa r ts 0

the pl ant and the e n vironm e nt ; they are also p r otective and 11 ,

ce rtai n r egions are useful fo r suppo r t The fu nction of rep ro .

d uctio n al so fall s upo n the se ti ss ues as is shown by the develop,

ment of the spo rangi a a ntheridi a and a rchego nia

, ,
.

The fi brO c a scu la r ti ss ue s se r ve in p ar t as a suppo rtin;

-
’

skeleto n ,
fo r which fu nctio n thei r rich ne ss in p r o se nch ym a

and thei r fi rm co n ti nuity ad m irabl y a dapt the m A n eq uall; .

impo r t ant fu nctio n h o w eve r is thei r co n d uc tiv ity si nce the;

, , ,

serve for the t r ansportatio n of the w ate r fo r ev a po r atio n by th .

le af ( tra mwira tio n ) and f o r the move me n t ( th rough the sieve

,

t ube s) of the u ndi ss o lved and indiff usible p roteid s The f u n da .

men ta l tiss ues are dev o ted eithe r to shari ng the speci al dutie
of the other syste ms as in the case of the sclerotic pare nc hy m
,
l

ab utti ng u po n the epide rm al ti ssue in the r hi z o m e ( p am .

the sc lerotic p r o se nc h y ma which a ppea rs to beh ave like the fi bro

v as c ul a r ti ssue s ; or to nutritive and m etab o lic f i mc tio ns as ll ,

the me so ph yll ( p 1 2 6 ) and t he pare nch y ma o f the r hiz o m e

. .

Th e Phy s i ol og y of R epr odu cti on I t is no t k now n w hethe r th

.

brake ever die s of old ag e B a r ri ng accide nt s gro w th at th

.
,

api c al b ud s see ms to be u nlim ited keepi ng p ,

the hi nder p a rt s o f the rhi z o me ( p 1 B ut

.

vid ual die s or no t a mple pro v i sio n agai nst the

,

is made in the ac t o f reprod uc tio n A l though .

pe ars to be usele ss to the i ndividu al and eve n ,

seriou s a nnual l o ss e s o f ma tter and e nerg et

y y ,

e v ery part of the pl ant di rectl y o r i ndi rectl y co

reprod uctive ger ms are caref ully prepa red ; a r e pr
sto c k of food s ufficie nt for the e ar li e s t s ta e s of
g
and are e ndo w ed w ith the pec uliar po w e rs an d

P teris a g u ili n a w hi c h i nflue nc e thei r life hi s tor

, y
-

and are b
y the m t r ansmitted in tu
are livi ng po rt i o ns of the pa re nt
po ses ; they co nt ai n a sh a re of protopl as m d ire c tl y
fro m the origi nal protopl asm of the s po r e f r om w hich
c ame ; and th us they serve to effe c t th at co nti nu
”
ger m pl asm to w hi c h we h ave already referr ed
-

w ith the e arth w o r m I n short r ep r od uc tio n is th

.
,
 P L A N T A N D A N I MA L COMP A R E D .
15 3

f unctio n of the plant I f we may p ar aph ra se the wo r ds of

.

M ich ael Fos te r the o é sphere is the goal of i n divid ual exi stence
,
,

a n d life is a c y cle begi nning with the o Osphere and conti nuall y
,

c o mi ng r ou n d to it aga in .

Compari son of th e Fer n and th e E a rt hwor m To the super .

fi cial obse r ve r the fer n and earth w o r m see m to h ave little o r

nothi ng in co mmo n except th at both a r e what we c all alive
, B ut .

whoeve r has st udied the p r ecedi ng p ages must h ave perceived

be neath manifold d iffere nce s of detail a fundam e nt al like ne ss
betwee n the pl ant and animal not o nl y in the sub stanti al ide n
,

t ity of the living matter in the two but also in the con st r uctio n
o f thei r bodi es an d in the p r oce ss e s b w hich the co m e i to
y y n

e xis te n ce . E ac h a rise s f ro m a s i ngle cell which is the re sult of

t he u n io n of tw o differe ntl y co nstituted cell s male and fe m ale
-

, .

I n both the p ri ma ry cell m ultiplie s an d for ms a m as s of cell s at ,

fi rst n ea rl y s im ilar b ut after wa rd s d iff ere ntiated in v a riou s di

r ectio ns to e nable the m to pe rfor m differe nt functio ns i e to ,
. .
,

eff ect a ph y s iologic a l di v i s io n of l abor I n both the ti ssue s th us

.
,

rovided are ass oc i ated m ore or le s s clo s el in to di sti n ct organs

p y
a nd s ste ms a m o ng which the va r io us ope ra tio ns of the bod
y , y
-

a re di s tributed A nd in both the ulti mate go al of i ndivid ual

.

e x iste nce is the produ ctio n of ge r m cel ls which for m the st a rt -

ingpoi nt of n ew and si m il a r cy cle s

-
.

This fu nd am e nta l like ne ss exte nds al so to m o st of the actio ns

ph siolog ) of the two orga nis m s B oth po sse ss the po w er o f
( y y .

a d a pting the m selve s to the e n v i ro nm e nt s in w hi c h the

y live .

B oth take in v a rio us fo rm s of matte r and e n ergy fro m the eu

v iro nm ent build the m up i nto thei r o w n li v i ng sub st an ce and
, ,

fin ally bre ak do wn thi s sub s t ance more o r le ss co mpletely i nto

s i m pler co m pou n ds b proce ss e s of i nte r n al co mb ustio n s etti ng
y ,

fr ee by thi s actio n the e nergy which maintai ns their vita l ao

t iv ity. A nd soo n e r o r l ater both give b a ck to the e n viro nm e n t
, ,

t h e matte r an d e n ergy w hi c h they h a ve take n fro m it I n othe r .

Wo r d s both effect an exch ange of matter and of e nergy w ith

,

t he e n viro n m e nt .

N everthele ss the pl ant an d the an imal di ffe r They d iffer .

w idel y in fo r m an d the pla n t is fixed a nd rel atively r igid whil e

, ,

t h e an im al is flexible an d m obile The body of the pl an t is

.

relati vel s olid ; th at of the anim a l co nta ins n u m erou s c avitie s

y .
15 4 THE B I OL O G Y OF A PL A N T .

The plant absorbs matte r di r ectl y th r ough the exte r nal su r fac e ;
the ani mal p ar tly th r ough the exte rnal and pa rtly th r ough an
i nter nal (ali me n ta ry) su rface The pl an t is able to ab sorb sim pl e
.

chemical co mpou n ds from the air and e a r th an d ki netic e nergy ,

from su nlight ; the an i mal ab sorb s for the mo st p ar t co mplex , ,

che mical co mpou n d s and m ake s n o nutritive use of the su n s ’

ki netic e ne rgy B y the aid of thi s e nergy the pl an t manuf ac

.

t ur es starch f rom si mple co mpou nd s carbo n dioxide and wate r ; , ,

the ani mal lacks thi s power The pl an t can build up p r oteids
.

fr om the nit rogeno us and othe r co mpou nd s of its food ; th e a ni mal

ab solutely requires proteid s in its food A nd by ma nu factu r ing .

proteid s withi n its livi ng sub sta nce the pla nt is reliev ed of the ,

n ece ss ity of c ar ry i ng o n a p r ocess of dige s tio n in o r de r to re n de r

the m diffusible for e nt r a nce i nto the bod y .

S till g re at as these diffe r e nces appea r to be at fi rst sight

, ,

all of the m with a si ngle exceptio n fade away u po n close r ex

, ,

a minatio n Thi s exceptio n is the p ower Of ma k ingf oods

. .

Pl ant s and a ni mal s diffe r in for m becau se thei r m ode of life

differ s ; but a wide r study of b iology reveals the exist e nce of in
numerable a ni mal s ( cor al s s po nge s h y d r oid s etc which hav
, ) ,
e ,
.

a close superfi c i al re se mblance to pl ant s a nd o f many plants ,

which re se mble ani mal s not o nl y in fo r m but al so in possess i ng

, ,

the po wer of active loc o motio n The sto mach of the wo r m as .

,

sho wn b its devel o p me nt is r e ally a part of the ge ner al oute r

y ,

surface w hich is folded i nto the b o d a nd the a nima l like t h e

y ; ,

pl ant therefo re r e ally ab s orb s its i nco me ove r its w hole su rf ac e

—o xygen through the gene ral oute r su rface othe r food matte rs
, ,

thr o ugh the i nfolded ali me nta ry su rface .

I n like manner it is easy to sho w that no t o ne of the di ffer

ences bet wee n the pl a nt and a ni mal is fu nda m e n ta ll im po r
y
t ant sa ve the p ower of ma k ingf oods The wor m m u s t hav e
-

co mplex r eady mad e food i ncludi ng p ro teid matte r S o m u st

-

the fer n ; but the fe rn is able to m a n uf a c ture thi s co m plex food

o ut of ver si mple co m pou nd s I n te r ms of e n ergy the wo rm
y .
,

requires read y made food r ich in po te ntial e ne rgy ; the fe rn

-

aided by the su n s e nergy can manufac tu re food fr o m matte rs

’
,

de v oid of e nergy .

He nce it appea r s broadl y speaki ng that the fe rn by th e aid

, ,

of s
ol a r e nergy is const r ucti v e and sto res u p e nergy ; the ea rth
,
 F OOD OF PL A N Ts A N D A N I MA L S . 15 5

wor m is de structive and di ss ipate s e nergy A nd thi s diffe r en ce

, .

beco mes of im m e nse i mpo rta nce in view of the fact that the
fe r n is ty pical in thi s r e s pect of all gree n pl ants as the earth ,

worm is ty pica l o f all a ni mal s .

I t w ill here afte r appe a r th at eve n thi s di ffe r e nce gre at as it ,

is is pa r tl y b ri dged over by colorle ss pl ant s like y e ast mo uld s

, ,

bacter ia etc wh ich h a ve n o c hl o roph y ll are the r efo re unable

,
.
, ,

to use the e ne rgy of light and he nce m u s t have e nergi z ed food

, .

B ut thes e orga nisms do n o t l ike ani mal s r eq uire proteid fo od

, , ,

bei ng able to extr act all needf ul e nergy fro m the si mpler fat s ,

ca r boh yd ra te s and ev e n fro m ce r t ai n salts NVh en w e co nside r

, .

th at the d isti nctive peculi a ritie s of a ni mal s can thu s be r educed

to the sole ch a rac te ri stic of depe nde nc e on proteid food we can ,

not doubt th a t the di fl erences bet w ee n pl a nt s a nd an i ma l s are of

immeasu r abl y less i m po r t a nc e tha n their funda me ntal like nes s .

It has bee n the object of the foregoi ng ch apters to give th e

stude nt a ge ner a l co nc eptio n of orga n i s ms w hether veget al o r
,

a ni mal ; of thei r s t r uctu re gro wth and m ode of actio n ; of their

, ,

po sitio n in the world of matter and e nergy and of their rel atio ns
,

to lifele ss thi ngs With thi s p reli mi na ry k no wledge as a b asi s

.
,

the stude nt is prepa red to take u p the progre ss ive s tudy of othe r
o r gani sms sel ected as co n ve nie nt ty pe s o r ex ample s I t is c on
,
.

v enient to begi n w ith low a nd s i m ple for ms of life and wo r k

gradually up war d s ; and it is e s pe c i ally de sir able to do so be

cause the re is reaso n to believe th at thi s co urse co rr espon d s
b ro adly with the path of actu al evol utio n .
 C HA PT E R XI .

THE UN I C E LL UL A R O R G A N I S MS .

has bee n sho wn in the foregoi ng p ages that the co m plex

IT
body of an adult fern o r e ar th w o r m o r of a ny of the h ighe r ,

fo r ms of life o r igi nate s fro m a s i ngle cell of m icro scopic si z e

,
.

Th is cell —the fertili z ed ovu m or o os phe r e—gives r ise by di v i

sion to new cell s whi c h in their tu r n divide g e ne ratio n afte r ,

ge ne rati o n until a full grow n bo dy is fo r m ed co mposed of

,
-

my ri ad s of ce ll s B ut the proc e ss o f cell divis io n does n o t in

.
-

thi s c ase go as f ar as co mplete c ell s ep a ra tion and the cell s do

-

not acquire a co m plete i ndividu ality They do it is t ru e ao .

, ,

quire a certai n i nde pe nde nc e o f s tru c t ure and f unctio n ; and

their i ndividu al c h ar ac te ri stic s may eve n depa rt widel y fro m
those of neighbori ng cell s (differe nti atio n) N eve rt heless they .

remai n clo sel y u nited by either mate r i al o r ph ysiologi cal bo nd s to

fo r m one bo dv The b o dy is no t howeve r to be regarded as
.
, ,

merel y an asse m bl age of i ndepe nde n t i ndividu al cell s The body .

is the ind ivid ua l ; its m o re or le ss pe rfect divisio n i nto cel ls is

o nl y a b asi s f o r t he ph ysi o l o gi ca l divi si o n of l abor ; of which
cell differe nti atio n is the o ut wa rd exp r e ssi o n
-

A ll thi s is tr ue h o w ever o nl y in the higher ty pes

, , A t the .

bottom of the sca le of life there i s a v as t m ul t itude of for m s in

whi c h the bod y co ns i st s no t of m an cell s b ut of o nl o ne and is
, y y ,

therefo re c o mp ar able in str uctu r e no t to the adult fe rn or ea rth

w o r m but to the ger m cell s fro m which th ese a ri se S uch for ms
,
-

are k now n as u n icellula r organi sms in c o ntradi sti nc ti o n to the ,

m ulti cellula r . L ike other cell s the u nicellul a r orga n is m s m ulti

l b
p y y divi sio n b ut di v i sio n is fo llo w ed soo ne r o r l ate r by co m
,
”

p lete sep a r atio n

; the daughte r cell s be c o m e e nti rel y di sti n ct and
-

i ndepende nt i ndividu al s and do no t re mai n pe r mane ntl y asso

,

c iated I n the m a true multicell ular bod the r efore eve

.

y is n r , ,

fo r med ; the cell is the in div id ua l a nd the body is u n icellula r

, .

15 6
 THE UN I CE L L UL A R B OD Y . 15 7

N eve rthele ss the one celled organ i sm pe r for ms all of the

-

ch a racte ri stic ope r atio ns of l ife A si ngle mass of protoplas m a

.

si n gle cell
,
u ni tes in itse lf the pe rfo r m a n c e of all the v a rio us

ele me nta ry fu n ctio ns which in the multicellula r for ms are distrib

uted am o ng m any cells ,
diffe re ntiated in to divers tissue s and
o rga ns . The u nicellula r fo rms ar e therefo re in a phy siologic al
“
sense as t rul y orga nisms as the multicellul ar fo rms ; and in
ma ny case s the u ni cellula r body show s a very considerable deg r ee
of diffe re ntiatio n am o ng its pa rts B ut the u n ice llular fo rms
.

a r e orga nis ms reduced to thei r lowest ter m s ; they p r e sent us with

the proble ms of life in the ir mo st ru di me nt a ry fo r m He nce .

the y may affo rd a kind of k ey to the more el abo r ate organi zatio n

We shall fi nd am o ng u nicellul ar fo rms r ep re se ntatives both

of animals and of plant s and to a detailed exami nation of so me
,

o f these we may n o w p r oce ed .

 C HA P T E R XI I .

UN I CE L L UL A R ANI MA LS ( Protozoa ) .

A . A m mb a .

( Th e P rt
o eus A n mai l ulc e .
)

r a
Gene l A cc o unt A ma ba is.a m i n ute o rga n i sm
fou nd in stagnant water in the sedi me nt at the botto m
,

ditche the rface of wate r pl an ta i n da m p ea r th , in

and o n s u

organic i nfusions of variou s ki nds —al mo st any whe re in sho rt in

s
-

, ,

the p re sence of moi sture orga nic matte r and othe r favo r able
, ,

conditio ns There are many s pecie s of A moeba so m e li vingi n

. ,

salt water othe r s in fre sh O ne of the l a rge st and co mm onest

,
.

f r e sh wate r for ms is A ma ba P roteus w h ich fo r ms the s ubject

-

,
*
of thi s accou nt .

A moeba occ urs in an active or mo tile state and a quiesce nt o r ,

ency s ted s tate NVh en active the bod y c o nsi st s ( Fig 8 4 ) of a

. .

m i nute naked m ass of prot o plas m which in the case of la rg e

sp ecimens is barel y vi s ible to the naked ey e half a m illi

m etre ( t i nch ) or le ss in le ngth
”
Thi s m ass c reeps o r r athe r
.
,

flows activel y about by the co nt in ual p rot r u sio n of lobes or p r oc

,

e sse s of its own sub st ance k now n as p seudop odia

,
These may .

be put fo rth from any pa rt of the su rface and agai n me r ged into
the general mass ; the body the r efore contin ually changes its
”
sh a pe ,
and he nce the na m e P rote u s .

Whe n the body is well ex t e nded the p rotopl asm is see n to

consi st of a clear periphe ral sub stance the ectop la sm and a cen , ,

tr al sub st ance the entop la sm filled with co ar se gran ule s w hi ch

, ,

give the body a highly char acte r istic gra nul a r appea rance so me
ti mes de sc r ibed as a gr ay colo r ”
Withi n the ectopl as m the
.

mo re fl uid e ntoplas m freel y flo ws as if c o nfi ne d in a t ube o r

,

O th r mmo n fo rm s are th e s ma ll er A rad iosa a nd A

e co . . eerrucosa . Th e
l arge A ( P elomyzra ) oillos a a nd A ( Dina moaba ) mirabilis are
.
no t i nf r q u nt
e e .

S ee L eidy Fre s h w a t er R h i zo po ds o f N o rth A m e ri ca

,
-

.
 THE PR OTE US A N I MA L C UL E . 15 9

th e
FI G t f l if
A mcs ba Pro eus , ro m e x 80 0 Th e a rr ow si i t i ti
n d c a e th e d rec o n of

d v u l ;
- .
.

p o op rt
la s mi c rr t
c u e n l
s ; n, n u c e u s ; c . tr t i v
r, c o n ac le u l ; f
ac f
o e . v,
f
oo -
ac o e

w v, a e
w t rv ac u o lh
e A s o w s t h e txt r
e u e r t pl
o f th e p o m
o B
asi .
u tl i s an o ne o

i ivi l f r i t t t th i ght f Fig

-

tr u r
. .

e u p w a rd en s a e r o
m l t h c r .

th e s a m e n d d u a o u n u e s a e

A h v t
a e s o p ped , m e v e rs ed , a n d t h e m ia n fl o w i s t w rd t h l f t
now o a s e e .
1 60 UN I CE L L UL A R A N I MA L S .

but the two s ub s ta n ce s are n o t sepa ra ted by any defi n ite

s ac,

bou nd ary li ne and p ass i mpe r ceptibl y i nto o ne anothe r The

-

,
.

xter l bo un d a r of the bod y is for m ed b y the outer m os t l i m it

e n a y
of the ectopl asm There is no me mbrane and the body is.
,

q uite n ked
a N e v erthele
.
ss the p r otopla s m ic m ass shows no
te ndency to mix w ith the su rrou ndi ng w ate r and pe r fectl y mam ,

t ai ns its i ntegrity ; it is an individual .

The for matio n of a p seudopod begi ns by the bu lgi ng out of

the ectopl as m to for m a rou nded pr o mi ne nce at so me poi nt o n ’

the s urface I nto its i nterior a sudde n gu sh of e ntoplas m then

.

t ake s pl ace and a stead y o ut wa r d s t r ea m e nsue s the e n toplas m ,

pu shi ng the e c topl as m before it and the sub s ta nce of the body ,

fl o wi ng i nto the p seudopod The whole su b s ta nce of the body

.

m ay thu s fl o w o nward i nto the p seudo pod which mea nwhile fo r ms ,

new p seudopod s and so the e ntire ani mal adva n ce s in the di rection
,

o f the fl o w ; or the p seud o pod afte r attai ni ng a ce r tai n siz e may

,

be withdr a wn i nto t he bod y by r e v er se ( ce nt r ipet al ) cu r re nts the ,

mai n bod y havi ng me anwhile flo w ed o nwar d in a nothe r directio n .

A s a rule the new p s e ud o podia are p ut fo r th n ea r o ne end

,

“
of the b o dy (he nce called ante r io r and the ge neral direc tio n

o f adv ance is therefo re fai rl y co nst a nt no t vague and i ndefi nite , ,

as is o fte n s t ated The directio n of fl o w fl uc tuate s ho w eve r

.
, ,

a bo ut a c ert ai n me an bei ng c o nti nu all y di verted thi s w ay or

,

th at by the for matio n of new ps e udopodi a Tho se w hich do not .

fo r m dire ctl y in the li ne of ma rch eithe r m e r ge little by little

with the adv anc i ng o ne s or are w i t hd r aw n b y r eve r sed cu r r e nt s
,

i nto the body I n the l atter case they ofte n le a ve sh rivelled

.

wart like re m nant s and a gr o u p of s i mil a r w a rt s is u suall y

-

,
“
fo und ne a r t he po ste r ior e nd of th e b od
y ( Fig 8 4 p) .
, .

Defi nite c h ange s in the ge neral d irec ti o n of adv ance are e ff ect ed
by the dive r sio n of the mai n curre nt i nto l ater al p s eudopodia .

A moeba feed s upo n min ute pl ants and a ni mal s or othe r o r

ganie particle s There is no m o uth and food matte r s are bodil y
.
,
-

i ngulfed (at no defi nite poi nt) by the p r otopl asm w hich close s
up beyo d the
n m *
The i ndige stible r e mai ns are pass ed out in
.

hi m d
T s ll ul r l i m nt t i n i f f req u nt urr n in m lls
o e o f ce a a e a o s o e oc c e ce so e ce

o f mult i ll ul r
ce w ll ain un i ell ul r
, as ni m l
e Ce ll x h ibi t i ngi t re
as c a , a a s . s e a

k n w n ph g y t ( t i ng ll ) nd th pr
o as a oc es ea i r f rr d t as p ha g y tosis
-
ce s a e oc es s s e e e o oc .

I t i bvi u l y n l y pr l ud t m t
s o o s o l l ul r di g t i n
a e e o ra c e
-
a es o .
 EN C Y S TE D S TA TE OF A M G B A '
.
1 61

an equ all y p r i mitive fashio n usually at so me point near the ,

“
poste r io r end B e side s so lid food stuf fs A mwba t ake s in a
.
-

ce rtai n qu antity of w ate r (alo ng with mi nute quantities of in o r

ga nie salt s dissolved in it) and it al so breathes by t aki ng in , ,

( m ai nl y by di ffu sio n) the f r ee oxyge n di ssolved in the wate r and

gi ving off c a rbo n di oxide .

S uch is A m azba in its active phase The q uies cen t or en .

cys ted st a te is e nte r ed upo n u nder co nditio ns n o t thoroughl

y
u nde r stood but p r obabl y of an u nfavo r able natu r e such as the
,
,

0
FI G .
-
A , A moeba d ivi i
d n gb y fi i n nu l u n tss o ( ft r L i d y )
, c e C Am s o s ee n a e e .
, oeb a
a ft er a. f l l i ti
ul m ea c o n s s n go f l r g d i t m ( it) ( A ft r L i dy ) L tt r
a a e a o c . e e . e e s as in
Fig 84 D, E y t d mmb n t i n i gf d m tt r ( ft r H w )
nc s e
A a, c o a n oo a e s a e o es
-
. . .

lac k of food d ry i ng up of po nd s and the like The pseudo

, , .

po di a a r e withd raw n m ove me nt cease s the body beco m e s , ,

sphe rical and su rrou nd s it self w ith a to ugh me mbr ane (cell w all ) -

( Fig 8 5 D ) The ani mal t akes no food and all of its acti v itie s
.
, .

are n earl su s pe n ded I t is like an a n i mal asleep or hibe rnati ng

y .

and in thi s s tate it m a lo ng re m a i n P rote c ted b

y its m em
y .

br ane it is able to r e si st de sicc atio n and upo n the ev apor ation of ,

“ ”
the su r r ou ndi ng w ater it may as a p article of d ust be tr ans , ,

po r ted by the w i nd s eve n to a great di sta nce Whe n agai n ,

.

pl aced u n der favo ra ble con ditio ns the protopla sm bu r st s its

e nvelope c r awl s fo rth fr o m it and reas su me s its active phase
, ,
.
 62 UN I CELL UL A R ANI MA L S .

S t r u ct ur e L y i ng in the e ntoplas m usuall y n ea r the pos

.
,
.

terio r extre m ity is a n ucleu s ( n Fig ,

havi ngthe fo rm of ,
.

a bi co nc ave di sk and largely m ade up of coa r se gr anul e s of

-

c h roma tin ( cf A m ceba is the r efo r e at o nce a si ngle

.

c ell an d a u nicellular o rga n i s m m orphologicall y e qu ivale nt to a ,

s i ngle ti ssue cell of a higher a n i m al or to the ge r m cell f r o m

-
-

which eve ry multicellul a r fo r m ar ise s The body of A mo eba is .

a one celled body

-
.

The p rotop la sm (cy toplas m) co nsi sts of a clea r basis and (in ,

the case of the e ntopl as m) of i nnume r able gr anules ext re m ely

diver se in for m and si z e and fr eque ntl y d ifferi ng in cha r ac te r ,

in diff ere nt i ndivid ual s O fte n they are in the fo r m of rh o m

.

hoid al c ry st alli ne bodie s ; in other cases they are r ou n d ed o r

i r regul ar Their pre c i se che m ica l co mpo sitio n is u nce rt ai n but
.
,

they are p r ob abl y co mplex orga nic co mpou nd s a p r oduct of ,

metaboli sm and s e r vi ng as re serve food matte r * -

.

Vacuoles The protopl as m ofte n co ntai ns rou nded vacuoles

.

o f which the three follo w i ng ki nd s may be di s ti ngu i sh ed

( )
a l V a ter v a eu oles -
Fig s 8 4 fill ed wi th wate r .
, ,

l yi ng in the e nt o pl as m and carried alo ng in its c ur re n ts .

b F d v a c uoles a l so l i ng in the e ntoplas m

( ) oo -

y co n ,
.

tai ni ng the so lid fo od matter s that have bee n i ngulfed Withi n -

the m dige stio n t ake s pl ace Whe n thi s p r o cess is co mplet ed

they approach the exterior—us uall y at so me po i nt nea r t he
.

po sterior end—the o uter wall bre ak s through and the inn utri ,

tiou s re mnants are ca st out the ectopl as m clo s i ng up the b r eac h ,

i mmedi atel y afte rward s Thu s A moeba has no m outh ali .

,

me nta ry c anal or a nus but the ge ne r al m ass of p r otop las m

, ,

pl ays the role of all th r ee .

( 0) Co ntra ctile va c uole (a v) U su ally s i ngle so meti mes

r
.
,

double ly i ng ne ar the poste rio r end and filled w ith liquid

.
, .
.

Thi s i s sharply di sti ngui shed fro m the other v ac uoles by its
r h yt h mical pulsati on expa ndi ng (dia stole) and co ntracti ng (sys
,

tole) at regul ar i nterv al s Du ri ng the dias tole the vac uole slowl y
.

fill s with liq uid w hich d r ai ns i nto it fro m the su r r ou nd ing pro to
pl asm A t the sys tok which is very sudde n thi s liquid is fo rci
.
, ,

bly expelled to the exterior through an ope ni ng th at breaks

In m sop i e sf A mmb t hec e snt p l m m y l
o nt in nn um r bl e
a e e o as a a so co a i e a .

gr in f nd t k n in f r m th x t ri r b ut th i s is n t th e case in A P r teus
a s o sa a e o e e e o , o . o .
 PH S Y IOL OG Y OF A M CE B A ’
. 1 63

th r ough the ectopl asm an d i mmediatel y afte r wards di sa ppears , .

The co nt r actile vacuole is al m ost ce rtai nly to be r egar ded as a

s i mple kind of excreto ry a ppa r atu s the w ate r which collects in
,

it co ntaini ng in solutio n v a rious products of destructive metabo

lis m whi ch are thu s passed ou t of the bod y * .

R epr odu cti on H oweve r abu n dant the food suppl y A m oeba
. -

n eve r gr ow s be o n d a certain m aximu m limit A fte r thi li m it

y s .

has M e atta ed the a m

°

n in ni a l soo ne r o r later divide s by fis sion “

i nto tw o smalle r A mazon? ( Fig 8 5 A ) Thus the exi ste nce of .

, .

a n indi v idu al A m oe ba is n o r mall y ter m i nated not by death but , ,

by resolutio n int o two n ew i ndivid ual s Thi s proce ss is the .

s i mple s t po s sible fo r m of aga m oge n e s i s and A moeba is n o t k n ow n ,

to m ultiply in any other way xf The fi ssio n of A moeba is a

p r ocess esse n ti all y of the sa m e n atu re as the di vi sio n of ordi nary
tissue cells a di vi sio n of the nucleu s p r ecedi ng th at of the
-

cy toplas m Whethe r the divi sio n of the n ucle us is of the i n di

.

r ect type passe s th rough the phe no m e na of k ary oki ne si s) is

n o t k n ow n by d ir ect obse r vatio n b ut there is som e r e ason to be ,

lieve th at it is so I n any case the succe ssive fi ssio ns of A moeba

.

a r e di r ectl y co m parable with the succe ssive cle av age s of the egg
o f a m etaz oon ( p The proge ny of the A m a ba how ever
.
, ,

sepa ra te and fo r m i ndepe nde nt i ndivid uals while thos e of the egg ,

cell re mai n inti matel y associ ated to fo rm a si ngle m ulti cellular -

i ndivid ual M o r ph o logi cally the r efo r e a m etaz oon is co m p a rable

.
, ,

n o t w ith a s in gle A m oeba but with a multitude of A m cebce , .

Ph y s i olog y The po ssible s i m plicit

.
y of a n i m al str u c tu r e is

well sho wn in A mceba which is m orphologic all y an an i mal re ,

duced to its lowe s t te r m s I ts ph y siologic al O pe r atio n s are cor

.

r es po nd ingl y p r i mitive and r udi me ntary ; and by an analy si s of

the m we m ay discove r wh at is e ss e ntial and fu ndame ntal in the
phy siology of ani mal s in ge ne ral A su r vey of the v arious activ .

itie s of A meaba sho ws that the s e may all be r educed to a fewf u n da

men ta l p lzys io log i ea l p rop erties of the p r otopl asm ! as follo ws ,

It m y b r a ll d th t th vi ty f th n ph ri di um in th rth w rm i
e eca e a e ca o e e e ea o s

i nt r
a ce-
ll ul r l ik v u l ( p
a , e a ac o e .

»
1 I t h b n
as s r t d th
ee t A m az b
as e n j ugt nd
e l th
a t i t m u lt i p l i
a co b y a es a a so a es

e nd gn u d i i i n
o e o s b ut th vi d n n b th th p i nt i i n n l u iv
v s o e e e ce o o e se o s s co c s e .

t I t i h rdl y n essa ry t r m rk th t in mm n w i th ll E ngl i h p k

s a ec o e a a co o a s -
s ea

ingbi l g o oi t we i nd b t d t F t r f th fi r t mpr h n iv l b r t i n
s s a re e e o os e or e s co e e s e e a o a o

o f th e fund ment l p h y i l gi l p r p rt i
a a x hi bi t d by A mwb
s o o ca o e es as e e a .
1 64 UN I CEL L UL A R A N I MA L S .

( )
1 Con tractility , by m ea n s of which m otion is eff ect ed .

This appear s most clea rl y when the an i mal is stim ulated by a

sudde n jar o r by an elect r ic shock which causes the bod y to
, ,

cont r act into a b all Th is prope r ty p r ecisel y like the co nt r action

.
,

of a muscle (p is the re sult of a m ol ecular r ea rra nge me nt

. ,

accomp ani ed by che m ical cha nge s which causes a cha nge of ,

fo rm in the m ass withou t alte r in g its bulk The action of the .

cont ractile v acuole is due to the co nt r actility of the su rr ou nding

protoplasm ; and in l ike mann e r th e cu rre nts wh ich cause the
p rotru sion and w ithd ra wal of pseudopods and so th e loco motio n ,

of the ani mal as a w hole are p r oduced by l oca liz ed co nt ractions

,

of the pe r iphe ral lay er of p r otoplas m which drive onwards the

more fl uid cent ral part s .

I rrita bility ( inc lud ingCo ordin a tion ) o r the powe r to

( )
2 -

be aff ected by and to r e spo nd to cha nge s o r st imuli act i ng

, ,

upon o r withi n the p r otoplas m The c h ange of shape foll owing .

the applic atio n of an elect r ic shock is ac tu ally effec ted by co n

t r actility but the power to be affected by the s hock and to a r o use
,

cont r actility is i rr itability To thi s p r ope rt y the ani mal o wes it

,
.
s

powe r of pe r for mi ng ad aptive ac tio ns in r e spo ns e to changes in

the e nviro nment and al so its power to c o ordin a te the va rious
,
-

actions of its own body To illu st rate : I t is a r e mark able fact

.

th at A ma ba is able to di sc ri mi nate bet wee n nutritiou s an d i nnu

’

t ritio us matte r s ing ulfi ngthe for m e r b ut rejecti ng the l a tte r

, , .

Ph ysiologic ally this di scri m i natio n is a d if erence of resp ons e to

( lif erent s timu li—he nce a phe no m e no n of irritabilit A gai n
y .
,

the v ario us actio ns (m ove me nt s etc ) of A ma ba de s pite thei r ,

.
,

app are ntl vagu e c har ac te r a re co ord ina ted to fo rm a defi ni te

y ,

whole ; and c o ordi natio n may be rega rded as a phe no me no n o f

-

ir ritability ch ange s in o ne part se rvi ng as sti m uli to othe r pa rt s

,

an d bei ng brought i nto o rderl

y rel atio n with the m The p r ope rty .

of irrit ability lie s at the b as e of all ne r v o u s ac ti v ity in highe r

fo rms (of p 6 7 ) and is co nce r ned in ma ny othe r actio ns
. .
.

(3 ) illet abolis m the m ost fu nd ame nta l of all vi tal actio ns

, ,

si nce it lie s at the root of all is the powe r of was te and re pa i r ,

the dest r uctive che mic al ch ange s in p ro to pl as m ( ka ta bolism)

w he r eb y e n ergy is set f r ee and the co nst r ucti ve ac tio ns a na bo
( ,
v

lism) th r ough which new pr o topl as m is b uilt and pote ntial

e nergy is stored (cf p The r e is eve ry reaso n to beli eve
. .
 PH YS I OL O G ICA L P R OPE R TI E S OF A M GJ
BA 1 65

th at the metabolic phe n o m e na of A maeba are bro adly speaki ng

'

, ,

si mila r to tho se of highe r an i mals The k atabolic ch anges are in

.
.

the long run proce sses of oxid atio n and although thei r p r oduct s
,

h ave no t yet bee n defi n itel y asce r tai ned in A m oeba there can be ,

no doubt that they co nsis t m ai nl y of ca r bo n dioxi de water an d , ,

so me fo rm of n it roge nou s matter ( ure a o r a r el ated sub stance) .

Mo st of th ese w as te m atter s are believed to be passed out (se

cretion ,
ac cretion b
) y m ea ns of the co ntr actile v a cuole but prob
,

abl y carbo n dioxide le a ve s the bod y by dif fu sio n th r ough the

'

ge ne ral s u r face (resp ira tio n in pa rt) .

The m ate ri al s fo r the co nst ructive proce ss ( a na bolism) are

de rived fr o m o rganic food matter s bodie s or frag me nt s of pl ants
- -

an d anim al s take n as food in the p r oc e ss of a limen ta tion and

,

absorp tion f ro m the w ater a nd the i n or gan ic sal ts di ssolved in

it and fro m the free oxy ge n that e nte rs by diff usion thro ugh
,

the ge ner al su r fac e ( resp ira tio n in pa r t) P roteid matter is an .

indi spe nsable co nstitue nt of the food and A moeba is the refore ,

an an im a l .

A li me ntatio n abso r pti o n sec r etio n d ige stio n and circul a

, , , ,

tio n all of whi ch are o nly the p r elude to meta boli s m but which
, ,

in the h igher a ni mal s are assig ned to diffe re nt organs ti ssue s , ,

and cell s a re he re pe rfor med by o ne an d the sam e c ell

,
The .

capt ure o f solid food here r equire s its e ntra nce i nto the cell ;
and the fact that proteid s ca nno t be ab sorbed by diffus io n n ec es

sita tes i nt racellul a r dige s ti o n w hich in t ur n n ece ss it ate s cell ular

defaeca tio n I t w ill be ob s e rved th at wh ile there is no loc ali z ed

.

o r pe r man e n t m o u th or a nus the w hole su rface of the c ell is

,

p o te nti ally m outh o r anu s I n sho rt the pr o topl asm here ex

.
,

hibits no t th e ph y siologic al di v i s i o n of l abor but its ab se nce ,

.

e rod u ctio n o gica ll y there is in th e

( )
4 G row th a n d R p L .

case of A ma ba no go o d gr o un d fo r a di sti nctio n betwee n the se

processe s an d metaboli sm ; for r eprod uctio n is dire ctly or i n di
rec tl a n effect of gro w th a n d gro w th is si mpl y a n exce ss of
y ,

anabolis m ove r kataboli s m P racticall y however the di sti no

.
, ,

tio n is n e ce ssa ry ; f o r th e te nde ncy of livi ng thi ngs to run in

cycle s of g r owth and r ep r oductio n is one of the ir mo st obviou s
and c ha r ac te ri stic featu res .

He r e as in all protopl as mic s tructu r es g r owth take s pl ace

, ,

th r oughout the mass by in tus sus cep tio n (p

,
n o t by the ad .
1 66 . UN I CELL UL A R A N I MA L S .

ditions of su pe r ficial l ay e r s as in the case with g r owt h by accre ,

tion ( i n orga n ic bodie s e

g cr y s tal s
) U n der f a vo r,
able co n di
. .
,
.

tion of nut ri tio n thi s p r ocess exceed s the de str uctive p ro cess so
th at the body i ncrease s in s iz e up to a li mit at which fi ss io n .
,

takes pl ace What deter mi n es th is lim it is u nk no wn but the

.
,

ca u se is perhap s in so me way conn ected with the geo met r ical

p ri nciple th at the vol ume of the cell i ncrease s as the cube of its
di amete r whereas the su rfac e by which it abso rb s nut ri me nt
, , ,

and othe rw i se c o m e s i nto r e latio n with the outside w o rld in ,

cre as es onl y as the sq ua re of the diam eter N o gr eat i nc rease .

in si z e therefo re is possible without de s t r oy i ng the n or mal equi

, ,

libriu m of the cell and he nce the pe riodic r eduction of si z e by

divi sion This pri nciple is howeve r too ge ne ral to be of much
.
, ,

v al ue Differe nt s pe c ie s of A mab a di ff e r in si z e li mit an d the

.
-

i mm ediate cause lie s in so me s ubtle r elation betwee n

and e nviro nm e n t th at c annot at pre se n t be m ade out I .

k no wn w hether o r not the A m oeba ever dies of old age .

These fund ame ntal phy siological prope r ti es of p r oto

pl asm lie at the bas i s of all ph ysiology and will be foun d ap ,

plic able to all for ms of life whethe r vegetal o r ani mal .

R elate d Forms represent a t i ve o f a very ex t ensive cl ass 0

. A moeba is a

Pro t o z o a k no wn as R h izop od a a ll c h a rac t e riz ed b y t h e po wer to f o r m ,

pseud po dia a nd ag
o reei ngwi th A mo ba in ma ny th er res pec ts O ne 0
,
e o .

t h e c o mmo nes t f res h wa t er fo r ms is th e g -

enus A roella ( Fig 8 6 wh i c h .
,

e v en in t h e ac t i v e ph as e is s urr o unded b y a bro wn h o rny memb ra ne

per fo ra t ed by a l a rg e r o unded o pe ni ngth ro ug h wh ic h pseudo

po di a a re pro truded Dzj fl ugia ( Fig 86 B ) a l so a co mmo n f res h wate
. .
, ,
-

fo r m bui l ds abo ut i t sel f a bea ut i ful v ase s h aped o r ret o rt s h a ped s h el

,
- -

co mposed o f s a nd
gra i ns o r ev en in so me cases o f di a t o m s h el ls I n
-

, , ,
-
.

A cti nop h r ys o r the , s un a ni ma l c ul e

"
( Fig 8 6 A ) t he pseudo podi a a re
-
”
.
, ,

s t i ff needl e s h a ped
process es radi a t i ngin ev ery direc t i o n
-
.

A mo ngth e ma ri ne f r ms t wo g ro ups ( o rders ) are o f

o

a nd i mpo r t a nc e v iz th e Fo r a mi n if era wh ic h s ec ret e a c al ca reo us

,
.
,

per forat ed by numero us po res a nd t he R ad iola ria wh i c h h a v e a s i l i , ,

s h ell . Many o f th ese fo r ms fl oa t at th e s urf ac e o f th e wa ter an d ,

c as t off s h ell s h a v e in f o r mer t i mes acc umul a ted a t t h e bo t to m in

-

e no r mo us q ua n t i t i es as to f r m beds o

w h i l e th e rema i n o f R adi o l aria h ave

s

f o r ma t i o n o f si l iceo us roc ks .
 FR E S H WA TE R R HI ZOP ODS .
1 67

FI G 86 G
. .
—r
o up o f co mm o n fr h w t r R h i z p d ( ft r L i dy ) A A tin p h y
es -
a e o o s a e e .
, c o r s
so l. th e
“
s un a n
-
im a c l u l fi l l d w i th v u l
"
e. e dac t i n i ngth r f d b d i
o es a n c on a ee oo -
o es
f urth i j u t b i ngi ngulf d Th u l u i n t
o s s e e . e n c e s s o s ee n .

b u i lt f d g r i n nd p ud p d i f x
o sa n -
a s a se o o a ar e

i nd ivi du l h w i gth p r t p l mi b d y u
a s o n e o o as c o s s

p end ed w i th i n th e s h ll ; v r l v u l
e se e a ac h wn b ut n nu l u
o es a re s o , o c e s.
‘
 C HA P T E R XI I I .

UN I CE LL ULA R A N I MA LS ( PR OTOZO A ) ( Continued) .

B . I n f us o ri a .

( P a ra me c ium , V rticell
o a , etc .
)

I N FUSO R I A are u nicellul ar ani mals fo und like A mwba

mi nute
in st agnan t water o r in orga nic in fusio ns ( see p 2 01 ) (he nce .

“
I nfusoria I n the leadi ng fea tu res of their orga niz atio n
they are clo sely si mil a r to A moeba and its allies f ro m which ,

they di ff e r ho weve r in havi ng a much highe r degr ee of di ffer

, ,

entiatio n in mo v i ng by m ea ns of c ilia i ns tead of pseudopodia

, ,

a nd in sho w i ng the fi r s t i ndica tio n of ga moge n esis ( am phi m ixis ) .

P a ra mozc ium (the slippe r an i malcule ) is an ac tively free

- .

s wi mmi ng for m ofte n fo und in m ulti t ude s in h ay infus io n o r -

water contai ni ng the de c o m po si ng re mai ns of N itella and othe r

w ater pla nt s
-
Vortic ella ( the
. bell a nimalc ule is co mmo nl y
-

att ached by a sle nder s talk to du c k weed ( L emna ) and o the r -

w ate r pl ants o r to other sub me rged objects ; at othe r ti mes it

-

,
.

bre ak s loose f r o m the s ta lk and swi ms fo r a while ac ti vely about .

The two fo rms a r e constructed u po n e sse ntially th e sa me plan ,

b ut Vo rticella sho w s in so me r e spects a much highe r degr ee of

differe nti atio n
Par ame c i um —The slippe r shaped bod y ( Fig 8 7) is cove r ed
.

-
.

with cilia by m ea ns of w hich the a ni mal r apidl y swi ms about .

Morphologi c ally the body is a si ngle cell h av ing the sa me g en ,

eral com po sitio n as in A moeba but posses si ng in additio n a deli

,
”
c ate su r r ou ndi ng me mb ra ne cuticle ) o r cell wall The -
.

differe nti ation of the p rotoplas m into ectoplas m and e ntoplas m

is ve ry sh a rpl y mar ked and the fo rm e r co nta i ns n u merous
,

peculi ar r od l ike bodie s (tricho cys ts) f ro m which lo ng

-

be thrown out Thei r fu nctio n is p r o ba bl y th a t of

.

prote c tion A s in A ma ba the p rotopla s m co n tai ns wa ter v ac u

.
-

o les and
f ood v acuoles (f m) ( both of which ar e ca r ried
-
 THE S L I PP E R A N I MA L O UL E — . 1 69

B
FI G . 87 .
—P ara mazci um ca udat um
A f r m t h l f t i d h w i.
n g t h,
l oS p t ; B
e e s e, s o e a na o ,

fr m th v n t l i d h w in gt h v t ibul n f ; r w d t h b dy
.

ace ar o s i n si e e o 1 11
o e e ra s e, s o e es e e

di t th
ca e di r t i n f p r t
e p l m i ec u rr o n t th
o u t i
o o d t
ash d i c t ic
n f e s. o se o s e e rec o o

w t r u rr n t
a e -
u d by th
c i li
e s ca se e c a
f v r v
.

a n ,n la p t; a tr t il v
s o u l ; f
c v, co n
.
d acu l : ew w
ac t
o es cfl
u l v, oo -
ac o es . v, a e a o es

m u th : m
o m u l u : mi mac, i r n u l
a c ro n u ; c e p h
s g u :c,
t b c o
T h
c e s as, oeso a s v , VO S l l e e

a n t ri r n d i di r
e o t d up w d
e s ec e ar s.
170 UN I CELL UL A R A N I MA L S .

about cu r ren t s in the e ntopl asm) and two ve ry large con tmac
by ,
t

t ile v a c u oles ( )
o v o cc up y ing a co n s ta
.
n t po s itio n o n e near eithe r ,

en d of the body The nucleus (as in I nfuso ri a ge ne rall y) is

.

d ifferen tiated into two di stin ct par ts v iz a la rge oval macro , .

,

n u cleu s ( ma c ) and a much smalle r s phe rica l micron ucleus (m ic )

do ble in s o m e specie s) l y i ng clo s e beside it
( u .

U nlike A moeba I a ra mwc ium posse sses a d istin ct mouth (m) ,

’

an d ( es op ha g u s ( a ) which ope n to the exte ri o r th r o ugh a n obli q ue

funn el sh aped depress io n k n o wn as the v es tibule ( v ) situat ed at

-

o ne side of the bod y Mi nute floati ng food pa rticles are d rawn.

-

by the cilia i nto the m outh and accu m ul a te in a cilia ry vo r tex at

the bottom of the ( e sophagu s F rom tim e to ti m e a bo lus o r .

food mass is then ce passe d bodily i nto the sub s ta nce of the en
-

to plasm form i ng a food vac uole with in which dig estio n takes
,
-

pl ace The i ndige stible re mnants ar e fi nally passed out no t

.

through a pe rm a ne nt ope ni ng or a nu s but by b reaking th r oug h ,

the protopl as m at a defi nite po int he nce kn ow n as the ( mal ,

sp ot ,
w hich is s itu ated nea r the hin de r end ( Fig The .

con tra ctile v acuo les of P a ra meec i u m are e s pec ially favor able f o r

st ud showi ng at the m o m e nt of co n t r actio n or just b e fo r e it

y , , ,

a p ro nou nced star shape with lo ng ca nal s runningout into th e -

protopl asm Through these liquid is supposed to fl ow into the

.

vacuole .

L ike A m oeba P a ra moec iu m occ urs both in an acti ve and in

,

an en c s ted state
y I n the for me r sta te it multiplie s by t ra ns
.

ve rse fission divi sion of both mac ron ucleus and m icro nucleuc
,

precedi ng or accompa ny i ng th at of the protoplas m ic bod y ( Fig .

88 A )
, U nde r fav o rable c o nditio ns di v i si o n may take pl ace o nce
.

in t we nty four ho urs or eve n ofte ner

-
Thi s p r ocess which is a
, .
,

ty pi c al case of agamoge ne si s may be r epe ated agai n and again , ,

thro ughout a long pe riod B ut it appe a rs fro m the celeb r ated .

re sear ches of Maupas that eve n u nder the mos t favo ra ble co n
ditio ns of food and te m pe ra ture the p r oce ss has a l imit ( in the
c ase of S tylo n ichia a for m rel ated to P a ra meec ium thi s l i mit , ,

is reached afte r abo ut 3 00 successive fi ss io ns A s thi s l imit i

) .
s

approached the an i mal s beco m e dwa rfed s how v a rio us s igns of ,

degeneracy and fi nall y become i ncapable of taki ng food The

, .

r ace gro ws old and dies .

I n nat ure ho w e v er thi s li m it is p rob abl seldo m i f eve r

,
y ,
 CON JUG A TI ON OF P A R A M OZOI UM ’

171

re ache d ,
an d the dege n e rative te nde ncy see ms to be checked
by
kn o wn as co nj uga ti on I n thi s proce ss tw o in dividual s .

mselv es side by s ide pa rti all fuse t o gethe r and remai

y n , ,

thus u ni ted fo r seve ral hou rs ( Figs 8 8 B D uri ng thi s .

, ,

unio n an exch an ge of n ucle a r m aterial is e ffected after which

,

sep a rate both m a cron ucleus and m icron ucleus now

'

FI G . 88 ,
—A . F i i
s s on o f P a ra mazci um . ( Fro m a p r e pa r ti
a on by G N . . C lk i ns )
a . mac ,

mae ro n uc le us ; mic m i c ro n uc l eus ; m m o uth

, , .

B .Fi rs t s ta ge o f c o n j uga tio n Th e a n i m a ls a re

th i r v ntr l u. a pp li ed by e e a s r

f ; th n ly
a c es h g th u f i th l rg m t f t h m i
e o c an e u l i s ar s e en a e en o e c ro n c e .

C C j ug t i
. on t th m m t f x h
a on a g f th m i r nu l i ( l
e o m g ifi d)
en o e c an e o e c o c e es s a n e .

Th m r u l i d g r t i g E h i d ivi d u l t i t w m i r u l i
e ac on c e are e e ne a n . ac n a c o n a ns o c on c e

( n w p i nd l
o h p d ) n f w h i h r m i i t h b d y w h i l th th r r
s e s
-
a e , o e o c e a ns n e o , e e o e c o s s es

v r t f u w i th th fix d m i r u l u f t h th r i d ivi du l ( A ft r M u p )
o e o se e e c on c e s o e o e n a e a as .

co nsis ti ng of mi xed materi al de r i v ed eq uall y fr o m both i ndivid

o n of the tw o a ni m al s is q ui c kly follo w ed by

In h i ndi v idua l th e mac ro nuc l eus b reak s up a nd dis appears Th e

eac .

mic ro nuc l eus o f eac h di v i des t wi ce and o f t he fo ur bo di es th us pro duced ,

th ree disa ppear: Th e four t h di v i des agai n i nt o two o ne o f wh i c h remai ns ,

in th e bod y wh i l e t he o th er c ros es o v er a nd f uses wi th o ne o f th e mi cro

“

,
s
nuc ei o f t h e o t h er i ndi v i dua l a f te r wh i c h t h e a ni ma l s sepa r a t e Th is
l ,
.

pr o c ess be i n r
g pec i r o ca l ea c h i nd ivi d ua l no w co nt a,
i ns a mi c r o nuc l eus con
1 72
t ri l f m h i d i v i du l T hi s mi c ro
q l m o f m a te a ro e ac n a
t ai ni ngan e ua
u n
.

a c
t i i v ri f r b d i t w o f wh i c h be
n ucl us no w
e di v i d e s w c e a n d g es se to o u o es o ,

r l i d t w m i c r o nuc l ei Fi ss i o n n e x t occ u rs a n d is th ere

c o me mac o
,
nu c e a n o .

l m nner
a f t er co nt i nue d i n t h e u s u a a .

a alogous to the u n io n of the ge r m

This is a p r oce ss clea rly n
I h w eve r be ca lled ga m o
cell s of highe r a n i m al s . t ca n n ot o
, ,

it is only co mpa rable with one of

the elements of g amo e o' n e s i s . I n the m eta z o o n 8 .

FI G —G r oup o f Vo rt icella . in
'
v ria o us tt i t ud
a e s. a tt h
ac ed to
t r pl nt
.

wa e -
a .

cell s (fertili zation) is follo w ed by a l o ng se ries

c le avage of the ovu m) the re lti g cel l bei n g associated t
( s
,
u n s

fo r m one new i ndividu al I n the I n f uso ria te mpo ra r y fu sio

.

o j g tio
( c n u a n) is like ws i e followed b y a s e r ies of cell div isio n s -

but the cell s beco m e en ti rel y sep r te each bei g

a a n a n i
i div du
n a
,

r t
Vo icell a a gree s w ith I a ra m wc i u m’
in ge n e ral s t ruct u r e bl ,

difi ers in m ny n a i te r e sti n g det a il s m o s t of which a,

r e the e x pref
 THE B EL L A N I MA L C UL E
-
'
.
17 3

FI G . I
(X gl
-
h
A eas in
d f Vorteicd la h i ghl y m o g n i fi d x nt t
, i l xi f a e . c , co rac e a s o

t lk ; ut i l ;
s a c, c nt t il v u l ; d d i k ;
c e c v, co
.
t pl m ; n t pl m ;
ra c e ac o e , s cc, ec o as e , en o as

D pi t m ; f
6 , e s o f d ve u l ; m
v, m u th
oo ; m - m ac n u l
o e u ; m
,
i m i ou ac , ac ro c e s c, c ro n

i v ib v x p in t t
.

l u ;
c e s p h g ; p p
u
as. (e so r t m a; st u l ; ,
uw we t r s o u el ; v, es e , a e -
ac o es , o a .

whi ch p istm n e n d p ri t m m e a t t n n d f th v t i b u l
e s o e ee a o e e o e es e.
7 174 UN I CELL ULA R ANI MA L S .

s io n of highe r diffe r e n tiatio n The body is pea r shaped o r co n -

cal attached at its ape x by a long slen de r stalk

,
The lat h .

consists of a sle nde r cont ractile axia l fila ment by mean s ,

1

which the sta lk may be th r own i n to a sp iral an d the body d ra w

dow n and an elastic shea th ( co ntin uou s with the ge ne ral cuticl
,

by w hich the stalk is st r aighte ned ( Fig The c ilia are cm .

fi ned to a thicke ned rim the p eris to me ( p ) s u rr ou ndi ng tl

, ,

b ase of the co ne whi c h may be te rmed the dis k A t o ne sit

,
.

the dis k is raised for mi ng a projecti ng angle covered with ci li

,
:

a nd k n ow n as the euis tomc ( 3p ) A t the sa me s ide the peristo n .

dips dow nwa rd s lea ving a s pac e bet wee n it and the episto m
,
.

Thi s space is t he ves tibule (v ) and i nto it the mouth ope ns 1 , .

it likewise is sit uated an a na l sp o t like th a t of P a ra mceci mz

The cilia produce a powerf ul vo r te x c e n te ri ng in the m outh l ,

me ans of whic h f ood is secu red q

The mac ro nu c leu s ( mac) .

lo ng sle nde r and ho rse shoe shaped ; th e small s phe ri cal mien
, ,
-

nucle us ( m ic) lie s nea r its middle po r tio n The re is usua l .

but o ne co nt ra c t ile v a cuole .

Vo rticella m ultiplie s by fission divisio n of the pro tO plas ,

bei ng acco mpanied by th at of the mac ro nuc

c leus ( Fi
g The. pl an e of fi ss io n is ve rtica l
( thu s

the peri st o me i nto h alve s) but exte nd s o nly th rough t

,

body le a vi ng the sta lk u n divided A t the clo se of the

, .

therefore the stalk bea rs two head s O ne of these

, .

att ached to the or iginal stalk w hile the o ther fold s in ,

s to me acqui res a seco n d be lt of c ili a a r ou nd its

,

b reak s loose fro m the ste m and s wi ms active ] ,

”
called motile fo rm Ulti matel y it attache s
.

lose s its second bel t of cili a develops a s talk and ass um es , ,

o rdi na r
y for m B y this p r oce ss dis pe rsa l of the s pecies is
.

s ured .U nde r u nfa v o rable c o nditi o ns s i mil a r m otile for ms

ofte n pro duced without p revi o us
inga s e co nd belt o f cili a droppi ng ,

seek mo re fav o ra ble s urrou ndi ngs Vo rt .

cy sted l o si ng its peris to me a nd m o uth

, ,

fo r m acqui ri ng a thick me mb ra ne a nd
, ,

this state it is sa id so meti me s to m ult

breaki ng up i nto
bodie s (sp ores)
 CON JUG A TI ON OF VOR TI CE L L A .
175

ln uc leus These a re fi nally libe rated by the bu rsti ng of the

.

me mb r a ne ac q ui re a cili ated belt and after swi mm i ng

,
for a ,
rtime b
eco me attached lose the ciliated belt and develop a stalk , ,

and pe ri s to m e .

Vorticella goe s th rough a p r ocess of co njugation which h

as
so me i nte r e st ing pe culia ri tie s ( )
1 C o n jugati o n a lwa y
s t a
ke s
.

place betwee n a la rge attached in dividu al ( t he a morog a mete and

)
a mu c h smalle r free swi mmi ng individual ( the m icro a mete
g )
-

. ma a

FI G . 91 .
—
Fi i s s o n a n d co n u j ga t i o n Vo r icella
of A t . . E a rl y t s a ge of fi i s s o n, s h wi n g
o

ii
di v s o n o f m c ro n uc leus i ( m ic ) a nd m a c o n uc r l e us ( mac ) ; pe ri ts o me . ( A ft er
B fl t sc h li ) .

B. , D C S uc c e ss e s a g
.
es iv t of fi i ss on ; in B a nd C t h e n uc l i h v m p l t ly di
e a e co e e
v id ed a n d s s o n o f th e fi i
ll b d y i i p r g ; ce -
o s n o r es s c v, c o n
. tr t i l v u l I n
ac e ac o es .

D fi i n i m p l t ; t h r i ght h nd i d ivi d u l h
ss o s co e e e -
a n a as acq ui d reb lt f l a e o oc o

m t r i li o ot d i r dy t
c a a w im w y1; an s ea o s a a .

E C n j ug t i n f
. o fix d m
a og m t ( m ) W i th fr
o a e a c ro a e e a a ee s w
-
i mm i n gm i c ro ga m et e
( mi) ; p p r i t m p
, pi t m
e ( A ft r G
s o if )
e, e , e s o e . e rec .

(Fig . E ) The microga mete is fo rmed ei ther by the unequal

.

fissio n of an o rdi na r y i ndivid ual th e s maller m oiet y bei ng set ,

f ree o r by two or m ore r apidly succeedi ng fi ssions of an ordi nary

,

i ndi v id ual o j gat o n is pe rm an e nt an d co m plete th e

( )
2 C i
'

n u .
,

bod y of the m ic rogame te be i ngw holl y absorbed into th at of th e

176 UN I CE L L UL A R ANI MA L S .

m ac rogam ete Withi n the bod y of the lat te r afte r c o mplica ted
.
,

c h ange s the n uclei fu s e togethe r and this is followed by fissio n

, , .

The analogy of co njugatio n to the fe rt iliz ati o n of the egg is he re

c o mplete The co nj ugati ng cells show a se xual difi erentiatio n
.
,

o ne bei ng like the ovu m la r ge and fixed the othe r lik e the , ,

s pe rmato z oon s mall and m otil e ,

.

A s in Pa ra ma ci um the mac ronuc l ei ent irel y di sappea r f us i on ta k es ,

l ac e be t wee n deri v a t i v es o f t h e mi c ro n uc l ei a nd f ro m t h e res ul ti ngbody

p ,

bo th mac ro nucl ei and mi cro nu l ei a re deri v ed c .

E ug
lena and Ot h er Si mpl er I nfu sor i a
fo rms l ik e . B esides
P a ra mccci um and Vorticella which bea r nume r o us cilia the re ,

a re ma ny I nf us o r i a which possess o nl y o ne la rge lash o r fl a g elé wm .

O f the se E ug le na which is so m eti mes fo u nd in stagna nt wa te r

, ,

se w age po lluted pool s etc is o ne of the m ost inte r estin g inas

-

,
.
, ,

much as it co nta i n s chlo r oph yll possesse s an ey e spot of red ,

-

pigme nt and u nde r ce rt ai n co nditio ns e xhibits ammbif orm

,

m ove me nts .

Compound or
“
Colonial For u m I n a nu mbe r of fo rms
”
,
.

c losel y r elated to Vorticella the i ndivid ual s z ooid s fo rmed

,

by fi ss io n do no t i mm ediatel y separate but r e mai n fo r a ti me ,

“ ”
u nited to fo r m a colo ny w hi c h may co nt a i n hu nd reds of
z ooid s Zodt/ mm n imz a c o mmo n s pecie s thu s fo r ms a beautiful
.
, ,

tree like o rgani sm c o ns i s t i ng of a si ngle ce nt ral s t alk with nu

-

m ero us b r anc hi ng offsh o ot s fro m its su m mit each t w ig te rmin at ,

ingin a z o oid The e ntire sy s te m o f bra nch es is trave rsed by a

.

c o nti nuou s co n t ractile a xi s Ca l c/zes i u m is si mila r but the axis

.
'

is i nter ru pted at the begi nni ng of ea c h b ranch I n E p is kc/ lie .

the e nti re axi s is no n co nt ractile -

S uch colo ni al fo r ms are of high i n tere st as i ndicati ng the

m anner in w hich true m ulti cell ul ar for ms may have a rise n .

Fro m the l atte r ho wever they differ no t o nly in the fact that
, ,

the as soc iati o n of the cell s is no t pe r manent but in the abse nce ,

o f an divi s io n of l a bor a mo ng the u nits

y .

Phy si ol og y Most I nfu so ri a are true ani mal s ag r eei ng wi th

.

A moeba in the e sse ntial featu r e s of their nutritio n and having ,

t he po w er to dige s t no t o nl p r oteid s but al so carboh d r ate s an d

y y ,

fats . P a ra maaci um and Vorticella arc herbivor o us fo rms ,

feedi ng upo n mi nute pla nts and e spe c ially upo n the bac te ri a ,
 CHL OR OP H Y L L CON TA I N I N G I N F US OR I A . 1 77

O th e r for ms are o mn ivorou s S tea ler B ursa r ia ) feedi n g , ,

both o n vegetable an d o n a ni mal food O ther s still are car .

niv o ro us and lead a predator y life ofte n attacki ng herbivo r ou s

,

for ms m uch lar ge r th an the m selve s preci sel y as is the c ase with
,

ca rn ivo r es a mo ng the mammalia Thus the u ni cell ular world

.

r eproduces in m i niatu r e the esse n ti al biological relations of

h ighe r ty pe s .

I t is a r e mark able fact that so m e specie s of I nfusori a

P a ra mwci um bursa ria Vorticella mwidia) co nt ai n n u merous
’

chl o r oph yll bodie s e mbedd ed in the e ntopl asm Much di sc us

-

i
.

sion has a rise n as to whether these bo dies are to be regarded as

an i ntegr al pa r t of the a ni mal i e diffe r e nti ated out of its o wn
,
. .
,

protoplas m o r as mi nute pla nt s livi ng sy mbiotic ally (ize as

, .

m ess mate s) w ithi n the an i ma l

-
I n the for m e r c ase ( which is
.

the m ost probable) the a n i mal would to a certai n exte nt be

nou r i shed afte r the fas hi o n of a gree n pl a nt ( cf p . .

I t wi ll no w be clear to any o ne who h as carefull y co nsidered

the phe no m e na desc r ibed in the foregoi ng page s th at the uni
“ ”
c ellul a r a ni mals are orga ni sms by right and no t merel y by ,

courtesy I n so m e of the I nfu so ri a for ex ample differe nti a

.
, ,

tio n withi n the si ngle cell may go so far as to give ri se to pri mi

tive se nse o rgans ( as in the case of the eye spot of E ug
-
lena ) ; a -

r udim e nta ry oesophagu s and defi nite mo uth (as in P a ra m wciu m

and Vor ti cella ; o r ga ns of l oc o m otio n ( cilia fl a g ella ; organ s
)
) ,

of exc retio n (co nt r act ile v acuoles) etc , etc . .

 C HA PTE R XI V .

UN I C E L L UL A R PL A N TS .

A . P r o to c o cc us .

( P rotococcus P leurococc
,
ua Ch lorococc us
, , Ha ma tococcua , et c .
)

U N I C E L L UL A R pl an ts like u nicellula r animal s ar e ve ry com

, ,

m on although as i ndividu al s m ostl y i nvi sible o n ac cou nt of thei r

,

m icroscopic si z e I n the m as s ho w eve r they are ofte n visible

.
, ,
“
e ither as su s pe n ded o r floati ng matter cau si ng tu r bidity in ,

liquid s ( yeas t ba cter ia d ia to ms demn itls etc ) o r discolo rations

, , , , .

o n t r ee tr unk s
-
ea rth sto ne s r oofs and fl o wer pots ( P ro
, , , ,
-

tococcus , Glazocap aa ,
'

U nder the te r m P rotococc us ( ”p o r o s, first, K o k k o s, berry)

w e m ay fo r o ur p re se nt pu rpose s i nclude a nu m be r of the simple s t
s phe r ic al fo r ms , ge ner all g r ee n in colo r an d of u nce rtain afiin
y
i tie s in cl ass ificatio n but very si mil a r in s t ructu r e livi ng for th e
, ,

mo st part in q uiet water s or o n m oi s t ea r th sto n es t r ee t ru nks , ,

-

o r old roofs or in w ater b utt s r oof gutter s and the l ike

,
-

,
-

, .

S o meti me s the color w hich the y exhibit is y e llowi s h gr ee n -

, g
s o m eti me s blui sh g r ee n and so meti me s thou h le ss ofte n reddish
-

, , ,

a cc o rdi ng to the s pecie s .

O ne of the co mmo ne s t and m o st co nspicuous is a s peci es

ofte n see n o n the shady side of o ld t r ee tru nk s whe re whe n -

a b und ant it for ms a gree ni sh du st like coating o r discolo r atio n

,
-

s c a r cel
y vi sible w he n d r
y but beco mi ng a rich b right g r ee n dur
ingprolo nged r ai ns o r a fte r w a rm showe rs I f pi ece s of bark .

covered w ith thi s for m of P roto cocc us are m oi s ten ed the g r ee n ,

ish c o ati ng may be ob se rved at an ti me I t is g r a nula r in tex

y .

t ure and afte r moi ste ni ng is e as ily l oose ned by a camel s h ai r ’

-

brush .

Morph ol og y .Mic r o scopical ex am i nation show s that the par

t icles detached co nsi s t of r ou n ded ello wish gree n
y
-

e ither s ingl o r
y
 P R O TO O C CC US . 179

E ach i gle cell is a co m plete i ndi vid ual capable of car ryi ng on
s n
,

a n in depe n de n t life I t fair l y r epres e nts the gree n pl ant such

.

(
as P teris) r ed uced to its lo w e st ter ms ( Fig . .

L ike A mwba a nd the I nfu so ri a P rotococcus at le ast in som e ,

s pecie s occurs both in a m otile o r active state in w hich it m ove s

,

ab out and a q uiesce n t o r n on mo tile s tate analogo us to the eu

,
-

c ysted s tate of the u nicellula r ani mal s I n the l atter the motile
. .

o r active st a te is the us ual or do mi nant co nditio n and the en

c s ted s tate is ra r el y as su m ed I n P rotococcus on the othe r
y .
,

h an d the m otile st ate is r ar e and the ordinarv activitie s of the

, ,

pl ant are ca rr ied o n in the n o n motile st ate -

S t r u ctu r e I n s t ruct ur e P rotococc us is a nearl y ty pical cell

.

(p . I t co n s i s ts e ss e n ti all
y of an approxi ma tel y s pherical
mass of p rotopl as m e nclo sed w ithi n a thi n wood y l ay e r of cellu
lose (cell wa ll or cell m e m brane ) and co nt ai ns a si ngle nucleu s
- -

, .

I t al so incl ude s on e or m ore chlo rop hyll bodies ( clzrom a top li ores) -

s able to m a nufacture its own

( p. 1 2 6 b
) y virtue of w hich it i
food s ve ry much after the fas hio n of the gr ee n cell s of P teris
, .

I n t he se fo r ms which po ss e ss a motile stage the l atter c on

s ists of a s phe rical egg sh aped or pea r shaped cell havi ng chro
- -

mato ph ores and a me mbr a ne through w hich two fl agella protr ude .

I n the ov al fo r m s the se a re placed n e a r the narro w ed end of the

c ell and in all case s the a re loco motor orga ns and propel the
, y
c ell s wiftl y th r ough the w ater ( Fig . .

R epr odu cti on The ordi na ry method of r eprod uctio n in the

.

u n i c ellul ar pl ants as in the u nicellul a r ani mal s is by cell divi sion

, ,
-

I n P ro to coccu a the s phere bec o m e s divided by a p a r titio n i nto

tw o cell s w hich eve ntu all y sep a r ate co m pletel y o ne from the
othe r Ve ry ofte n ho weve r the sep ara tio n bei ng i nco mplete
.
, ,

o r po stpo ned u ntil after each d aughter cell h as in t u r n bec om -

divided groups o r agg r egate s of cell s a ri se w hich sugge st the

,

first ste ps in the for matio n of ti ssue in the develop me nt of highe r

for ms I n the end however s epa r atio n is tot al and co mplete
.
, , ,

a nd each cell is therefore no t a u n it in a bod b ut is it self a

y ,

body and an indi v id ual (see p .

( Fig .

The d aughte r cell s th us produced are the y o ung or off sp r i ng

-

, ,

which h ave the power to gr o w and ulti mately to divide in their

t ur n . U nder favor able ci r cu msta nce s generation may thu s fol
low g e ner atio n in quick succe ssio n E ach y oung cell is actuall y .
1 80 UN I CE L L UL A R PL A N TS .

FI G . 92 .
—
P t
~
ro o c oc c us ( Pleurococc us ) f m th b rk o f n lm t r
ro e a a e ee. in ac ti v v gw
e e

t i n n d h wi ng ggr g t i
o a s o a e a on i t m
n o f ll
a ss e s oA Pl u ce s. , c rococcus in th d ie r

co nd i t i B A m m on .
, s c occ s h w i ng d g n u d ivi i n i n t
o en o e o s s o o t wo ce ll nd
s a

i nt f u D E F m t il f
o o r. . , , o e orms of Protococc us w a f ter O o h n)
-
.
 N UTR I TI ON OF PR O TO O C CC US .
81

o ne half of the pa r e n t cell an d co n ta ins a moiety of wh atever

that co ntained Here the r efo r e as in A moeba the proble ms
.
, , ,

of he r edi ty u nco m p licated by the occu rr en ce of sex are 1 educed

,
,

to thei r lowes t ter ms .

I n so m e ki nd s of P roto co cou s the quiesce nt cell s unde r

,

special ci r cum sta nce s w hi ch ar e not well u nder stood give ri se

, ,

to the mo tile fo rm s (z oo sp ores) r efe rred to above C ilia or .

,

rather fl agella a r e fo rm ed an d the p rotoplas mic mass with its

, ,

in cluded ch r o matopho r es swi ms actively about in the water .

A fte r a ti m e th ese m otile cell s m ay co me to r e st lo s e thei r fl a ,

ge lla and divide i n to two or mo r e daughte r cell s each of which -

in its t ur n may beco m e a m otil e cell and r epe at the p r ocess or , ,

unde r o th e r co nditio n s develop i nto the ordi nar y quie sce nt cell
, .

I n so m e s pecies of P ro to coc eus in which there is a motile

stage an othe r fo rm of rep r od uctio n a kin d of rudi me n tary ,

ga moge nesis h as bee n obse r ved I n th is proce ss two of the

,
.

motile c e lls (ga m ete s) meet fu se ( oo nj ug a timz

) lose their fl
, agell a , ,

beco m e e ncysted ( see p and ulti matel

y give
. ri s e to the
o rdina ry ce ll s of P rotococcus both no n motile and motile , .

Th is p r oce ss howeve r has no t yet be e n observed m the species

, ,

u nde r co nside ratio n .

Phy s i olog y O ur actu al kn o wledge of the ph ysiology of

.

P rotoc ocmls is ve ry small B ut the s tud y of comp arative pl ant

.

physi o logy giv es every r easo n to believe th at the e ssenti al phy s

io log ica l o pe ratio ns of th is s i mple pl an t are fund ame n tally of
as in the higher
g r ee n pla nt s such as P teris ,
.

N ut r iti on The i n com e of P rotococc us w he n grow i ng in

.
,

l ts na tural h abitat o n tree b ra n che s m oi st brick s and the like

-

, , ,

is diffic ult to de te r m i n e B ut as it is able to live al so in ordi

.

na r ra i n water w e a re able to se t do w n its prob able in co m e

y
-

co nditio ns with so m e degree of accuracy There .

th at it ab sorb s w ate r and c arbo n di oxide by dif

g h the cell ulose wall an d th at the se sub st ance s ,

the manufactu re of starch w hich if stored up , , ,

mak es its a ppea r an ce in the fo rm of small gr anule s within the

Th is proce ss ta ke s place only in the l ight and
‘

c h r o matophor es .

through the age n cy of the chlo r ophy ll an d is atten ded by a ,

sett ing free of oxy ge n precisely as in P teris N it r oge n is prob .

ably derived fro m n itrates o r ammon i ac al co mpo und s mi nute ,

1 82 UN I CE L L UL A R PL A N TS .

q uantitie s of which are dissolved in the w ate r and other neces ,

sa ry salt s (sulph ate s chlo ride s pho s ph a tes etc ) as well as f ree
, , , .

oxyge n a r e procu red fr o m the sa m e sou r ce These sub stances .

may be de r ived fro m d ust blo wn or w ashed by the rain in to the

w ate r or fro m the w all s o f the vessel
,
To the p r ocess of star ch
.

m aki ng atte nded by the a bso rptio n of C O and

,
and t he ,

“
libe ration of O the ter m ass i mi l atio n is ge ne rally give n
,
.

L ike othe r pl ant s m oreove r P rotococcus p r ob abl y b r ea thes

, ,

by ab so r bi ng free oxyge n and setti ng f r ee C O ( r e sp ,

The i nc o me and outgo of P roto oocous may then be

by the followi ng diag r a m :

Aislmllatlon

I ncome

Respir ti
a on

Itshould be u nde rs tood that this o nly r ep r ese nts the b

outli ne s of the proce ss and u nde r the si mpl est co nditio ns .

quite po ss ible that u nde r othe r co nditio ns P rotococcus m

m ore co mplex food s The fac ts r e mai n ho w eve r ( 1
.
, ,

P rotoco ccus is
action is on the
co mplex compou n ds ( ca rboh y d rate s p r oteid s) out ,

o nes I n these re spect s it shows a co mplete cont ras t

.

which is o n the w hole de structive b r eaki ng down co ,

pou nds i nto si mple r o ne s and is i nde pe nde nt

,

derives ene r gy fr o m the pote ntial e ne r gy of

r el atio ns bet wee n P roto cocc us and A moeba
epito me of the rel ations betwee n P teris and L umbricus ,

bet wee n gr ee n pl ants and ani mal s ge n e rally .

Th e Funda mental Phy si olog i cal Pr operti es of Plants I n .

siderin
g the ph y iology of A moeba we fou nd it poss ible to
s
 PR OTO CO CC US A N D A M G J
BA COMP A R E D . 83

duce its vita l activitie s to a few fundamental physiological proper

ties nam el y co nt ractility irritability metaboli sm gr owth and
, , , , ,

r epr od uctio n co m mo n to all ani m als A little r eflection wi ll,

, .

show that the sa m e prope r ties are manife sted al so by P roto

co ccu s C o nt r actio n and i r ritability are difficult to wit ne ss in
.

the quiesce nt stage of P roto coccus but ob v iou s en ough in the ,

r arer motile fo r ms M et aboli sm g r owth an d reprod uction o n

.
, ,

th e othe r ha nd a r e evide n t acco m p an i me nt s of nor mal life eve n

, ,

in the q uiesce n t co nditio n A n d preci sel y as P roto coccus diff er s

.

fro m A m oeba in r e spect to co nt ractility and ir ritability of which ,

it po ss e ss es r el atively little so pl ant s in ge ne r al d iffer in these

,

respects f r o m a n i mal s in ge ne ral A n i mals are e mi ne ntl y con .

t ractil e an d i rr itable while pla nts a r e but feebly speciali z ed in

,

these di r ectio ns O n the other hand as we have alre ady seen

.
,

in co mpa ri ng P teri s with L u mbri cus ( p an d as we see .

o nce m ore in co mp ar i ng P roto coc c us with A m oeba in re spect to ,

metabol is m the g ree n pla nt is p r e e mi ne ntl y constructive while

,
-

the ani mal is p re emine ntl y de st ructive of organic matter , .

I n the ir m od es of n utritio n as stated above A moeba , ,

and P rotococcus r ep r e se nt tw o ph ysiologic al ext r emes We .

pass no w to the s tu dy of Y e as t s and B acte r ia which are pl ants ,

des titu te of chlo rop hy ll and in a ce r tai n se nse may be r ega r ded
as occup i ng a m iddle g r ou n d betwee n the se ext r e me s
y .

Ot h er Forms . i nnumera b l e s pecies o f uni cell ul ar green

Th ere a re

p l a n t s A.v as t r
g p p
o u o f ec u l i a r b r o w ni s h f o r m s c o v er ed wi th t ra nspa rent

g l a ss -
l i k e cell s co m po s e d o f s i l i c eo u s m a t eri a l is k no wn as th e Dta to

mac ece o r d ia to ms I n th es e th e c hl oro ph yll is mask ed by a b rown pig

.

ment b ut is ne v er th el ess pr es ent

,
A no th er g ro up is th a t k nown a s the
.

Des mid ia or des mid s Th ese o f t en h a v e th e i ndi v i dua l cell s pec ul i arl y
.

c o ns t ri cted in t h e mi ddl e so th a t at fi rs t s i g h t th e t wo h al ves a ppea r to b e

two sepa rate cell s Mo re c l osel y resemb l i ngP rotococcus in many respec t s
.

are so me me mbe rs o f th e Cy a nop h y ceae o r

“
b l ue green al gae a mong
” -

wh i c h C h roococcus a nd G loeoc ap sa di ff er f rom P rotococc us c h i efl y in th e ,

for mer cas e in h a v i nga b l ue gr een i ns t ead of a yell ow green pigment

,
-
-

and in th e l a t ter no t o nl y in t h i s r es pect b ut a l so in th e fac t th a t the

, , ,

s i ng l e cells are widel y sepa rated by t ransparent mucil age .

 C H A PTE R XV .

UN I CE LL UL AR PL AN TS ( Conti nued ) .

B . Y east .

( Saccha romy cea)

U N DE R the gene r al name of ea s t a r e included so me of t
y
“ ”
si mplest fo rms of vegetal life . S o m e y east s a re Wild 1 ,

ingupo n fe r me nti ng f r uits o r in f ru it juices and co mmo ,

FI 93 —
Y t ll B rwr t t iv l y v g t t i ng Th l rge i nt r
’
G . . ea s -
ce s. e e s ( to p ) y ea s ac e e e a . e a e

va c uo l
es a n d th e s ma ll f t d
a - ro p s are s h wn
o , l b ud in v ri u t g
as are a s o s, a o s s a e
de v elo p m en t
, a nd th e c ell -
w a ll . N uc l i n t vi ib l
e o ( Hi gh ly m gni fied )
s e. a .

“ ”
occu rri ng in the air ; othe rs a r e domesticated o r cultiv att ,

such as those regularl y e mploy ed in b r ewi ng and in bak i ng .

“ “
I f a bit of y east cake ( ei t he r co mp r essed
-
o r d r iec

y east) is mixed with w ate r a milky flu id is obtain ed whi ,

closel y r ese mbles the se called bake r s or b rewe r s y eas t -

’ ’
.
 C
S TR U TUR E OF YE A S T .
1 85

Mic r oscopi cal examination p r oves that the milk appearance

y
of li q uid y easts
is due chiefly to the p r ese nce of my r iads of
mi nu te eg g shaped s uspe nded bodies and th at p r essed y east is
-

almost wholly a m ass of si mila r fo r m s These are the cells of .

which is the r efo r e esse ntially a mass of u nicellul ar organ

Fo r r easo ns which wi ll soo n appea r y east is u nive r sally

FI G OL —Y t r b tt
B re w e s ( o o m ) y ea s s o w t h i ng tru tur —p r t pl m
‘
. eas -
o ells . s c e o o as , cell
wa l v
ls, ac uo les , f a t dro ps
-
( N uc le. i n t h wn )
o s o .

r ega rded as a plant ,

and the sin gle cell is often spoken of as the

Morph ology Th e pa r ticular

.
y easts which we shall conside r ar e
the co mmo n cultiv ated for ms of com
merce The cells of an o rdin a ry cake
.

o f p r essed east are s phe ri cal s he

p
y ,

r eid el o r eg
, g shaped in fo rm and co n
-

s ist of a m ass of protopl as m e nclo sed

withi n a well defi ned cell wall -

By -

a pp r op r iate t r eat m e nt the l atte r m ay

be shown to co nsi st of cellulose ; and

it is d istinctl y thicker in old or resti ng zg
cells than in y ou ng o nes o r those vig
F
pfij
p

f br w r y
fi 22123 32;
t ( S rm my
g g
o e e
’
s ea s
r 01

acc
3

o
3

ces

o rous l growi ng Withi n the g ra nul a r

y .

p r o toplasm (cy top la s m) are us ually a number of v acuoles (c on

tainin g sap) an d m i n u t e sh ining do ts ( prob ably fat droplet s) but -

,
1 86 UN I CE L L UL A R P L A N TS .

no chloroph y ll is p r ese n t and no sta rch U ntil r ecentl y the yeast .

cell was supposed to be de stitu te of a nucleus but it is now kn own ,

that eac h cell p r obably possesse s a l ar ge and ch ar acte ristic nucle us .

This howeve r can be de m o ns t r ated onl y by s pecial reage nts and

, , ,

is r a r ely o r neve r seen in the livi ng cell ( Fig .

R epr odu cti on The o r di nary mode of r ep r oductio n

.

is by a modifi catio n of cell divisio n c alled budding -

.

FI G . Oil —Th e N uc l e i
t ll nd t h P o s f B udd i ng ( D wn by J H
of Y ea s -
ce s a e r ce s o . ra . .

E m r t n f m p i m n p r p r d b y S C K i th J ) Th u p p e r l f t h nd figure
e o ro s ec e s e a e . . e . r
. e e -
a

sh w t h nu l u in p i m n t r t d w i th D l fl ld h wm t xy lin Th
o s e c e s a s ec e ea e e a e
’
s a o . e
oth r fi gur in th upp r w nd th in t h l w r ( fr m l ft t ri ght) h w
e es e e ro a o se e o e o e o s o

ce ll in u s iv t g f budd i ng t g th r wi th t h p p n p i ti n nd
s c c ess e s a es o . o e e e a ea ra c e, os o , a

m v m t f t h nu l u
o e I t wi ll b
en s o b v d th t th b ud i f rm d b f r the
e c e s . e o se r e a e s o e e o e

n u l u d ivi d
c e (I h m t xy lin m th d )
s es . ro n -
as a o e o .

favo r able circu msta nces in actively gr o w i ng y east a l ocal bulging

of the wall takes plac e u suall y n ea r but no t p r ecisel y at one , , ,

pole of the cell P rotoplas m p r esses i nto thi s dil atatio n o r

. .

”
bud and exte nds it still fu rthe r A t thi s time we have still .

b ut one cell altho ugh it no w co nsi s ts of two u nequal pa rt s and

,

the se paration of a d aughte r cell is clear l y fo r e shado w ed E ve nt -

. .

ually the co nnectio n be twee n the two pa r ts is s eve red and th e .

daughte r cell or bud is detach ed from the o rigi nal o r pa re nt

-

cell ; but detach me nt may o r may not occu r u ntil afte r the bud
 FOR MA TI ON OF S P OR E S I N YE A S T . 187

has begu n to p r oduce d aughter cell s in its tu r n and more than -

one bud may be bo rn e by eithe r or both pare nt or daughter

cell s I n ve ry r apid growt h the co nnection may persi st between
.

th e cell s eve n du ri ng the fo r matio n of se veral ge neratio ns of

bud s ; but th is is u nu sual and in case s w here a nu mber of cells ,

re m ain a pp a re ntl u nited togethe r for mi ng tree like for ms there

y -

is ofte n no r eal co n nectio n the cell s sep arati ng re adil on agita

y ,

E ndospores y e ( A scospor es)

as t s in additio n to the . S o me
meth od of r ep r oductio n by budding exhibit another mode k now n

FI G . 97 .
—r
S po es o f Y ea s t ( A sc o s po r es ) . Th r ee a nd tw o -
ce ll d t e s a ge of r
sp o e f or
ma ti on in S . cc rev f
s irr .

as en do
g o
eno usr as co s ore omm tio n
pd iv isio n
f U nder certai n .

circu m sta nce s not y et e ntirel y unde r stood there are for med
within the y ea s t cell two th r ee o r fou r r o unded shl mngspore s
-

, , .

Th ese beco me s u rr ou nded by thick w alls an d th us give ri se

eve n tuall y to a group of d aughter cell s within the origi nal cellu -

lose sac To the latter the ter m as cus (sac) has been applied
.
,

an d to its co n tai n ed da ughter cell s the ter m a scosp o res -

.

I t is not yet a ll o wed by all bo t an s ts th a t th is te r mi nol ogy wh i ch im

i ,

pl ies rel a t io nsh ip

a of yeas ts to th e A sco mycet o us f ung i is sound ; but it ,

is co mmo nl y us ed .

ascospore is capable u nde r fa vorable c ircumst ance s of

E ach
sp r outi ng and starti ng a n ew serie s of ge ner ations of ordi nary
y ea s t cells-
I t s hould
. be partic ul a rl y ob served that the e n d o

s pore s of e as t are rep r od uctive bodie s and th at the proce ss of

y ,

their for matio n is o ne of m ultiplication—no t merely one of de

“ ”
fe nce or prote ction as is the c ase with the so called spores
,
-

Qf bac te r ia de sc ri be d bey o nd ( p .
1 88 UNICELL UL A R PL A N TS .

Phy siolog other o r ga ni s ms the y east plan t occu

L ike all
y
-

pie s a defi nite po sitio n in s pac e and ti m e ; it possesses an en

v ironm ent with w hi ch it m u st be in h a r mo ny if it is to li ve ,

fro m w hi ch it derives an i nco me and to w hich it co ntribu tes an ,

outgo of matte r and e ne r gy ; it manufactu r es its o wn su bst ance

fro m food s an d like all livi ng thi ng s it wastes by
oxidatio n of its sub sta nce (ka tab o lis m) I t is not obviou sly co n .

t r actile or ir r itable but it is h ighl y m etabolic and r ep r oductive

,
.

Y east and it s E nvir onment Y east is an aq u atic fo rm and .

, ,

as might be suppo sed c ultivated y eas t th r ives best in its u s ual

,

h abitat the juice s of fruits s uch as apple s o r g rapes and the

, , ,

wate ry ext rac ts of sp r ou ted seeds su c h as ba r ley co rn and , , ,

rye ( wo r t mash ,
I t lives however m o r e o r less success
, , ,

fully in many othe r place s ( such as the dough of b r ead) and can ,

eve n e ndu re much d ryn ess as is show n by the co mme rcial ,

dri ed y east-
”
I t appea rs to p r efer a te m pe ra tu r e f r o m
.

°
2 0 to 3 0 C
°
it is u sually killed by boili ng bu t if dri ed it can
.
, ,

e nd ure high te mpe ratu r e s I ts actio n is i nhibi ted by very low .

te mpe r at ur es but like m ost livi ng thin gs it e ndu r es low te mpe r

,

a tu res better tha n high I t is killed by ma ny poiso ns ( a nti

.

septi cs) .

I ncome O wi ng to its indu st r ial i m po rta nce y east has bee n

.

perh ap s more thoro ughl y s tu died in r espe c t to its nutritio n t han

an othe r u ni cellul a r orga ni sm A n d e t it is i m poss ible to
y y .

give ac cu r ate s ta t i sti c s of its nor mal inco me and ou tgo I t is .

believed th at the ordi nary i nc o me of a yeas t c ell livi ng in wo rt -

( the w ate ry extract of sp r outed bar ley g rai ns) co nsi s ts of a dis -

yg g
°

s olv ed ox en b 7 aztro eno us bod ies allied to pr o teids but d iffu si

,
,
-

ble and able to p ass th rough the c ell ul ose w all ; 0 car bo hydrates , ,

es ecia ll
p y s ug ar
y m a tters a nd d s al ts o f v a rio us kinds , .

I t was s up posed for a lo ng ti me by P asteu r and oth e r s tha t

'

y east co uld di spe nse v nth free ( di ss olved ) oxy ge n in its dietary .

I t no w appe a rs th at thi s fac ulty is te m por ar o nl y an d th at if

y ,

y eas t is to th ri v e it mu st like all othe r li v i ng thi ngs be sup , ,

plied at le as t occasio nally w ith f r ee ox y ge n

, , .

Met aboli sm O ut of the i nco m e of foods ju st d esc ribed y east

.

is able to bu ild up its o wn peculiar protoplas m a na bo lis m and

( ) , ,

d o ubtle ss to lay do wn the d ro plets of f at which ofte n appe a r in

,

it There is go od reaso n to believe that its su bstanc e also b reaks

.
 N UTR I TI ON OF Y E A S T .
1 89

with the production of ca r bon di oxide w ater and n it r o , ,

was te (ka ta bolis m) and the co ncomitant libe r ation of

,

e nergy .The w o r k to be do n e by the y east cell is plai nl

y
-

li mited The m anufactu r e of new and of surplu s protopl asm

.

and the p r o t r usio n of bud s r equi r e work p artl y che mic al , ,

p a r tl y m ec ha ni cal ; bu t m o s t of the libe rated e n erg prob bl

y a
y
appea r s as hea t I n poi nt of fac t gr eat activity of y east is
.
,

acc o mpani ed by a r ise of te m pe ratu re as may beprov ed by ,

pl ac inga ther mo m ete r in ri si ng dough or fer menti ng fruit

”

Outgo . B ar r ing
the outgo of e nergy al r e ady me ntioned and ,

the pro bable exc r etio n of ca r bo n dioxide and nitroge nous waste ,

but little can be sa id co nce r n ing the outgo of a y e ast cell The -

o rdina ry excretio ns a r e so masked by the pre se nce of foreign

matters in the li quid s w hich y eas t i nh abit s th at little is kn ow n of
the r ea l cou rse of eve nt s To the co nside ratio n of co nditions
.

which e ntail these diffi cultie s we may n o w p ass me r ely p ausi ng ,

to c a utio n the s tude n t agai nst the suppo s itio n th at the evol utio n
of ca r bo n dioxide in fe rme ntatio ns r epre se nts to any great ex
te nt the no rmal r esp ir atio n of the y east cell s .

Mi ner al N ut r i ents of Y east I t h as bee n s ho w n ( pp 1 4 8 1 8 1 )

. .
,

that P teris and P rotococcus inasmuch as they po ss e ss chl o rophy ll

,

ca n li ve u po n s i m ple in orga n ic matter s such as C O H O and ,, , ,

n it r a tes out of w hich the a r e able to ma nufact ure for the m

, y
s elv es e nergi z ed foods su ch as st a rch Y east is unable to do
‘

thi s as might be supposed f r o m the fact that it is de stit ute of

,

chloroph y ll A nd y et y east doe s not require proteid re ady

.

made as all t ru e an i mals do for experi me nt s h ave shown th at it

,

ca n live and g r ow in a liqu id co n ta i ning o nly m i ner al matter s

plus so m e such co m pou n d of n it r oge n as am mo niu m t artr ate

Upo n a m uch le ss co mplex organic compound
of nit r oge n such as a nitr ate it cannot thrive th us show i ng its ,

i nfe rio rity in co ns t ru ctive power to P rotococcus and all green

plan t s o n the o ne hand an d its supe r iority to A mazba and all
, ,

a ni mal s o n the other

,
.

Pas t eur s fl uid co mposed o f water and sal ts amongwh ic h I S a mmoni um

’

, ,

t a rt ra t e ( bo v e) wi ll s uffi ce to s uppo rt yea s t I t wi ll s uppo rt a muc h mo re

a ,
.

v igo ro us g ro wth if sugar be added to it B ut if a mmo ni um ni t rat e i s sub

.

s t i t ut ed f o r a mmo ni um ta r t ra te yeas t wi ll re f us e to gro w in th e fl uid .

 90 UN I CELL UL A R P L A N TS .

Y east is 3 Pla nt
The superio r const r uctive faculty of y east
.
,

j ust desc ri bed separ ates it fundam e ntally fr o m all animfl s in

,

r e spect to its physiolog y an d allie s it closel y to all plan ts ,

I ts .

i nferiority to the chlo r oph yll bea r i ng plant s or pa r ts of plants on -

the othe r hand in no wise sep ar ate s it fu ndamen tall y fr om

,

plan ts ; fo r it mu st no t be forgotten th at the powe r eve n o f ,

pl ant cel ls to utiliz e mi n e r al matte rs as raw materials an d fr o m

-

the m to m anufacture food s like sta r ch ordi na ril y resides exclu ,

siv el
y in the chloroph y ll bodie s and is ope r ative o nl
y i n the ,

prese nce of light I t follo w s the r efo r e that most of the cel ls
.
, , ,

even of the so called g ree n pl ant s and a co nside r able po rtion of

-

the co nte nt s of the so called gr ee n cell s mu st be destitute of -

th is synthetic po we r C o nside ratio ns of this kind show how

.

exceedi ngl y locali z ed and s pecial the s ta r ch maki ng fu nct ion is -

,
“ ”
even in the g r ee n pla n ts ; and y east p r obably co mpares ve ry
favo r abl y in its sy nthetic po we r s with ma ny of t he colo r les s ce lls
of such pl ant s or eve n with the colo r less p r otoplasmic po r tio ns
,

of chro matopho r e bea r i ng cells -

.

B ut yeast is vegetal r athe r than ani mal m o rphol ogicall y as ,

well as ph ysiologicall y I ts st r uctu re m o r e n ea rl y r es e mbl es

.

that of so me u ndoubted pla nts ( fungi ) than any ani mal I ts .

wall is co mpo sed of a va riety of cellulos e called fu ng us cellulose ; ,

-

and cellulose though occasio nall

, y occu r ri ng in a ni mal st r uctu r es ,

is bro adly s peaki ng a vegetal co m pou nd

,
Fi nall y in its
,
.
,

method s of r ep r od uctio n by buddi ng and by spo res y eas t is , ,

allied r athe r to pla nts tha n a ni mals .

Top Y east . B ot tom Y eas t . I n the p rocess of b rewi ng two well

mark ed va ri et ies of yeas t occur k no wn as to p a nd bo tt om yeas t ,
“ " “ "
.

The fo r mer is used in th e ma k i ngo f E ng l is h ale s to ut a nd po rter ; the , ,

l a tt er in th e mak i ngo f German o r l ager beer The to p yeas t is cult i .

v ated a t t h e o rdi na r
y s ummer te mpera t ure o f a roo m wi th o ut s pec i al at ,

tent io n to t emper at ure th e l a t te r in roo ms a rt i fi c i all y coo l ed so th a t even

in s ummer i ci c l es o f ten h a ngf rom th e wa lls
, The t wo yeas ts a l so s h o w .

o bv i o us difi erences in f o rm s i ze a nd s t ruc t ure a nd h o w muc h th ey mus t

, ,

difi er in th eir f unc t io n is pla i n f ro m th e v ery di ff erent prod uc t s to wh ic h

th ey gi v e ri se .

W ild Y e as ts . B es i des th e lt i v a ted yeas t s th ere are

co mmerc i a l or c u

a l so wi l d yeas ts a nd to th em are due in th e ma i n th e ferme nta t io ns o f

,

appl e j ui ce o f
-

, g rape j uice a nd o th er fr ui t j uices A d rop o f s weet cider

-

,
'

s h o ws under th e mi c rosco e a
p g ood ex a mpl e o f o ne o f th ese s peci es ; a nd
Pas t eur l ongag o p ro v ed th a t th e o ute r s k i ns o f ri e f
p gp r a es a nd o th er r ui ts
 VA R I E TI E S OF Y E A S T . 19 1
‘

are apt to h a bo r r ll s in th e dus t wh i ch l odges upon th em Mo re

yeas t ce
-

recently it h as bee n s h own th at wi l d yeas t s o f t en l i v e under appl e t rees -

upo n t h e s ur f a c e o f t h e ea r th I n a dry t i me th e wi nd eas i l y l i f t s the dust

.

co nta i ni ng th em a nd co n v eys th em o v er g rea t di s t ances ( of A moeba

*

.
,

I nf uso ri a Th e do mes t ica ted yeas t s of to day are prob ab l th e de

y
-

scendants o f s i mi la r wi l d y eas t s .

R e d Y eas t O ne o f th e fi nes t o f th e wi l d yeas t s is th e so call ed “ red

.
-

t h i c h is f ur th e rmo re v er y easy to s t udy R ed yeas t and many

”
y eas ,
w .
,

ro w l ux uri a ntl y upo n a j ell y made b

o th ers no t red g , y th ick eni ngb eer
,

wo r t wi th co mmo n g el a t i ne I n th i s wa y.
“
pu r e c u lt ur es — th a t is c ul ”
,

tu res f ree f ro m o th e r s pec i es o f yeas t s o r b act eri a and co nsi st i ngo f o ne

, ,

k i nd on l y—c a n be eas i l y made and s t udi ed The mi croscope s h o ws th a t .

th e cel ls o f red yeas t wh i c h fo r m red do ts upo n s uc h j ell y a re no t th em

, ,

sel ves co l o red b ut t h e pi g

,
men t a ppea rs to lie be t ween th e cell s as in th e ,

case o f t he mi racl e g e rm ( B acillus p rodig ios us ) .

Ferment a ti on To the processes where y east is employ ed to

. .

p r oduce che mical ch anges in various dome stic agricult ural and , ,

i n du st r ial ope r atio ns the ter m f ermenta tion or more often ,

“
erm ent a tio n ad
”
alcoho lic
f is applied I n the , r ai s i ng of bre .

o r cake in b r e w i ng cide r mak i ng etc

, , y east acti ng upo n
-

,
.
,

suga r p r oduces fr o m it an abu nd ance of alcohol and c arbon

dioxide B oth p r oducts are sought for in brewi ng and carbon
.
,

dioxi de is especially des ired in bread maki ng -

.

B ut alc oholic fer me ntati o n is o nl y o ne exa m ple of a large

class and y east is o nl y o ne of many fer ment s We may there
,
.
,

f o r e po s tpo n e f urthe r c o ns ide ra tio n o f fer me nt atio n to the n e xt

,

R ela ted Forms . been s h o wn by th e research es of Hansen th at

I t h as
o rdi na ry co mmerc i a l yeas t is s el do m o ne s i ng l e species as was fo r merl y ,

s uppo sed b ut ra th er a mi x t ure o f se v er a l spec i es

,
I t is th erefo re no .

l o nger sa fe to spea k o f commerc ial yeas t as S accha ro myces cerevis ia unl ess ,

c a re ful e x a mi na t i o n b y th e moder n meth ods h a s s h o wn it to b e s uc h a nd

to det er mi ne wh at S pec i es ex i s t in a ny pa r t i cul a r speci men is o f t en a l abori

o us a nd di ffi c ult ma t ter .

I nas muc h as th e na t ura l po si t i on o f yeas t in th e veg etal k i ng dom is

not es t a b l is h ed bey o nd a ll do ub t it is i mpo ss i b l e to s ta t e pr eci sel y wh a t

,

a e its nea
r r r l
e a t i v es T h e re
. a re n um er o us uni ce ll ul a r c o l o r l ess pl a n t s b u t ,

t h ey are no t n ecess ari l y c l osel y rel a ted to yeas t and th e s tudent must not
a nis m
co nc l ude f o r pl a nt s a ny mo r e th an f o r a ni ma l s th at beca use an o rg

uni cellul ar it is necessa ri l y a t th e v ery bo tt om of th e s ca l e of l i f e .

 C H A PT E R X VI .

UN I C E LL UL A R PL A N TS ( s tinued) .

C . B ac te ri a .

( Sch izomy cetes )

THE smallest and the m ost nu me r ou s of all li v i ng thin gs a re

,

th e bacte ria B acte ri a occu r al mo st eve ry whe re : the y a re lifted

.

in to the at m os phe r e as dust pa rticle s in it they float and with its ,

cu rre nt s they are d rive n about ; w ate r— both f re sh and sal t

o fte n co n tai ns l a r ge nu mbe rs of the m ; and the uppe r lay e rs of

the soil teem w ith the m B ut they a r e m ost abun da nt in li q uids

.

c o nt ai ni ng di ss ol ved o r ga n ic m atte rs e s peci all y such as have stood

,

fo r a t ime —fo r ex ample s tale milk and sewage th ese fl uids

, ,

o fte n co n tai ni ng millio ns of i ndividual bacte ria in a single cubic

centimet re .

I n re s pe ct to thei r abu nda nce in the s u rface lay e rs of the

e a rth (o ne g ra m of fe rtile soil ofte n co n ta i ni ng a m illio n o r m o re) ,

a nd the wo r k w hich they do the re in p rodu c i ng the o xidatio n of

o rga nic m atte rs and ch a nge s in the co m po s iti o n of the soil bac ,

te ri a may w ell be co m pared with e a rth wo rms (c f p They . .

a re al so of m uch ge ne ra l i n te re st beca use so me a re

kn ow n as di sease ge rms -
”
Mos t bacte r ia h o wev e
.
,

a rasi tic but s a ro /i tic i e live upo n dead o rga nic matte rs
p , p p y ,
. .
, ,

a nd therefore are no t merel ha rm le ss but po sitivel y useful in

y ,

re nde ri ng back to the i norga nic w orld u sele ss orga nic

S o me specie s such as the v in ega r bacte ria a r e c om
i m porta nt .

I n sy ste matic bota ny b ac te ri a co nstitute a well defi ned group -

t he S chiz o my cetes (fi s s io n u ng i their ea r allies bei ng the

f ) n ,

Cya nop /i y ceaz o r blue gree n algae

-
.

Mor ph olog U nder the m ic roscope bacteri a ap pear as

‘

y .

m i n ut e r o d s ( B a c illi) ( Fig .b all s ( 000027) ( Fig o r s pir als .

( p r illa ) ( Fig
S i . s o m eti m e s a t re s t b ut ofte n at lea s t in , ,

t h e cas e o f the r o d s a nd spi ra l s in active m oti o n L it t le or no

,
-

.
 S HA PE S OF B A CTE R I A .
193

st ructu r e can be made out in the m by the begi nne r to whom ,

they u sually appea r at fi rst sight like p ale transluce nt or wate r
y ,
b i ts of p rotoplas m I n ve stigatio n has show n however that the
.

y , ,

pos sess a cell wall (p r obably composed of cellulose) and a non

-

ho moge neous protoplasm Unlike P rotococc as but like east

y
.

,
cells the ce lls of bacte r ia co nta i n no chlo r oph ll
,
y N uclear mat .

E m . 99 .
-
B ac illi f rom
B ay I t in
'

f us ion ( uns
t a

ed) . l Th e fi a m en
t t s a

t h l f t in
e e di ti n a con o

o f t iv v g t t i
ac e e e a o n.

Th m i dd l fi l m n t
e e a e

Fm .as B ac us M eg
. i ll t i
a er um . f rm i g p r
o nTh s o es . e

R eds ( un sta n ed ) in i a v ri
o us fi l m t t th ri ght
a en o e

ag g reg a t io n s a s c o m m o n ly se en t i fi
c on a n s p r ve s o es

wi th h i h
a r ft r th i
g po w e a e e r l d i th rw i
en c o se n o e se

t i ti
c ul v a o n b ilin o u l on a nd emp ty ll th w ll ce s, e a s

hi i l g i
w le ra p d y ro w n gan d m ul of wh i h bulg pr b c e, o a

tip ly in g b ytr v iv
a n s erse d i b ly f ro m th e a b so rp

d on . tion o f wa e tr .

ter is p r ese nt eithe r scatte red about o r if the vie ws of B utschli

, , ,

be accepted co m posi ng m ost of the p r o topla smic body itself

,
.

Many bacte ri a be a r appe nd ages in the sh ape of flagella o r

cili a ; but the se can onl y be demonst rated in special cases and ,

by s peci al methods They a r e believed to be locomoto r organs ,

a nd in so m e cases have bee n see n in active m

.

otio n ( Fig .
194 UN I CE L L UL A R PL A N TS .

The mi nute ne ss of bacte ria is ext rao r di na ry Many bacilli are .

n ot m ore th a n 00 5 m m (” n
i i nch ) i
.
n le n gth o r m o r e tha
. n 001 .

mm (1 m n inch ) in b r eadth
. 7
i
S om e a re ve ry m uch smalle r . .

M ost bacte ri a a re at so m e tim e f ree fo r m s ; but like othe r

u nicell ul ar o rga nis ms many of the m have the powe r to pass
fro m a fr ee swi mm i ng (s wa r ming -

) i nto a q uiesce nt
( res ting )
condition I n the l at te r so me u nde rgo a pec ulia r cha nge in
.
,

which the cell wall be c o me s m ucilag -

i n ou s and by the aggrega ,

tio n of nu me r ou s i ndividual s o r by r epeated divi sio n lu mp s of

jelly like co nsi ste ncy (zoog
-
loea ) a r ise I f the jelly mass tak es .

the shape of a sheet or me mb r anous ski n ( as happe ns in the

mother of vi nega r) it is so meti mes desc ri bed as M ycoderma
- -

(f g u n u s sk in ) ( Fig -
.

R epr oducti on The b acte r ia i nc rease in nu mbe r s solel y by

.

t ransve r se divi sio n G r owth ta kes pl ace and is follow ed by t rans

.

ve rse divi sio n of the o r igi nal c ell usuallv i nto halv es E ach h alf , .

the n like wise gr ow s and divides in its tu rn I n this way m ulti .

plicatio n may go o n in geo met r ical progr essio n and with almo st ,

i nc r edible r apidity I t has bee n sta ted that such r epeated divi.

sio ns may follow onl y an hou r apa r t and on th is basis it is easy ,

to compute the e no rm ous nu m be r s to which a si ngle cell may

give r ise in a si ngle day .

I f sepa ratio n afte r divi sion is co mplete st r ictl y u nicellula r ,

fo r ms arise I f actual sepa ratio n is postpo ned lo ng r ods chains

.
, , ,

o r plate s ( in the case o f cocci )

may appea r D iff e r e nt names .

ar e give n to the r e sulti ng fo r ms .

S trep to cocc us is a mo nilifor m

o r necklace like a rrange me nt ; -

S tap hy lo cocc us s i ngle cocci ,

D ip loccocc us cocci in pairs ,

L ep to thrix a filame nt of ,

bacilli ; S a rcina a plate of ,

cocci r ese mbli ng a ca r d of bis

cm t two o r m ca r ds
i Fm lOl —S h rt
01
0—
‘
o re
FI G . 10 Mi r . c oco c c . . o
’

( u t i d ) tr m b y
n s a ne B i ll i o (u supe r pos ed ; etc etc
a ac n .
, .

“mm “ St i n d ) f m S o me bacte r i a p r o
1
Spor es a e ro
h y mm i n
.

a o .

duce so called sp ores (end o -

sp ores
) in the followi ng way : The co nte nts of the cell
 S P OR E S OF B A CTE R I A .
19 5

withd r aw f r o m the wall and co ndense i nto a (u sually oval

on e en d of the cell
)
leavi ng the re st of it e mpt
,
y
thi s ti m e th at the cell wall -

is be s t s ee n The co nde nsed mass

.

no w be c o m e s d a rk and O paque appa ,

ren tl
y fro m the depo sit upo n it self of a
g r eatly thicke ned and peculia r w all ; it
r efu se s to ab so r b s tai ns which the o r igi
nal cell would have take n and beco mes ,

e xceedingl
y r e sista nt to extre m e s of
heat Cold and dr yn ess ( Fig 1 05 ) TO FI 102 —
) ? Th M th r f G . .
'

e o e -
o .

th ese spo r e s the G er mans give the V i g r Th d g f ne a e e e o a

fi lm f z gl f m th r o oo o ea o
s x c e l l e n t te r m Da uersp oren i e m mgg i t pp r
,
. .
, o fl as a
o

ea
e

s
u d r h gh p w
en d u r z n i Th n e a o er e
9
,

b t ri i mb dd d ac e a a re s een e e
8 p 07 e 8 j lly wh i h th y
’
i th n e e / c e

ofte n called h v r t d a e sec e e .

g
res tin spo r e s Whe n bro ught u nder .

favo r able co nditions the se sprout ,

an d the ordi nary bacteri um cell

,

havi ng bee n prod uced gro wth and ,

fi ss io n proceed as before O bviously .

the se S pore s are very different in

fu nctio n from those of P teris (p .

si nce they are prote c tive

m e r el y and not r eproductive They ,
.

co rr e spo nd doubtle ss to that ph ase , ,

of ani mal life which is k n o wn as the

e ncysted st ate A nother m ode .

of spo r e for mation in b acteri a is that -

kn ow n as the prod uction of a rt/tro

sp ores in w hi c h a lo ng sle nder cell ,

may beco m e constricted and det ach

daughter cell s fro m one or both e nd s -
.

Thi s is obviously a speci al c ase of

B t ri Th
ac e u nequal cell divi sio n but if it exi sts
a . e -

,
b i l lu f t y p h i d f v r h w i ng at all wh i ch has bee n dollb ted it
ac s o o e e s o
(
,

c i li ( Fr m “ w m n p r p r d
a . o o e e a e

by Dr wn b y J H clea r l a pproache s agam oge ne si s

y
a o

in su ch f or ms as P teris .

Ph y s i olog I nco m e M etab oli sm an d Out go The bacteria

y .
, ,
.
19 6 UN I CE L L UL A R PL A N TS .

s how a su r p risi ng dive r sity in the precise co nditions of their

nut r itio n and it is therefo r e difficult to m ake fo r the m a
,

sati sfacto ry gener al state me n t A s a g r ou p however and dis .

, ,
.

r egar di ng fo r the mo me nt ce rtain i mpo r ta nt exceptio ns they are ,

to be r egarded as colo r less pl ants livi ng fo r the mo st part u po n

co mplex orga nic co mpou nds f r o m which they derive thei r in .

co me of matte r and e ne rgy and which they deco mpose i nto

si mple r co m poun ds poo re r in pote n
tial e ne r gy I n so doi ng the y .

b r i ng about ce r tai n che mical

cha nge s in the s ubsta nces u po n
which they act which ar e of the
highe st th eo retical i nte r est and ,

so meti m es of g r eat pr actical im

po rta nce Pe r haps the mos t pecal .

ia r featu r e of the ph y siology of

bacte ria is the fact th at w hile they
a r e the mselves i ndividually i nvisi
ble they coll ectivel y produ ce ver y ,

F lG104 —
. S p i r i ll m u
. u nd ul S pir l co n s p i cu ou s a n da.
l mpo m nt C h ang
a
es
b t ri d p l y m m d D w in thei r e nvi ro nmen t Fo r ex am
ac e a ee i e ra "
tr m t h fi r t p h t g r ph i p
.

o e s o o a c re re

nt t i n
se a f b t ri
o v r pub ple Vi nega r bacte ria aC t U PO“
o ac e a e e ,
“Sh a “ th t f “ h m K M alcohol in cide r etc ) and by a
(
e r a O O “
in C h B tt d g l87 6 )
.
‘ ,
o n s e r e, .

p r ocess of ox i dat i o n slowly co nve rt

it i nto ac etic acid and water thus ,

o, on e , no .

He re it is not the bac te ria that a r e m o st conspicuous but the ,

e ffect which they p r odu ce I t is clear that the alcohol can be .

only one fac to r in the nut r i me nt of the o rga ni sm because it ,

co ntai ns no nitroge n and the abo ve r e actio n ca nnot r ep r ese nt

,

more th an a ph ase in the nutritio n of the bac te r iu m That thi s .

is i ndeed the case is p roved by the fact th at if the co ndi tions be

so mewhat ch anged the sa me bacteria may go fu r the r and co nve r t
the acetic acid itself into ca rbo nic acid and wate r

C , H, O , O, 2 C O, 2 H, O .

C he mical chang
es of this kind in which the effect u po n th e en
 FE R ME N TS A N D FE R ME N TA TI ON . 1 97

v ironment is mo r e co nspicuous than and out of all propo rtion to ,

,
the change in the age nt a r e in some case s k now n as f ermen
ta t/ions and the age nt effecti ng the change is described as a
,

f ermen t S m m g
o e fe r e
. nts a re or a n i zed o r liv in
g and so me are ,

FI G .
—B i ll u m g t i um ( x Gill) S p r f rm t i n d g rmi t i n A
10
6 . ac s e a er . o e o a o an e na o
p i f r d f rmi g p l k M B th m b ut h u
.
.
a a r o o b ut 2
s o n s o res, a o
’
o c oc P . .
, e sa e a o an o r
l t a er C . h ur l t r t i l l Th p r i C w r m tur b y
, one o a e s i g; th
. e s o es n e e a e e v en n e o ne
app t ly b gu i t h th i rd up p
a ren e l l f A d B di pp d th l l i 0
n n e er c e o an sa ea re e ce s n
w h i h did
c t t i p r w r d d by 9 M D fi
no co n a n s ll d d wi th thr
o es e e ea P . .
, a ve c e -
e ro ee
rip p r pl d i
e s u tr i
o es , t lut i ac e ft r d y i gf
n a n v ld y t en so o n, a e r n o r se era a s, a
P JJ E th
. m p im , b ut
e sa m F th
e s m
ec b ut 4 M G p i r f
en a o a.
, e sa e a o P . .
, a a o
ordi ry d i t iv v g t t i
na ro s d m ti
n ac ( A f t r D B ry )
e e e a o n an o o n. e e a .

a n or a gmized o r lif eless O f the fo rmer the vi nega r bacteri um .

and y eas t a r e good ex ample s O f the l a tte r the dige stive fer .

me nts l ike p ep s in p ty alin and tryp s in and ce rtai n vegetal

, , , ,

fer me nts like dias tas e of malt a r e famili ar i ns ta nces

, .

A s a rule the bac te r ia se e m to prefe r neutral o r slightl y

alkalin e nit r oge n ou s food s They the r efo r e decompose more .

r eadil y meats m ilk and substa nces ( such as beef tea) made of
, ,
-

anim al m atte rs ; le ss r eadily acid f ru its ti mbe r etc I f in the , , .

cou rse of thei r ac ti v ity they deco mpo se m ea ts or fi sh eggs etc , , ,

.
,

with the p r oductio n of evil smelli ng gase s o r put r id odors the -

p r oce ss is k no wn as p utrq f a c tio n R arely b acteria i nv ade the .

,

a ni mal ( o r pla nt) bod y and act upo n the organic matter s which .

they fi nd the re I n s uch case s diseas e may re sult and the

.
,

bacte ria co ncer ned are the n k now n as disease g erms .

B ut w hile b acteri a ap pea r to p refe r highl y organi z ed nitrog

eno us p roteid fo od the a r e n o m e ans depe nde nt upo n it
( ) y by ,
.

E xpe rime nts h a ve show n th at ma ny s pecie s can thrive upo n

Pasteu r s fluid a liquid co n tai n ing o nly a mm o ni um tartr ate and
’
,

ce rtai n pu rel y i no rga nic sub stance s ; and one bacteri um at leas t ,
19 8 UN I CE L L UL A R PL A N TS :

been said already to p rove tha t the bmteria

pl ants can live u po n ino r gan ic food B ut if .

just r efe rr ed to a re c o rrect bacte ria a re not on] ,

spite of thei r lac k of chlo roph y ll some at ,

able lik e gree n plan ts to ma n uf acture their own

,

the raw mate rials of the

this fac t in studies of the

to be dege ne rate fo rms They may ha ve bee n the

.

fo rms of life .

A s was the cas e with y eas t and P rotococc us it is ,

diflicult to make any p r ecise state me nt co nce rning the

outgo
al way
so me
n m o re favo rable ca se s th e inco m e appea r s
g en . I
p rote ids fats and ca r boh y d rates o r thei r eq ui valen ts
, ,
.

f r eely used unde r so me ci rcu mstances ; and fats ( wh en

and p rote ids pepto ni z ed o r othe r wise alte red might
, ,

abso r bed I t is p r obable tha t soluble

.

the bac te ria w hi c h dis so lve and make

, ,

s uch as mea t o r white of e

gg;
a ki nd of exte rnal digest io n .

that bact e ria can live and mul

terials so sma ll as al most o r q ui te to elu
it is fai r to say that they are amo ng the m ost
r eage nts .

I t must not be in ferred from wh at h as been said above t hat has ten
a l ways o xi dizi ng ag ents B roadly s pea ki nga nd in the l o ngrun the
.

s uch a nd in th is res pec t th ey rese mble a ni ma ls

, L i ke the lat ter t he
.

una bl e ( beca use o f wan t o f c hl o ro ph y ll ) to utilize so la r energy a n d 1 ,

f o re mus t o b ta i n th eir energy by o x idi zi ngth eir food Y et under 01 .

c irc ums ta nc es bac t e ria ac t as red uc i n

gagen ts as f o r exa mpl e when
.
, , ,

red uce ni t ra te s to a mmo nia Th is ac tio n o nl y ta kes pl ace bo rrow

.
,

t h e presence o f o rg a nic ma tter a nd a ppea rs to be merely an incid

,

e ff ect th e o x en o f t he ni tra te bei ngneeded f o r t he o x i dati o n o f ca

, yg
Wha t at fi rs t sig ht appea rs to be a n ex cept io n t herefore pro v es in th
, ,

to be a pa rt o f a g enera l la w t ha t bacte ria l i k e ani ma ls a re oxit

, ,

ag ent s are dependen t f or th eir ene r

, gy upo n t he potentia l energ y of
f oods a nd are una ble to uti l ize so la r energ
, y (p .
 M E THODS OF S TE R I L I ZI N G 199

I t h asrec entl y been s h o wn th a t many bac t eri a under circums tances

o th er wis e f a v o r a b l e a re k i ll ed by ex po s ur e to sunl i ht
g .

R ela ted Forms A cc o rdi ngt o o ur presen t i deas o f cl ass i fi cat i on th

.

e
bac teri a fo r m a so me wh a t i s o l a ted g ro up th eir nea res t rel at i v es bei ngthe ,

s li me mo ul ds ( M yxo mycetes ) a nd es pe ci a ll y t h e M y xoba cteri a o f Th a x ter o

-

n ,

t h e o ne h a nd a nd t h e Cy nop h y ceafz t h
,
a
“
b l ue gr n o fis si on al gae e -
ee r ”

o n th e o th er N ei th er o f th ese h o we v er need be co nsi dered h ere

.
, , .

Why B act er ia are Consi der ed to be Plant s The b ac teri a were .

fo rmerly regarded as in fu so ri al ani malcule s (because they abou nd

.

in i nfusio ns a nd many have the po w er of activ e move me nt

,
) .

They a re s till rega rded by so m e as a ni mals Most biologi sts .

ho w eve r r egar d the m as pla nt s becau se they can live w itho ut

, ,

proteid food ( which no a ni mal s o far as k nown can do) and , , ,

because in their m ethod of r eprod uctio n and in thei r gro wth

fo r ms they m o re nea r ly rese mble the Cy a nop /ty cew th an they do
an anim s l The re is also r easo n to thi nk that thei r cell wall is
y . -

c o m posed of cellulose
. .

B act eri a a ndironment The rel at io ns of o rga ni s ms to tem

th eir E n v .

perature a nd mo i s t ure h a v e been mo re th oroug hl y s t udied f o r the ba c teri a

t h a n f o r a ny o th er uni c ell ul a r o rg a ni s ms o n a cco unt o f th eir bea ri n u on
gp
moder n th eo ri es o f i nfec t i o us dis eas e I n g enera l t empera t ures a bo v e
.
,

7 0 C are fa ta l to o rdi nary ba c t eri a In g enera l as is s h o wn by c o mmo n

°
. .
,

e xperi ence wi th t h e k ee pi ng o f foods in co l d s to rage bact eria are be

“
,

n umbed b ut no t k i ll ed by modera te c o l d B ut in speci al cases pa r t i en .

,

larly wh e n th ey a re d ri ed sl o wl y b ac t eri a may wi th s ta nd ev en prol o ng

, ed

bo i l i ngo r f ree zi ngo r t h e ac t i o n o f po iso ns so th a t t h e r emova l or des t r uo ,

t io n o f th e l as t t races o f b ac t e ri a l l i fe is o f t en v ery di ffi cult .

S t erili za ti on a nd P as teuriz i ng Th e r emo v a l o f all t races of l ivi ng

.

ma t ter f ro m a ny s ubs t a nce a nd in pa r t i c ul a r th e des t r uc t i o n o f all b ac

,

teris l l i fe is k no wn as s teriliza tion

, To f r ee o rg a ni c s ub s tances f ro m th e
.

la rg er f o r ms o f l i fe is a c o mpa ra t i v el y eas y ma tt er ; but b ac t eri a a re s o

mi nute a nd so ub i q ui t o us th a t sca rcel y a ny th i ng is n or mall y f ree f rom

t h em a nd th ey a re s o h a r dy th at it is ex ceedi ng
,
l y di ffi c ul t to des t roy th em
wi th o ut at th e sa me t i me des t ro yi ngth e s ub s t a nces wh i c h it is desired to
s t eri l ize Th ey a re no t no r ma ll y present in th e l i v i ngt i ss ues o f pl ant s or
.

a ni ma l s wh i c h are s ea l ed a a i ns t th eir en t ra nc e by s k i ns o r epi th el i a ; b ut

g
a f t er th ese are b r o k en o r cut o en ( as in wo unds) b ac t eri a speedi l y i nv ade
p
the t iss ues O rdi na ry ea r th as h a s b een sa id a b o v e t eems wi th b ac t eri a
.
, , ,

wh i c h are eas i l y dri ed and di ss emi na t ed in dus t dri v en by th e wi nd Wh at .

e v er is in co nta c t th e ref o r e wi th th e a ir o r expos ed to dus t o r dir t is nev er

, ,

f ree fro m b act eri a a nd mea t or mi l k wh ic h in th e l i v i ngani mal are nor

,

ma ll y s teri l e if e x posed to t h e a ir soo n beco me co nt a minat ed wi th bac teri a

,
.

S teriliza tion ts uch as is req uired to p res er v e ca nned g oo ds f or ex a mpl e) ,

2 00 UN I CE L L UL A R PL A N TS .

may be efi ec ted by h ea t ti nued a fter cooling by excl us ion

and co n , ,

ger m l a den a ir Dis inf ed io n wh i c h is th e d es t ruc tio n of bac terial li fe

-
.
,

powerf ul po i sons is a no t h er fo rm o f s teril iza t ion S t ill anot h er is fl it ;

, .

t io n th ro ug h media imp ervious to g er ms s uc h as occ urs in th e we

,

k nown c l a y or po rce l ain wa te r fi lters

, , I n th e l as t case t h e po res o f t
-
.

fi lt er are l a rge enoug h to all ow th e wa ter very s l o wly to pass b ut too s m ,

f or th e bac t eri a .

I n s o m e c ases es pec i all y t h ose in wh i ch d isease p rod uci ng ( p a thog

,
-
en

g er ms may be presen t and yet i t is i m poss i bl e to use po iso ns and undes ii

b le to us e a h i g h t em pera t ur e Pas teuriza tio n is reso rted to
, Th i s on .

s i s ts in h ea ti ng to a t em per a t ure ( us ua ll y 7 5 hig h enoug h t o des tr

°

t he pa rti c ul ar p a t h og eni c ger ms s uppo sed to be p resen t b ut not h i ,

eno ug h to a l ter th e diges t i b ili t y or o t h er va l ua bl e p ro per t i es o f th e liq u

in q ues t io n.

F or th e medi c a l ec o no m i c a nd s a nita ry as pec ts o f p ro bl ems r el at

. .

to th e b ac teria r e fe re nce m us t be h a d to t he num ero us t rea t i ses u]

,

Ba cterio lo gy pe r h a ps t h e yo unges t a nd certa i nl y o ne of th e mos t imp

, ,

ta nt of t h e b iol o g i ca l s c i e nc s
,
e .
 C HA PT E R X VII .

A HA Y I N FUS I ON .

I a wisp of ha y is pu t in to a beak e r of wat er a n d th e mix

F

ture a ll owed t o s t a n d i n a wa rm p l ace th e r e is soo n fo r m ed w h a t

is k n o wn as a hazy mf uswn M ic r oscopica l exam i n at io n of a
.

drop of th e li quid a t th e end of th e fi r s t h ou r o r t wo r evea l s

h ttle o r n o thin g a n d if th e bea k e r be h e l d up t o th e l ight th e
,

Bu t aft e r so m e h ou rs a m arked
fou n d t o h ave take n p l ace T h e l iquid o r igi na ll y .
,

cl ea r h as beco m e c l oud y and a d r op of i t exa m in ed mic rosco

, , p
ically wi ll be fou n d to be swar m i ng with bac t e r ia A day o r .

t wo l a t e r th e c l oudi n ess m ea n w h i l e i n c r easi ng th e m ic r oscope

, ,

ge n e ra lly r evea l s no t o n l y swa rm s of bac t e r ia bu t a l so n u m e r ous ,

inf uso ria A t th e sa m e t i m e th e co l o r of th e l iquid h as deep

.

o ned i t b egin s to appe a r t u r bid a scu m fo r m s o n th e s u rface

, , ,

a nd th e odo r of h ay w h ic h was p r ese nt a t th e ou t se t is r ep l aced

, ,

by th e l ess ag r eea bl e o do rs of pu trefac tion T h e sim p l e ex .

rim ent of b r i ngi n

p e
g to ge th e r h a y a n d wa t e r h as in fac t set i n , ,

m o t io n a co m p lica ted se ries of p hy sica l c h e m ica l and bio l og ica l

, ,

p h e no m e n a .

Th e Composi t i on of a Hay I n fus i on A h ay i nfusio n consist s

.

o f t wo p rin cipa l co n st i t ue n ts h ay a n d wa t e r Bu t nei th e r of

,
.

th ese is c h e m ica lly pu re H ay is o n l y d ried grass w h ic h fo r

.

w ee ks a n d eve n m o n th s was e x posed i n th e fi e l d t o wi nd and

C ove r ed with th e l a tt e r —
, ,

d u st o f t e n th e pu l ve r ized m ud of

road s a n d r oad s id e poo l s —h ay is r ic hl y l ade n wi th d r ied bac t e r ia

.

a nd o th e r m ic ro o r ga n i s m s w h ile wa t e r suc h as is o r di na ri l y
;
-

draw n fr o m a tap freque ntl y co n ta i n s n o t onl y an abunda nce of

,

f ree o xy ge n an d va r ious sa lt s in so l u t io n bu t als o n u m e rous bac ,

t e ria i nfuso r ia a l gae di a t o m s and o th e r m ic r o org

, , , ,
a msms m -

s u spe n sio n I n th e m aki ng of a h ay i n fusion th e r efo r e n u m e r

.
-

, ,

o us fac t o r s co ope ra t e a n d a se ries of co m p licat ed r eac t ion s

,

fo ll ow on e a n o th e r in rapid suc cessio n A t th e s ta rt both .

2 01
202 A HA Y I N F USI ON .

h ay and wat e r ar e in s ta t e of co m pa r a t ive r es t o r e uilib

1.
q
rium ,
but upo n b ri ngi ng th e m t oge t h e r ac tio n a n d r eac tion
begin .
Fi rs t th e du st o n th e h ay is we tt ed and soaked
, ,

a n d any m ic r o o r gan i sm s i n i t o r ad h e ri n g t o th e h ay a re se t free

-

and floa t i n th e wa t e r ; n ex t th e wat e r fi nd s i t s way i n to the,

s t e m s a n d l eave s of th e h ay cau s i ng th e m t o swe ll a n d r esu m e

,

th ei r o rigi na l fo rm A t th e sa m e t im e va r iou s so l u bl e cons titu

. .

e nts of th e d ead gr as s s uc h as sal ts , suga rs and s o m e nitrog

, ,

eno us subs ta n ces difl use ou t wa r d i nt o th e wa t e r w hile f ro m

,
’
,

suc h ce ll s as h ave bee n c r u sh ed o r b r oke n ope n du r i n g dr ying

o r h and l i ng so l id p r o teid o r s t a rc hy su bs ta n ces m ay pass ou t and
,

m i ng l e wi th th e wa t e r T h e se si m pl e p hy sical r eac t io ns obv i

.

ousl i n vo l ve a dis t u rba n ce o f th e che m ical eq uilibri u m of th e

y
wat e r O rigi nall y ab l e to suppo rt o n l y a l i m i t ed a m o un t of l ife
.

( suc h as e x is t s in d r i nki ng wa t e rs
) i t is n ow a soi l e n-
r ic h ed ,

by w h a t i t h as gai ned f r o m th e b ay The bac te ria ex tr e m e ly .

,

se n sit ive t o va ria t io n s in th ei r e n viro nm e n t an d e s pecially to ,

th ei r food supp ly imm edia t e l y p r oceed t o m u l t ip l y e no rm ousl y

-

, ,

so th at a bio l ogica l r eac tio n fo ll ows c l ose l y o n th e h ee l s of the

c h e m ica l c h ange Bu t as a r e su lt of th ei r m e ta bo lic ac t ivity th e
.

bac t e r ia se t up e xt e ns ive c h e m ica l c h a n ges w h ic h in t h ei r t u rn ,

i nvo l ve ph y sica l dis t u r ba n ces Fo r e x a m p l e th e di sso l ved o xy .

,

g en wi th w h ic h th e l iquid was sa t u r a ted soo n disappea r s so th at ,

m o re oxy ge n m us t th e r efo r e diffuse i nt o th e l iquid fr o m th e

, ,

a t m o sp h e r e C a r bo nic acid is ge ne ra t ed in excess a n d so m e

.
,

m ay pass ou twa rds t o th e ai r A l so as a resu lt of th e vi ta l .

,

ac t ivi ty of th e m ic r o o r ga n i sm s th e te m pe r a t u r e of th e i nfusio n
-

m ay rise a f r ac tion of a degr ee above th a t of th e su rr ou n d

inga t m osp h e r e .

We ar e co n ce r ned h oweve r c h iefl y wi th th e biol ogica l

, ,

r esu lt s I n con seque n ce of th e ex h au s t io n o f t h e oxy gen supp l y

.

i n th e l owe r pa rt s of th e l iquid m any o f th e bact e r ia w h ic h ,

r equi re abu n dan t ox y ge n fo r th ei r g r ow t h (aerobes ) fi nd

th ei r way t o th e su r fac e w h e r e so m e pass i n t o a k i nd of ,

r est i ng s t age (z oog lcea ) a n d fo r m a scu m o r skin on .

th e su rface of th e l iquid O th e r s fo r w h ic h f r ee ox y ge n is no t
.
,

n ece s sa r y o r t o w h ic h i t is eve n p rejudicia l a n a erobes l ive

( ) a n d ,

thr ive in th e d eepe r parts of th e beake r Bu t m ea n t i m e an .

, ,
 PH Y SI OL O GI CA L C YCL E IN THE I N FU I S ON . 2 03

o th e r p h e no m en o n h as occu rr ed Th e i nfuso r ia o r igi nally few .

,

in n u m be r fi n din g th e c o n ditio n s favo r abl e h ave m u lt ip l ied

, ,

e no r m ou sl y a n d afte r a day o r t wo m ay be see n dart i ng i n and

, l

ou t a m o ng th e bac t e r ia e speciall y n ea r th e su rface and feedi ng

, ,

upo n th e m Am o ng th e i nfu so r ia h oweve r ar e so m e w h ic h

.
, ,

feed upo n th ei r fe llow s so th a t we soon h ave th e h e rbivo rous ,

in fu s o ria pu r s ued b y carmv o rous fo r m s th e w h o l e scene i ll us ,

tratingin o ne fi e l d o f th e m ic r o scope th a t s tr uggl e fo r exist e n ce

w hic h is o ne o f th e fu n da m e n ta l fac t s of bio l ogy .

O bvio us l y th is c h ai n of l ife is n o s tro nge r th an i t s weake st

,

pa rt T h e h ay is th e sou r ce of th e food supp ly fo r all th ese

.
-

fo r m s a n d th i s su pp l y m us t eve nt uall y becom e e xh aus t ed

, .

W h e n th is h ap pe n s th e bac t e r ia cea se t o m u lt ip l y th e h erbiv o

, ,

rous i n fuso ria w h ic h depen d upo n th e m pe r ish o r pass i n t o a rest

ings e crth e ca rn ivo rous i nfuso ria l ikewi se st ar ve and all th e
, ,

bio l og ica l p h e no m e n a m us t ei th e r co m e t o an end o r ch ange

th e ir c h a r ac te r .

Up t o th is poi nt th e ac t io n is pu re l y destr uc t ive Bu t soone r .

o r l a te r m ic r oscopic g ree n p l a nt s may appea r on th e sce n e ,

P roto eocc us i t m ay be o r i ts al lies and a constr uc t ive ac t io n

‘

, , ,

begi n th e was t e p r oduc ts o f th e a ni m a l s and of th e bac t e ria be

,

i ng r ebui lt b y th e gr ee n p l an ts i nt o co m p l ex o rganic m att e r B y .

this t i m e also th e disso l ved o r ga nic m att e r wi ll h ave bee n

, ,

la rge l y e x tr ac ted fr o m th e l iquid th e bac t e ria fo r th e m ost ,

pa rt devoured by th e i n fuso r ia and th e l att e r m ay m o re o r l e ss ,

co m p l e t e l y h ave give n way to l a rge r fo r m s — t o rh izopod s rot i ,

fe r s s m a ll wo r m s and th e l ike T h e pu trefy i ng i nfu sio n h as

, ,
.

r u n i t s cou rse and th e o rdi na ry ba l a ce of at u r e h as bee

,
n n n

r es to red .

T h e n cefo rwa r d a n app r oxi m at e equi l ib r iu m is m ai nt ai ned .

T h e gr ee n p l ants bui l d co m pl ex o rgan ic matt e r and st ore up

th e e n e rgy of l ight T h e an i m al feed upo th e pl a t o r
s n
.
n s ,

upo n o ne a no th e r b r eak dow n th e co m p l ex m att e r a n d di i

ss
,
,

pa t e e n e r gy T h e eve r p r ese nt bact e r ia b reak dow n

.
-
a ll th e
r efuse ex tr ac t so l ub l e o rga nic m a tt e r fr o m th e wat e r d aco m
,
,

d en d
pose th e dead bodies of th e a ni m al s o r p l a t n s a n i n th e ,
,

i i nfuso r ia T he
i t may be th e m se l ves fall vic t i m s t o devou r ng
,
.

p hysio l ogica l cy c l e is co m p l e t e .
2 04 A HA Y I N F USI ON .

A h ay in fusio n th us affo r ds i n m in ia t ure a pic t u r e of th e l iv

i ng wo rl d T h e g ree n p l ant s a re co nstruc tive an d i n th e sun
.
,

lig ht bui l d up m a tt e r s ric h i n po t e n tia l e ne rg

y Th ese as food:
.

suppo rt co l o rl e ss p l a nts (s uc h as bac te r ia) o r a ni m als O n th ese

.
,

agai n h e rbivo rou s a n d ca r nivo r ou s an i m al s feed ; a n d so

, ,

wo rl d at l a rge as i n th e h ay i n fu sio n o mn ivo r ous as we ll at

, ,

carn ivo r ous anim al s i n th e l o ng r u n feed upo n h e r bivo rou s

a ni ma ls a n d th e l a tte r u po n p l an ts—e i th e r co l o rl ess o r gree n
, ,

w h ic h th us s tan d as th e bul war k be t wee n a nim als a n d s tarvatio n .

 A PPE N DI X .

S U GG E S T I O N S FOR L AB O R AT O R Y S TUDI E S AND

D E M O N S T R ATI O N S .

Th e L abo ra t o ry
D i rec t io ns in G e ne ral B io l ogy publish ed ,

a n d cop yr ig ht ed b y P r of E A A n d r ews of Joh ns Hopkin s

. . .

Un ive rs i t y wi ll be foun d ex tr e m e ly usefu l and p ra ctica l A l so

, .

t he fo ll owi ng : H u xle y an d Ma rt i n s P r ac t ica l B io l ogy (Howes

’

a nd S co tt
) a,
n d th e acco m pa ny i n g A tl as of B io l ogy ”
b y H owes ; ,

M a rs h a ll and H urs t s P rac t ica l Zool ogy

“’ ”
C olt on s “
P rac t ica l
,
’

” “
Zoifl o g m ”
Dodge s E le “
’
y ,
B u p u s s I n ve rt eb ra t e Z o ol ogy ,
’

m enta r y P rac t ica l Bio l ogy ”

B r ooks s Handbook of I nve rt e
,
“ ’

”
b r a t e Zool ogy . A cco rdi ng t o ou r expe rience th e per iods fo r ,

th e cou r se s h ou l d be so arr a nged as t o affo r d l abo rat o ry wo r k

. a nd r ec i ta t io n s o r quizzes i n abou t th e p ropo rt ion s of three t o

tw o (f o r e xa m p l e thr ee pe riods of l abo ra t o ry wo r k and de m o n

,

s tref io n to t wo of quiz) fo r a h al f y ear

,
-
.

C HAP TE R N T R O DUCT OR Y )
I .
( I .

It co n ve nie n t t o give a t th e ou t se t o ne o r m o re p ract ical

is
l esso ns o n th e m ic roscope affo r di ng th e st udent an oppo r t u ni ty t o
,

lea rn i t s diffe r e n t pa rt s u se i ts adjust m e nt s t est th e m agnify i ng

, ,

p owe r of th e va r ious co m bi na t io ns e t c A good

,
objec . t fo r a
fir s t exa m i n a tio n is a h u m a n h ai r w h ic h se rves as a con venient
,

st a ndar d of siz e f o r co m pa r i s o n wi th o th e r th i ngs O th e r good .

objec t s a r e s ta rc h e s th e sca l es fr o m a bu tt e rfly s wi ng (ske t c h

,
’

u de r d ffe r e t powe r a d r op of m i l k o r b l ood n d powde r ed

la n i n s
) ,
a ,

carm m e o r ga m boge r ubbed up in wa t e r ( t o s h ow th e B r ow n ian

m ove m ent) T h e st ude nt sh ou l d co m par e th e sa m e object as

.

see n u n d e r th e sim p l e a n d th e co m pou d n m ic r oscope ( t o sh ow

205
2 06 A PPE N D I X .

r eve r sa l of th e im age in th e l att e r) and s h ould du ri ng th e cou rse

,

l ea rn t h e use of t h e ca m e ra l ucida ( A bbe s ca m e r a of Zeiss th e ’

, ,

best) T h e st age m ic r o m e te r m ay a l so be e xa mi ned a t th is t i m e

.
-

o r l a t e r and th e s t ude nt ta ught to p r epar e a s ca l e ( see A n d rew s)

,

by d rawi ng t h e l i ne s wi t h ca m e r a o n a ca r d un de r d iffe re nt
, ,

powe r s ( A 2 D ,
2 D +4 of Zei ss) an d labe lli ng eac h
, , ,

wi th th e n a m e s o f l e n se s and ac t ua l size of th e spaces as s ta t ed ,

o n th e m ic r o m e t e r .

P e nci l d rawi n g s h ou l d begi n as soo n as th e fir s t s peci m e n is

-

in focu s and s ke t c h es s h ou l d be m ade f r o m th e ve ry fi rs t e x e r cise

, ,

o nwa r d of eve ry t h i ng rea lly s t udied I t is abso l u te l y i ndis

,
.

pe nsab l e to keep a la bora to ry no te book w h ic h ought a t any tim e

-

t o give t a ngib l e evide n ce t h a t th e l abo ra t o ry s tu d y is bea ring

f r ui t ; an d in th e ve ry fi rs t l abo rat o ry exe r cise a begi nn i ng s h ould
b e m ade i n th is di r ec t io n .

The p re l i m i na ry m ic r oscopy of o n e o r t wo l abo ra t o ry pe ri

ods co rre spo ndi ng t o th e t i m e s pe nt in co nfe r e nce s u po n th e first
,

c h ap t e r of t h e t ex t book l eads n a t u r ally up t o th e ea sy rni cro

-

,
«

scopica l st udie s r equi r ed in co n n ec tio n wi th t h e seco n d c hap te r .

C HAP T E R II .
( S T R UC T U R E or L I V I N G O RG AN I S MS .
)
T h e l abo r a t o ry wo r k m ay be m ade v e ry b rief an d si m pl e ,

an d t h e fac ts s h ow n l a r ge l y b y i ll u str at io n T h e p r i ncipal .

o r ga ns o f a p l a n t and of a l ive o r dissec t ed a nim al m ay be sh o w n

a nd so m e of t h e m o re obvi ous t i ss ue s poi n t ed ou t A frog u n de r .

a be ll gl ass and a flowe ri ng p l a nt (ge ran iu m ) in b l osso m p l aced

-

, ,

side by side 011 t h e de m o nstr a t io n ta b l e wi l l se rve to sugg -

es t
m at e rial s fo r th e l i st s of o rga n s and th e co m pariso ns ca ll ed fo r .

T h e ski n of a Ca lla l eaf is eas i ly s t ripped off and de m o n

strated t o th e n aked ey e as o n e fo r m o f t i ss ue I t m ay th e n be .

cu t up and di stribu t ed f o r mi cro sc O pic s t ud y a n d fo r p roof t h a t

i t is co m po sed of ce lls ( D u r i ng th i s p r ocess ai r is ap t to rep l ace
.

wa te r l ost b y evapo ra t io n and m u st be di sp l aced by al co h o l ,

,

w h ic h in t u r n m u s t be r e m oved b y wa te r ) .

Fo r a fir st m ic r oscopica l exa m in at io n o f t issue th e r e is n o

be tte r object th a n th e l eaf of a m o ss (a s pecie s h avi n g th i n b r oad
l eaves sh ou l d be c h ose n) o r a fe rn p ro th a ll iu m O th e r good .

objec ts a r e th i n se c t io n s of a po ta t o t ube r fro m j us t below th -

e
 LA B ORA TOR Y ST UDI E S AND DE M ON STRA TI ON S . 2 07

m f ( n ed wi th di l u t e iodi ne to sh ow n ucl ei and st arch

aee t a i s

grains ) an d fr og s o r n ewt s b l ood m ixed wi th no rm al salt so lu

,
’ ’
,

t io n an d exa m in ed ei th e r f r e s h o r sl ightly s tained wi th di l u t e

,

Th i n sec tio ns o f pi th (e l de r fr o m w h ic h th e air h as ,

bee n d isp l aced b y al co h o l give good pic t u r es of t i ssue com po sed

,

of e m p t y ce lls F r e s h o r al co h o l ic m u sc l e fr o m th e fr og s l eg
.
’
,

g yen tl t eased ou t s h ows m u s cu l a r t i ss ue to be co m posed of e l o n

,

ga t ed ce lls (fi b r e s) Fi na lly th e st uden t m ay p r ove th at h e

.
,

h imse lf is co m posed of ce ll s b y ge ntly sc r api ng th e i nside of his

lip o r ch ec k wi th a sca l pe l m ou n t i ng th e sc r api ngs on a sl ide ,

a n d aft e r addi ng a dr op of Delafi eld s h aem a t oxy l in cove r i ng ’

, ,

a n d exa m i n i ng in th e usua l way .

To sh ow th e l ife l ess m a tt e r in l ivi ng t i ss ue i t suffices t o ex

’
a min e fr og s bl ood o r h u m an b l ood ; sec t ions of po tat oes es ,

c iall if l ig htl y s tai n ed wi th iodi n e ; sec t ion s of ge r a ni u m st e m s

p e y
( P e la rg o n lum ) w h ic h usua ll
,
y s h ow c r y s ta l s in so m e of th e m o r e

pe rip h e ra l ce lls ; ca rt i l age s ta i ned wi th iodi ne in w h ic h th e l ife

, ,

l es s m a tr i x r e m ai n s u nco l o red ; or p repa red sec t ions of bone in ,

w h ic h th e s paces o nce fill ed by th e l ivi ng ce ll s a r e new b l ack and

o paque be in g fill ed w i th du st in th e g ri ndi ng o r wi th ai r
, ,
.

C HA P TER III .
( PRO T O P L A SM AND TH E C E L L )
.

N ak ed ey e E xa min a t i on of Pr ot opla sm
-
A d rop of p r o t o .

p l as m is rea dily ob tai ned f r o m o n e of th e l ong (i nt e rnodal ) ce ll s

of _ ZVltd la af te r r e m ovi ng th e supe rfluou s wat e r and snippi ng off
'

o ne end o f t h e ce ll wi th sci sso rs Th e ce ll co ll apse s an d th e .

d r op fo rm s a t th e l owe r (cu t) end I t m ay be tr ansfe rr ed t o a

.
.

( dry) s l ide a n d t es t ed f o r i t s viscidi

. t y b y t ouc h i n g i t wi th a

n eed l e M ic roscopica ll y i t is i nstruc tive c h iefl y by i t s l ack of

.

m a rked str uc t u r e .

Th e P a rt s of th e Cel l T h e s tr uc t u.r e of th e ce ll i s beau t i

fu ll y sh ow n i n p r ope rl y s tai ned and m ou nt ed p repa rat ion s of un

N elella
e
f t r i l ized s ta r fi sh or sea
- u r c h i n eg g s o-
r of apica l bud s of ,
.

I f th e s e a re not avai l ab l e po ta t o ce ll s o r cart i l age cell s do ver y -

we l l ; o r sec t ion s o f epi th eliu m gl ands e t c m ay be sh own , ,

.
,
.

’ ’
The c l ass m ay a l so m ou n t a nd d r aw f r og s o r n ew t s b l ood

e ll s p r epa r ed an d doub l e s t a in ed as fo ll ow s T h e b l ood is sp r ead

c
- .

,
208 A PPE N DI X .

o ut eve n l y on a s l ide a n d d r ied cau tious l y ove r a fla m e S tain .

wi t h hie mato xylin fo r thr ee minu t es ; was h th o r oughl y wi th wa te r ,

a dd s t r o ng aqueous so l u t io n of eosi n allow to s t an d o n e m i n u te ; ,

was h th is t i m e ve ry r apid l y re m ove th e e x cess of wate r q u ickly,

w i th fi lter pape r p r essed dow n ove r th e w h o l e s l ide ; d r y r apidly

-

a nd e x a m i n e wi t h l ow powe r I f successfu l m ou nt in ba l sam ; if

.

t he s peci m e n is no t pi nk e n oug h add m o r e eos in and was h still

m o re r apidl y th a n befo r e In good s peci m e n s th e ce l ls k eep .

t h ei r fo rm pe r fec tl y th e c y top l as m is b rig ht pi nk a n d th e n uc l eo

, ,

p l as m is l ig ht pu r p l e .

E pide r m i s f ro m y o u ngl eaves o f h o t h ous e li l ies A f rican -

l ily ,
C h i n ese l i l y an d e s pecia ll y lil y of th e va lley) yields
,
- - -

c e ll s s h owi ng fi n e ly th e ce ll wa ll n uc l eu s a n d ( in favo rable -

, ,

c ase s cy t op l as m I f s t ai n ed wi t h ace tic acid a nd m e th y l gree n

) .
-

t h e n uc l ei a re h ighly co l o r ed ; wi th Delafi eld s haem a to xyli n th e ’

c y t op l as m is m o r e easily s ee n .

Cell div ision s or Cleav ag

-
e are ea s i l y o b se r ved i n segm e n tin g

o v a o r in f r e s h s peci m e n s of P ro to c oc cus ( F lew coc oons de

)
-

tec h ed fro m m oi s t e n ed pieces of ba r k w h ic h bea r th ese a lga .

( S ee p .

S t a ge s in th e c l eav age o f th e o v u m m ay be see n in th e seg

m en ting eggs of f res h wa t e r s nai ls ( P hy sa P la norbis ) w h ic h
-

a re eas i l y p r ocu r ed a t a l m o s t any t i me by k eepi n

g th e a n im a l s i n

a qua ria The o l d egg m ass e s s h ou l d be r e m oved so as t o e n su re

.
-

th e eggs bei ng f re sh O r a supp l y of p rese r ved segm e n ting eggs

.

st a r fis h r h m y t r m n tr t th rl y
( -
sea u ,
c i n-

) a be kep fo de o s a in g e ea
s ta ge s .

Pr ot opla sm i n Mot i on T h e be s t i ntr oduc t io n to p r otop l asm

.

in m o t io n is affo rded b y a s upe rfi cia l exa m i na t io n of A moeba

( fo r p r ocu ri ng A moeba see above C h ap te r X I I ) I f A mwba is , .

n o t avai l ab l e y ou n g l ivin g t ips o f N itella o r Cha ra m ay be u sed .

A n ac ha ri s a nd Trades ca n tia a re usefu l and oft e n ve ry beau tiful , ,

bu t l ess eas y t o m an age as a r u l e I n m ou n t i n g N i tella o r

, .

Cha ra ca r e m us t be take n n o t to c r us h th e ce ll s and as fa r as ,

possib l e pa l e f re sh s peci m e ns r a th e r th a n da r ke r a n d o l de r o n es
s h ou l d be c h ose n I f A na c ha ris is t o be s t udied th e y ou ngest
.

l eave s sh ou l d be se l ec ted f ro m th e buddi ng e nd s an d n o t as is , ,

s o m e t i m es r eco mm e nded l ea ves w h ic h a r e beco m i n g y e ll ow

, .

The m ove m e n t in t h e ce ll s of A na c ha r is l eav es of te n begi ns

 L A B O RA TO R Y ST UDI E S AN D DE M O N STRA TI ON S . 2 09

o n l y aft e r th e l eaf h as b ee n m ou n ted fo r a h al f h ou r or m o r e ; -

but w h e n o n ce es tab l i s h ed af fo r ds o n e of th e m o st beau t ifu l and

st ri ki ng exam p l e s of p r o t op l as m ic m o t io n I f Tra desca n tia is to .

be used ca r e m us t be take n to h ave if po ssib l e flowe rs just open

, , ,

o r ope ning T h e m o rn i ng is th e refo r e p refe rab l e fo r wo r k o n

.

th is p l ant H ig h powe r s a r e n ecessa ry

. .

In all th ese fo r m s th e m ove m en t s m ay oft e n be st i m u l ated b y

p l acin g a la m p n ea r th e m ic roscope or by ca u tiously war m i ng
th e sli de ove r th e l a m p c h i m n ey C iliary ac t ion is easi ly sh ow n
-
.

i n bi ts o f th e gi ll s take n f ro m fr es h c l am s m usse l s o r oy st e r s or , , ,

in ce ll s sc raped f r o m th e i nside o f th e fr og s (e soph agus A ’

.

str iki n g de m o n str at io n is easi l y give n by sl i tt i ng open a fr og s ’

( )
’
o r t u rtl e s oesop h agus l e n gth wise p in n i n g o u t fla t m
,
ois t e n i n g ,

wi th n o rm a l sa lt so l u t io n and p l aci ng t i ny bi t s of m oi st ened cork

,

o n th e su r fa ce T h e p rog
. r essive m ove m ent of th e co rk bit s is -

th e n ve ry obviou s M uscu la r co n tract i l i ty is easi ly sh own by

.

re m ovin g th e ski n f r o m a f r og s l eg di ssec t i ng ou t th e sciat i c ’

,

n e rve cu tt i n g its uppe r end and th e n st i m u l a t i ng th e l owe r end

, , ,

if po ss ibl e b y co n tac t wi th a pai r of e l ec tr ode s o th er wi se b y

, ,

pin c h in g i t wi th fo r ceps I f th e necessa ry appa r at us is avai l ab le

.

th e regu l a r m usc l e n e rv e p r epa r at io n m ay be sh own (see Fost e r

-

“
an d L a n gl ey s P r ac t ica l
’

Foo d st uffs Con t a i n E n er g

-

y T h is m a y be s.h ow n i
( de m
n

onst ratio ns
) b y s rin k ing fi nely
p p o wdered an d tho ro u hl
gy
dried st a r c h s uga r o r flou r upo n a fir e o r upon a p l at i nu m di s h
, , ,

o r piece o f foi l h ea ted t o r edn ess ove r a sm all flam e O i l s and .

d ri ed an d powde r ed a l bu m e n ( p r o t eid ) m ay be si m i l arly m ade t o

bu r n wi th alm o s t e x p l o sive v io l e n ce if app l ied in a st at e of fi ne
divi sio n in p r ese n ce of ai r .

Th e Ch emi cal B as i s (a ) P roteids ; Coag .

u la tion ; R ig or M o r

tic ; R ig or Ca lori e W h i te .o f egg m ay be s h

-
ow n -

( in de m o n s tra
t io n) an d m ade to coagulat e in a t e st t ube h u ng down i nt o a -

b eake r of wa t e r u n de r w h ic h is pu t a fla m e A th e r m o m e t e r in .

th e t e st t ube m ay be read off f r o m t i m e t o t im e as th e experi

-

m e n t adva nce s u nt i l fi n a ll y co agu l a t io n begi ns w h en th e t empe r

,
,

a t u r e is n o t ed The dea th st iffe i g ( g

.
n n r i or m- or ti s
) com e s o n

ve r y quick l y in f rogs ki ll ed wi th c hl o rofo r m H eat st i e i g ff n n .

-

r i or ca lor ic is we ll s h ow n b y i m m e r si g l eg of a decap i
( g )
“
n o n e

t a ted f r og in a b eake r o f wa t e r a t 4 0 C T h e o th e r l eg °
re .

s
2 10 A PPE NDI X .

m ai ns n o r m a l and aff o r ds a va l uab l e m ea n s of co m pariso n I t .

is n o t wo rth w h ile t o m ake m a ny c h e m ica l t es ts of p r o t eids a t

th is poi n t .

( )
b C a rbohy dra tes A usefu. l de m o n s tr a t io n m ay be m ade

o f va rious s t a rc h es s uga rs an d g ly coge n T h e iodi ne t e s t may -

,
.
,

be app l ied if des i r ed I f t i m e all ows th e m ic roscopical appea r

.
,

ance of po ta t o s ta r c h co r n st a r c h Be r m uda a rr ow r oo t e tc
- -

may be dwe lt u po n in th e l abo ra t o ry wo r k C e llu l ose is we ll

, , .
,

-
.

s h o wn in fi lte r pape r o r abso r be n t co tt o n

-
.

c Fa ts A de m o ns t r a t io n of a ni ma l fa t s a n d vege ta bl e oi l s
() .

m ay be m ade if t 1 m e a ll ow s T h ey m ay be exa m i ned microsc op

.

ically i n a d r op of m i l k i n a n a rt ifi cia l e m u l sio n m ade by sh ak

,

i n g up swee t oi l in di l u t e w h i t e o f egg o r i n f r es h fa tty t issue

- -

r m s u bcu t a n eou s t issue o f m ouse r t s of f r og I t

( f o o fa b odie , ) is -
.

h a rd l y wo rt h w h i l e to exa m i ne th e se s ubst a n ce s c h e m ica lly bu t ,

a few si m p l e t e s ts m ay be app l ied if de si r ed .

Dialy s i s
. A de m o nstr a t io n of di a ly s i s is easi ly m ade by in
ve rt i ng a b roke n t e s t t ube t y i ng th e m e m b ra n e ove r t h e fla ring
-

e nd fill i ng th e t ube to a m a r ked poi n t wi th s tr o ng sa l t o r g l u

,

cose sol u t io n a n d i mm e rs i ng i t i n a beake r of dist i ll ed wa t e r

, .

A f t e r an h ou r o r so th e fl uid wi ll be fou n d t o h ave r ise n i n th e

t es t t ube agai n st g ravi ty
-
.

Temper a t u r e a nd Pr otopl asm T h e p r ofou nd i nflue nce of

.

t e m pe rat u re o n p rot op l as m is we ll s h ow n by th e f r og s h ea rt
'

D ecapi ta te a fr og a n d des tr oy t h e spi na l co r d E x p se th e . o

h ea rt and cou nt th e bea ts a t t h e r oo m t e m pe ra t u re T h e n pou r .

u po n th e h ea rt iced n o r m a l sa lt so l u t io n A gai n cou n t th e bea ts . .

°
N ex t pou r u po n i t n o r m a l sa lt so l u t io n h ea t ed t o 3 5 C T he .

n mnber of bea ts wi ll fo ll ow t h e fa ll a n d r ise of t e m pe ra t u r e .

C HA P T E RS I V To V III .
(THE E A R T H W ORM ) .

L a rg
e ea rt hworms m ust be sa t isfac to ry r e su lts can
us ed or

n o t be expec t ed P a in s sh ou l d th e r efo r e be t ake n to p r ocu r e

.

th e l a rge L ter res tris ( no t th e co mm o n A llo lobop hora mu cosa )

.

w h ic h is r eadi l y r ecog nizab l e by th e fl atte n ed pos t e r io r end .

Th is species is n o t eve r y w h e r e co mm o n ; h e n ce a supp l y sh ould

be p r ocu r ed and kep t in a coo l p l ace i n ba rre l s h al f fu ll of ea rth ,

o n th e su rface of w h ic h is p l aced a qua n t i ty of m oss T h ey will .

 LA B ORA TO R Y S T UD I E S A N D DE M N O S TRA TI O N S . 2 11

th u s l ive fo r m o nth s L t rres tris m ay be ob t ained m gr eat

. . e

n u m be r s be t wee n A p ri l a n d N ovem be r by sea rching fo r th e m ,

a t n ight wi th a l ant e rn i n l ocal it ie s w h e re n u m e rous cast ings

s h ow th e m t o a b ou n d
( a r a th e r h eav y bu t ric h soi l wi ll be fou n d
m o s t p roduc t ive) T h ey wi ll th en be fou n d ex tended fr o m th ei r
.

bu rr ows l y i ng o n th e s u rface o f th e g rou nd and m ay be seized

,
,

wi th th e fi n ge r s C o n side rab l e dex t e r i ty is needed and i t is

.

n ecessa ry t o tr ead ve r y softl y o r th e wo r m s t ake al a rm and in

s ta n tl
y wi th d r a w i n t o th ei r bu rr ow s .

Fo r di sse c t io n f r e s h s peci m e ns a re fa r p r efe r ab l e for m ost

pu r poses th oug h p rop erly p re s e r ved o ne s answe r th e pu rpose
, .

F r e s h speci m e ns s h ou l d be n ea rl y ki ll ed by bei ng p l aced fo r a

s h o rt t i m e ( abou t fi ve m i nu t e s) i n 7 0% al coh ol and th e n str et ch ed,

o ut to th ei r u t m o s t ex t e nt in 5 0 % al co h ol i n a dissect i ng pan -

th e two e nd s bei ng fas t e n ed by pi ns T h ey sh ou l d th en be at

.

o n ce cu t ope n a l o ng th e m idd l e do r sa l l i n e wi th sci sso r s th e

,

flaps pi nn ed ou t and th e di ss ec t io n cont i nued u nde r th e 5 0%

,

a l co h o l
.
( T h e y m u s t be comp letely cove r ed wi th th e l iquid ) .

B y this m e th od th e m i n u t e s t de t ai l s o f struc t u r e m ay be ob
s e r ved ,
an d m a ny of th e di ssec t io ns sh ou l d be don e u n de r a

wa tc h m ake r s l e ns
’
.

Fo r p r ese rva t io n ( eve ry de tai l of w h ic h sh oul d be att ended

to ) a n u m be r o f l ivi ng wo r ms a r e p l ac ed in a b road ve sse l fill ed
t o a dep th o f abou t an i nc h wi th wat e r A l i ttl e a l coh o l is th e n
.

cau tious ly d r opped o n th e su rface of th e wate r at i nt e r val s u nt i l

th e wo rm s a r e stupefi ed and beco m e pe r fec t ly m ot ionl e ss an d r e
laxed (th is m ay r equi r e an h ou r o r t wo) Th ey a r e th en trans
.

fe rr ed to a l a r ge s h a ll o w vesse l co nt ai ning ju st e nough 5 0%

a l co h o l t o cove r th e m and a r e ca r efu lly straighte ned out and
,

a rr a n ged side by side A ft e r an h ou r th e weak al coh ol is re

.

p l aced by s tro nge r w hic h sh ou l d be c h anged once o r twice

a t i n te r va l s o f a few h ou r s ; th ey ar e fi nally p l aced in 9 0?
al co h o l w hi c h s h ou l d be libera ll u sed T h e tr oub l e de m an de d
, y . .

b y th i s m e th od wi ll be fu ll y r epaid by th e r e su lt s Th e wo rms .

s h oul d be qui t e straight fu ll y ex t ended and p l u m p and th ey

, , ,

m ay be used ei th e r fo r di ssec t io n or f or m ic roscopic study .

Fo r th e pu rposes o f sec t io n cu tt i ng wo rm s sh ou l d be ca r efu lly

-

wa s h ed an d p l ac ed in a m ois t ve sse l co ntai ni ng p l e nty of wet

h lter pape r t o rn i n t o s hr eds
-
T h e wo r m s wi ll devou r th e paper,
.
212 A PP E N D I X .

w h ic h sh oul d be c h a n ged seve ra l t i m es u n t i l th e pape r is voided

,

pe rfec tl y c l ean The wo rm s a r e th e n p r ese r v ed in th e o rdin a ry

.

way and w h e n p r ope rly h ar de n ed a r e cu t i n to s h o rt pieces

, ,

s ta i ned wi th bo r ax ca r m i n e i m bedded in pa raffi n an d cu t in to

-

, ,

sec t ions wi th th e m ic ro to m e
Th e l ivi n g wo rm s sh ou l d fir s t be ob se rv ed —th ei r sha pe
.

m ove m ent s be h av io r to s t i m u l i pu l sa tio n of th e do rsal v essel

, ,

( t i m e th e pu lse and v a r y th e ra t e b y te m pe r a t u r e c h a n ges ) .

We ll p rese r ved s peci m e ns sh ou l d th e n be ca refu lly s t udi ed fo r

-

th e ex t e r na l c h a r ac t e rs ( d r aw thr oug h th e fi n ge rs t o fee l th e se tae) .

( S ke t c h ) O b se r ve ope ni n gs T h e n ep hr idial ope n i n gs ca nn o t be

.

see n bu t if p re se r ved wo r ms be soak ed so m e h ou r s i n wa te r and

,

th e cu t ic l e pee l ed o ff th e y m ay be c l ea rl y see n in t h i s A .

ge n e ra l di ssec t io n of a f re s h s peci me n s h ou l d n ow be m ad e ,

a nd th e po s i t io ns of th e l a r ge r o r ga ns st udied ( Make pa rt ial .

s ke t c h t o be fi ll ed ou t af te r wa rd s as in Fig
, ,
T h e a l im e n tary
.

c a nal a nd ci r cu l a to r y o r ga n s sh ou l d no w be ca r efu ll y s tudif xl .

E ve n th e s m a ll e s t of th e b l ood ve sse l s m ay easi l y be wo r ked o ut

-

u nde r th e l e ns b y u s i ng fr e s h speci m e ns ( ki ll ed in 7 05‘ a l co h o l

and af t e r wa r d s di s sec ted u n de r w a ter) an d ca r efu lly t u r n i ng as id e

th e a l i m e nta ry ca na l .

The a l i m e n t a ry ca na l s h ou l d aft e r wa r d s be cu t thr oug h be

h in d th e gizza rd a n d g r adua ll y dissec ted away i n f r o nt e x pos in g ,

t h e ne r ve co rd and th e rep r oduc tive o r ga ns ( was h away dirt wi t h

-

a pi pe t t e) N o g rea t diffi cu lt y sh ou l d be fou nd i n m aki ng ou t any

.

o f t h e pa r t s excep t i ng th e t e st es T h ese a re diffi cu l t t o fi nd in

,
.

m a t u re wo r m s bu t may be fou n d w i th ease i n th ose w hic h h ave

,

n o m edia n se m i na l vesic l es ( u s u all y th e cas e wi th speci m e n s h av

ingn o c l i te ll u m) .

Th e co n te n ts of th e se m i na l r ecep t ac l e s and ve s ic l es f r o m a
f r e s h wo r m sh ou l d be e xa m i ned wi t h th e m ic r oscope R e m ove .

a n ova ry wi th fo r cep s and s m a ll cu rved s ci ss o rs m n t n ter

( ) ou i wa , ,

and s t ud y t ai ed i n a l u m c r m e an d m oun ted in ba l sam

( S
. n a i n -

th e ovary is a bea u t ifu l objec t ) T h e st ude nt s h ou l d a l so re

.

m ove a f re s h n ep hr idia l fu n n e l and pa rt of a n ephr idiu m and ,

s t ud y wi th th e m ic r oscope h s m ay h ave t o be s h ow n by th e
( T .i
de m o nstrato r bu t sh ou l d n eve r be o m i tt ed as t h e ci l iar y ac t io n
, ,

is o ne of th e m os t s tr iki ng thin gs to s e e ) A ca refu l di ssec t io n .

of t h e a n te r io r p art of th e n e r vo us sy st e m sh ou l d als o be m ade .

 AND D E M ON STRA TI ON S . 21 3

s t udy of m ic r oscopica l sec t io ns

p r epa r ed sec t io ns sh ou l d be kep t
v en

deve l op m e n t is too diffi cu lt to st udy bu t ,

st ratio n s m ay be give n by th ose w h o h ave

I n th e n eig h bo rh ood of P h i l adelp h ia egg

in g r ea t n u m be r s in o l d m an u re h eap s -

O ne en d of th e cap s u l e sh oul d be sl iced off

sc a l pe l and th e co nt e nt s d rawn o ut u n de r ,

e m ou t h ed pipe tt e
-

T h e m ass m ay th en be
.

in wa t e r u n de r a suppo rt ed cove r gl as s an d s t udied -

m ic r o scope Th e e m b ry o s m ay be p r ese r ved in

.

fluid and ei th e r s t udied w h o l e in th e p rese r vin g fluid

,

harde n ed in a l co h o l a n d cu t i n t o se r ie s of sec t io ns .

C HA PTE RS IX TO XI .
( TH E C OM M O N B R A K E .
)
E xcep t w h e n th e g r ou n d is f r ozen P teris m ay be dugup and
th e l abo ra t o ry i n a fr e s h s t a t e F r ond s m ay be .

in m id s u mm e r a n d co nside rab l y fr es h en ed ( by a
wa r m wa t e r) w h e n n eeded t o be u sed (in
ill u str a t e th e ae r ia l po rt io n of th e p l ant .

ined a t co n ve nie nce a nd kep t in weak

th e B ody . i ll u strat e th is o ne whole

To ,

po ss ib l e be a t h a nd fo r exam i na t ion
,
.

o un d po rt io ns m ay th e n be ske t c h ed

io ns B r a n c h e s roo t s and old l eaf st a l ks

.
, ,
-

ide nt i fi ed and ske t c h ed

,
.

the R hiz o me sh ou l d first be m ade ou t wi th

T h e l a t e r a l r idge s wi ll be de t ec t ed by th e cl ass ,

he a s ked t o d raw th e c r o ss sec t io n as see n wi th

-

Fo r th is p r e l im i n a ry wo rk eac h s t uden t sh ou l d
of rh izo m e t wo o r thr ee i nc h es in l eng t h (C ar e .

e d rawi ng has been co rr ec tly

dissec t ion wi th jack k n ife or -

i nfe r e nces as t o t h e c h a rac t e rs

o us p ul py woody
, , ,
2 14 A PPE N DI X .

fo r th e m os t pa rt easy but de m a nds m uc h t i m e I f t i m e

, , .

l ows c ross s ec t io ns of r oo t s m ay be m ad e a n d m ou nt ed in bal sa

,
-

T h ey a r e r eadi l y cu t i n pi t h S ec t io ns o f th e rh izo m e m ay
.

m ade f reeh a nd wi t h a r azo r o r be tt e r wi th a m ic ro t o m e : l , ,

th e o ld st e m s a r e exceedi ngl y h a r d a nd liab l e t o i n ju r e *

k n ive s .

The Fre ud or L eaf m ay be ob tai n ed in f r ui t in Ju l y a

A ugu st and p r ese rved in al co h o l F r o m i t sec t io ns of leafi .

m ay easi ly be go t by i m beddi n g i n pi th E pide r mis is ob tai1 .

wi t h so m e diffi cu lty ( by begi nn e rs) aft e r sc r api n g F r es h f e .

l eaves fr o m h o th ouses a n swe r th e pu r pose as we ll a re easie r ,

ge t and m o re a tt rac t ive R ea ll y good sec t io n s o f fe r n l eaves

,
.
-

n o t easy fo r begi n ne r s t o m ake Th ey s h ou l d be kep t o n h a ] .

S po ran gia m ay be ob ta i n ed i n abu n da n ce f r o m alcoht

s peci m e n s of P te r is o r upo n h o th ouse fe r n s eve n i n midwint
'

, ,

S o m e of th e m a ny species of P teris fou n d i n h o t h ou ses answ -

eve ry pu r po s e T h e th i n edge of a scal pel sl ipped u n de r th e 1

.

r i pe i ndu siu m re m ove s th e la tte r an d ge n e rall y al so l o ng ra n ks

,

s po r a n gia i n a ll s ta ges of deve l op m e n t In so m e spo ran gia spe .

m ay be fou n d S po ra ngia a n d s po res

.

bu t ca re m u s t be t ake n to se l ec t f r ui t do
'
-

o r t oo y ou n g .

Spr ou t i n gt h e S por es To o bta i .

ings po r e s and pro th allia f ree f ro m

foll owi ng p r oc edu r e : Fi ll s eve r al s m a ll fl owe r
bee n th o roughl y c l ea ned i ns ide an d o ut wi th ,

S te r i l ize th e w h o l e by baki ng i n a n ove n o r a h o t

S e t th e po ts i nt o l a rge ( po r ce l ai n) di sh es capab l e of 11
a nd keep th e bo tt o m of th ese di sh es cove r ed to th e

i nc h wi th wa t e r ; cove r th e po ts co m p l e t e l y wi th
A ft e r t we nty fo u r h ou rs o r afte r th e san d
-

co m e t h o r oughly we t i n side a n d o utsi de, ,

a nd the ou ts i des of th e po t s wi t h spo r e s

h ouses by sh aki ng fe rti l e fr o nds ove r w h i t

be take n t o ge t sp o res and n ,

week o r l o n ge r ( so m e t i m es
l ay e r of sa nd is r e m oved t o
ined fo r sp r ou t i ng spo r es .
 Fai l i n g th ese p r o th a ll ia m ay a l m os t a l ways
,
be fou n d i n fe rn
h ouses o n th e t ops o r sides of th e pot s and especially on
th e
,
m ois t ea rth u nde r th e be n c h es C ar e sh ou l d be t ake
n n ot t o
. ~

c o n fou n d p r o th a lli a wi th th e l ig ht e r
gree n and re l at ive ly coa rse
live rwo rt ( L u n u la ria oft e n fou n d i n h o th ou es
) s .

Th e Sex ua l Org a n s of Pr ot h allia Wi th good clea n

.
speci
m e n s th ese a r e easi ly foun d wi t h a rath e r l ow powe r
High e r .

powe rs a re n eeded t o make out de tai l s I f th e ar c h ego nia and .

Fert i liza t i on T h is is n o t easy t o ob se rve bu t th e att e m p t

.

,
m ay be m ade by exa m in i ng successive ly a n u m be r of ve ry fr esh
an d v igo r o us p r o th a ll ia i n di ff e r e n t s t ages T h ey m ust be .

m ou n t ed ca r efu ll y ( not fl ooded wi th wat er) an d spe r m at ozoids ,

are ge n e r a ll y m o r e ea s i l y fou n d swi mm i g abou t aft e r th e speci
n
m e n h as bee n m ou n t ed a littl e w h i l e .

E mb ry olog y E x cep t in i t s gene ra l feat u res th is is t oo dif

.

,
fi cult f or th e beginn e r H e m ay h oweve r obse r ve th e l at e r
.
, ,

s tages by st ud y i ng o l d p ro th a ll ia wi th th e y ou ng fe r n just ap
pea r i n g and y ou n g fe rns wi th th e o l d p ro th all ia st ill ad h e r ent
, .

Ch l or ophy ll a nd St a r ch Vigo r ous p r o th all ia affo rd exce ll e nt

.

exa m p l es of ce ll s bea r i n g c hl o r op h y ll bodies in w h ich stareh is

-

e asi l y de t ec te d . S om e of th e m a rgi n a l cell s sh ou l d be exam i ned

wi th th e high e st powe r attention beinggive n t o th e c hl oro
,
‘
p hy ll bodies a n d th ei r a rra nge m e nt In favo rab l e cases one m ay
-

obse rve th e opa que r od l ike or ova l gr ai ns i n side th e l att e r

-

a n d p r ove by r eage n t s th at th ey a r e sta r c h grai ns .

T h e s t ude n t s h ou l d al so exam i ne at th is poi nt th e l arge

, ,

c hr o m a t op h o res o f N itella w h ic h m ay be ob tai ned by p ressi ng

,

o ut a d r op o f th e co n t e n t s f r o m an i n t e r n oda l ce ll addi ng dilu te

,

i odi n e so l u t io n a n d exa m i n i n g wi th a h igh powe r I n favor

, .

a bl e cases as m a ny as a dozen s t a r c h g rai ns st ai n ed b l ue m ay be , ,

fou n d i n side a si n gl e e ll ip t ica l chlo rop hy ll body -

.
2 16 A PPE N DI X .

C HA PTE R XII) .
( tE B A
A M .

A mo eba is o n e of th e m os t cap r icious of a n im al s ,

a n d disappea ri ng wi th i n exp l ica b l e sudde nn ess a n d as ,

cann o t be fou n d a t th e t i m e w h e n n eede d u n l ess special p repar a ,

t io ns h ave bee n m ade in ad va n ce I t is n eve r safe to tru s t to .

c h a n ce fo r a supp ly of m a te r ia l I t is equall y u n sa fe to tru st to

.

th e m e th ods u sua ll y p r esc ribed Am os bee m ay h oweve r oft e n .

-

, ,

be p r ocu red in abu n da n ce a n d wi th t o l e r abl e ce rta i n ty as fo llow s :

A m o nth or si x weeks befo r e h a n d co ll ec t co nside rab l e quant i t ies
of wa t e r p l a nt s (e s pecia lly i Vitella o r Cha ra ) f r o m v a rious poo l s
-

o r sl ow di t c h e s wi t h a n abu n da n ce of sedi m e n t f r o m th e bo ttoin

, .

I t is i m po rta n t to se l ec t c l ea r quie t poo l s co n t ai ni n g an abu n

,

da nce o f o r ga nic m a t t e r ( suc h as des m ids dia t o m s etc in , ,

.
,

sedi1n ent) — 11 o t t e m po ra ry r ai n poo l s o r suc h as a r e

-

i no r ga nic m ud ( di rt was h ed i n by r ai n) T h e m a t e r ia l .

cu r ed s h ou l d be distr i bu t ed i n n u m e r ous ( 1 0 t o 2 0) o pe
di sh es ( ea rth e n wa re m i l k pa n s) a n d a ll owed to s ta n d
l abo ra t o ry in va r iou s p l ace s —
-

s o m e e x p osed to th e su n o th e r s i n ,

th e s h ade T h e co n t e n t s of m a ny pe rh aps a ll of th e vesse ls

es a n d swa r m wi th l ife—fi rst wi t h
.
, ,

wi ll u n de rgo pu tr efac t ive c h a ng

bac t e ria l a t e r wi th i nfu s ori a —
,
a n d wi ll th e n g r adually b eco m e
c l ea r agai n a in a h ay i n fu s io n T h e sedi m e nt s h ou l d no w be
s
-
.

exa m i n ed at i n t e r va ls a n d A mwbce a re al m os t ce rt ai n to
,

soo ne r o r l ate r i n o n e o r m o re of th e vesse l s

, .

A ra dio ed appea rs fir s t bu t th ese s h ou l d o n l y

.
,

fou nd im possib l e t o p rocu re A P ro teus w h ic h is fa .

,

and m o r e i n te r es t i ng E x pe rie nce wi ll s h ow th

.

poo l s al way s y ie l d a c r0p of A mwbw w h i l e o tl ,

W h e n o nce a p r oduc t ive sou r ce is fou n d all troub l e

I f possib l e a sedim e n t sh ou l d be se l ec ted th a t
A mwhw . I t is ve ry discou ragi n g fo r s t ude n ts t o
th ei r t i m e l ooking f o r th e a n i m a l s i n s t ead of a t
co ver -

g las ses s h ou l d be used an d th e m a te ria l ,

pipe tte fro m th e ve ry su r face of th e sedi m e nt

deepe r l ay e rs) W h e n firs t m ou n te d th e an i m als
.

trac t ed a n d o nl y beco m e fu lly e xt e n ded afte r a

,

s ke t c h es s h ou l d be m ade a t s ta ted in te rval s th e ,

p ro t opl as m ca refu ll y s t udied th e pu l se of ,

 LA BO RA TOR Y S T UDI E S A N D DE M N S O TRA TI ON S . 217

v acuo l e t i m ed ( va ry b y va ry i ng t e m pe r at u r e) and th e eff ect of ,

t appi ng th e cove r gl ass n o ted -

I t is p rac t ically u sel ess t o look

.

fo r fi ssio n fo r e ncy s t ed fo r m s o r f or th e ex t e rna l Ope ni ng o f th e

, ,

co n trac t i l e vacuo l e ; and th e ing ulfi n

g o f food o r pas si ng ou t
o f wa s t e m a tt e r s is r a re l y see n T h e fo r ma t io n of p seudopodia
.

s h ou l d be ca r efu ll y s t udied A f t e r exa m i n i ng th e l ivi ng ani m al s

.

th ey sh ould be k i ll ed and st ai ned wi th di l u t e iodi ne .

A rcellot is a l m o st a l way s an d D ifi ug ia som e t i m es fou n d

,
,

w it h A m aebe T h e se fo rm s m ay be exam i ned fo r co m parison

.
.

I t is de si r ab l e a l so t o co m pa re w h i t e b lood co r pusc l es w h ich -

may be O b tai ned ei t h e r by p ri cki ng th e fi nge r o r be tt er from a , ,

f rog o r n ew t A d rop of b l ood ,received upon a sl ightly wa rm ed

.

s l ide, s h ou l d be cov e red an d seal ed wi th oi l a rou nd th e edge of

th e cove r gl ass Th e w h i t e co rpu sc l e s a re a t first r ou nded bu t
-

s oo n begi n to s h ow c h a nge of fo r m N o co ntr ac t i l e vacuo l e no

( .
,

d iff e re nt ia t io n i nt o ec t op l as m and e n t op l as m oft e n n o n uc l eus ,

C H APT E R
XII (I N FUS OR I A ) . .

P a ra/meeeia a r e a l m o st ce rt ai n t o appea r in th e ea rl ie r stages

o f th e A moeba cu lt u r e s a n d in s i m i l a r deco m po si ng l iquid s o r
,

i nfu sio ns a nd t o e ns u re h avi ng th e m a l arge nu m be r of ve sse l s

,

an d ja r s co n t ai ni ng a n exce ss o f vege t ab l e m a tt e r sh ou l d be p re
pa red a m o nth o r m o re befo reh a nd Th ei r succe ssfu l st udy is .

ve ry easy i f th ey a re p r ocu red i n i e ry la rg e n u m bers th e wa t e r

( f

s h ou l d be m i l k y wi th th e m o t h e r wi se i t is p rac t icall y i m po ssib l e

) ,
.

Thr ee s l ide s of th e m sh ou l d be p r epa red an d set aside fo r a sh o rt

t i m e ( u nde r cove r p r efe rab l y in a m oi st c h a mbe r) to all ow th e
, ,

a n i m a ls t o beco m e q u i e t O n e sl ide sh ou l d co nt ai n si m p l y a
.

d r op of th e i n fu s o ria l w at e r ; a seco nd th e sa m e wi t h th e addi ,

t io n o f a l i ttl e powde r ed ca rm i ne ; t o th e th i rd add a d r op or t wo

o f an aq ueou s s o l ut io n o f c hl o ra l h y d r a t e ( m ade b y d r oppi ng a

c ry s ta l o r t wo i n t o a w a t c h gl a ss of wa t e r) -
Th e fir st sl ide .

s h ou l d be s t udied fir s t ; an d i t wi ll u sua ll y be fou n d th a t aft e r a

.

t im e th e a ni m al s c r owd abou t th e edge s of th e cove r oft e n ly i ng ,

n ea rl y o r qui t e st ill I f th i s is not th e case th e speci m ens para

.
,

ly z ed b y c hl o ra l m ay be st udied The ca r m i n e speci m ens will

.

s h ow beau t ifu l food vacuo l e s fill ed wi t h ca r m i ne ; a nd by ca r eful

-

s t ud y th e fo rm a t io n of th e v ac uo l e s m ay be O b se r ved .
2 18 A PPE N DI X .

T h e ge ne ra l s truc tu re s h ou l d be ca ref ully s tu died th e con ,

tr ac t i l e vacuo l es pa r t icu l a rl y e x a m i ned ( th e y a r e see n bes t in d yi ng

”

speci m e ns o r in t hose pa r a l y zed by c hl o r a l) and dividi n g o r co n ,

ug
j a tin
g i ndi v idua l s l ooked fo r ( th e y a r e O ft e n a b u n da nt
) Th e .

o n l y r ea ll y diffi cu l t poi n t is th e n uc leus w h ic h ca nno t be we ll ,

see n in th e l ivi ng a ni m a l I t m ay be c l ea rl y see n by m oun t ing

.

a d r op t o w h ic h a l i t t l e di l u t e iodi ne o r 2 56 ace t ic acid h as been

,

added T h e fo r m e r s h ows th e ci l ia we ll t h e la tt e r th e tric h o

.
,

cy s ts O s m ic acid a nd co rr osive s ub l i m a t e a l so give good p rese r

.

va t io n T h e i n t e rn a l c h a n ge s du ri ng fi ssio n and co njuga tio n

.

m us t be st udied i n p repa red s peci m e n s m ou n t ed in ba l sam S uch .

p repa ra t io ns a r e oft e n of g rea t beau t y and i nt e r est .

Vo rtieel/a m us t be soug ht fo r o n duc k weed o r -

o r o n floa t i ng s t ick s and th e l ike Zo iitha mn ion

, .
,

etc .
,
a re l iab l e t o appea r a t any t i m e i n th e aqua r ia A ll th ese .

fo r m s a r e eas i l y st udied C o njuga t io n is ve ry ra r e l y see n but

.
,

fi ssio n and m ot i l e fo r m s a re co mm o n T h e m ac r o n uc l eus is .

especia lly we ll sh ow n in dea d o r dy ing spec im e n s .

C HA PTE R X I V .
( P RO TOCOCC US .
)
P ro t ococcus ( P l
is fou n d i n ab u n da n ce o n
e u rococ eus
)
n o r t h e rly side of o l d tr ee s in m a ny pa rts of th e U ni ted S ta t es .

In case i t ca nno t be ob ta i n ed in a ny regio n i t m ay be p r ocu red ,

du r i ng 1 8 9 5 and 1 8 9 6 f r o m P r of S edgwick I ns ti tu te of
,
.
,

Tec h no l ogy B os t o n Mass by m ail T h e l abo ra t o ry wo r k wi th

, , , .
-

i t is too easy t o requ ir e co mm e nt S ee h oweve r A rth u r .

, , ,
“ ”
B a r n es C ou lte r s
’
P l a n t D issec t io n ( H e n ry H o lt C o .
,

N ew Y o rk) .

C HAP TE R X V .
(Y E A ST
)

Bak e r s b r ewe rs co m p ressed a n d d r i ed y eas t m ay be had

’
,
’
, ,

in th e m a r ke ts B r ewe rs y eas t is t o be p r efe rr ed as co m

’
.
,

p r essed y eas t cake s co n t ai n s ta r c h bac t e r ia a n d o th e r e xtran eou s

-

, ,

m at t e rs A ll of th e ki nd s m ay be cu l t iva t ed t o good adva n tage

.

i n wo rt ( t o be O bta i ned a t b r ewe r ie s) o r in P as teu r s fluids ( S ee ’

.

Hux l ey and Ma rt i n c h ap t e r o n Y eas t ) Wi l d y eas ts m ay be

, .
 LA B OR A T O R Y ST UDI E S A N D DE M N O S TRA TI ON S . 219

fou n d b y exa m i ni ng swee t cide r m ic r oscopically Fo r th e fo l .

lo wm gm e t h od s o f de m o n str at i ng n uc l ei in y eas t and ob t ai n i g

n
ascospo r es we a re i n de bted t o M r S C K ei t h Jr . . .
, .

To Demon str a t e N uclei in Y ea st A ny g ood ac t i v e ly gr owi ng

. -

y eas t wi ll a n swe r bu t a l a r ge ( b rewe rs ) y eas t is p r eferab l e Mix

,
’
.

a li t tl e of th e y eas t wi th an equal am ou nt of t ap wat e r in a t e st -

t ube an d s h ake th o r oug hl y A dd an equa l vo l u m e of He rm ann s

.
’

fl uid and sh ake agai n A s soo n as t h e y east h as se t tl ed pou r Off

.

th e supe r na ta nt li quid and was h t h e y eas t by deca nt a t io n Tr an s .

fe r s o m e O f th e ce ll s to a sl ide fi x by d ry i ng st ai n by Heide n , ,
’
h ai n s i ro n h ae ma t oxy li n m e th od (see Centra thla tt f ur B a cteri
-

olo ie
g xiv , pp 3 5 8
. was h dehy d rat e with a l co h o l
.
,

fo ll ow wi th ceda r oi l and m ou nt in bal sam I n successfu l speci

-

, .

m e ns th e e ff ec t is ve ry sa t i sfac t o ry ( S ee Fi g . .

A S i mpl er Meth od To de m onstr at e n uc l ei in y east m or e

.

uick l y and ve ry eas i l y th e fo ll owi ng m e th od m a be used : B oi l

q y
( in a t e s t t ube
) f o-
r a m o m e nt an i n fusio n of v er
y v igo r o u s y ea st
i n wa t e r p l ace a d rop o f t h e boi l ed i nfu sio n on a sl ide add a
, ,
“ ”
d r op of v ery di lu te D ahl ia so l u t io n cove r and aft e r o ne o r , ,

tw o m i nu te s exa m i ne wi t h a h igh powe r Th e nuc l ei in m ost of .

th e ce ll s wi ll be eas i ly discove rab l e .

To Obt a in A sc ospor es in Y ea st I t has been usuall y recom .

m e n ded to e m p l oy fo r th i s pu r po se b l ock s of p l a st e r of — P aris -

We h ave fou n d th e fo ll owi ng m e th od m o re tr us t wo r t hy

“ ”
~
T h e y ea s t to be used sh ou l d be th e top y east u sed in al e
b r ewe r i es I t s h ou l d a lso be ac t ive l y gr o wi ng and fr e sh I f
.
.

f re s h y eas t ca nno t be O b t ai ned so m e m ay be r evived by cu lt iva ,

t io n f o r 2 4 h ou r s a t 2 5 C in wo rt and a l i ttl e of t h e th ick sedi

°
.
,

m e nta ry po rtio n m ay th e n be p l aced in a ve ry th in l ay er on d ry

filter pape r w h ic h h as p r eviou s ly bee n ste r i l ized by baking
- The .

fi lte r pa pe r is th en p l aced o n a l ave r of cott o n abou t

-
i nc h in
th ick ne ss ly i n g 011 a p l a t e o r sauce r th e co tt o n h avi ng p r eviou sly ,

bee n th o roughl y we tt ed wi th col d st e ri l ized t ap wat e r The -

.

w h o l e is cove r ed by a be ll gl as s a nd set in a r ath e r war m p lace

-

th e cou r e o f t wo o r thr ee da y s po r e wi l l be fo un d
(
°
2 5 I n s s s

in m a n y of th e.
ce ll s T h e l owe r.
th e t e m pe r a t ur e th e l o nge r is
”
“ y ea st 1 8
th e t i m e r equi r ed fo r spo r e fo r m a t io n I f bott o m .

“ ”
u sed i nst ead of top y east a m uc h l o ge r t im e r equi r ed
n i s an d ,

th e r e s u lt s a r e fa r m o r e u n ce rtai n .
22 0 A PPE N DI X .

C HAPTE R X VI .

For th e st udy o f Bac t e r ia i t is ve ry desi r ab l e to h ave a l a r ge

spe ies a n d fo r t h i s pu r po se th e r e is n o n e be tt e r th a n B ac illus
c

meg a teri u m w h ic h m ay be O b t ai n ed f r o m a l m o s t a ny b ac teriolo i

, g
ca l l abo rat o ry and g r ow n i n th e boui ll o n u s ed by bac t e r io l ogi s t s .

D u r i ng 1 8 9 5 and 1 8 9 6 i t m ay be ob t ai ned fr o m B os to n (8 60
above ) .T h is fo r m is ve ry l a r ge and p roduces spo r es r e adi l y , .

”
S ee D e B a r y L ec t u r es O 11 B ac t e r ia ; S t e r be r g B ac t e r io l
( ,
n ,
” “
g
o y ; A bbo t t P r i n cip l es of Bac te r io l og y ;
,
e t c ) T h e pro .

l o nged st udy of bac t e ria is n o t sui t ed to begi n ne rs Vineg ar

ba cteria m ay be see n in th e m o th e r o f —
.

vi nega r by p re ss i ng a bit -

of i t ou t u nde r a cove r sl ip a nd exa m i n i n g wi th a h ig h powe r .

T h e je lly of m o th e r O f vi nega r is a good exa m pl e of z oog

- -

loaa
’

T h e w h i t e scu m w h ic h ap pea rs O 11 aqua ria and i nf usio n s is of

th e sa m e ge n e ra l c h a rac t e r (z oog lwa ) .

C H A PTE R XV II .
( A HA Y I N FUS I O N
) .

To m ak e a succes sfu l h ay i nfu sio n ca r e s h ou l d be tak e n t o

use wat e r co n tai n i ng n u m e r ou s a n d va riou s o r ga n is m s and th e re ,

fo r e di s t i ll ed wa t e r s p r i ng wa te rs a nd we ll wa t e r s a r e in ge ne ral
,
-

,
-

t o be avoided Tap wa t e r s h ou l d a l so be avoided if i t is de rived

.
-

f r o m sp ri ng s o r we ll s The be s t wa t e r fo r t h e pu r pose is th a t
.

d r aw n f r o m po nd s r ive rs l ake s o r o t h e r s u rf a ce sou r ces

, , , .

C l ea n di t c h o r poo l wa t e r is exce ll e n t T h e c h oic e O f h ay is l ess .

i m po rt a nt bu t i t is we ll t o avoid o l d h ay and h ay th a t is ve ry
,

wood y Th e i nfu sio n s h ou l d be wa r m ed bu t n o t h ea t ed o r

.
,

boi l ed I t m ay be kep t in a beake r in diffu s e day l ight , e g in

. . .
,

a n o rt h wi ndow th e b eak e r bei ng l oo se l y cove r ed

,
.

I N S T R UME N T S AND UT E N S I L S .

T h e s t ude n t sh ou l d h ave access t o th e fo ll owi n g a rt ic l es

A co m pou nd m ic r oscope wi th t wo ey epieces a n d l ow a n d
h igh powe r objec t ives abou t 1 in and i in o r objec t i ves . .
,

Mo s t of the pp r t u nd r g n t h r m ntio n d m y be obta i ned f ro m

a a a s a ea e s e e e e a

an y fi rs t -
cl ass d ea l r in p h y i c l nd m i c rosc o p i ca l pp r t u g fro m Th e
e s a a a a a s, e . .
,
 L A B O R A TOR Y s r UDI E s A N D
D EM N O STRA TI ON S .
‘

221

A a n d D of Zeiss , o r 1} a nd
5 i n c h of Bausc h and L om b ;
.
st i ll
hig h e r powe r s a r e desi rab l e) .

A sim p l e dissec ti ng m ic r o scope ; a desi rab l e

fo r m is an o r di
n ar y wa tc h m ake r s l e n s p r ovided wi th a support An o rdi n a ry ’

pocke t l e ns ; gl ass s l ides ( 3 X

-

cove r gl asses wat c h c ry st a l s -

-

,
s m a ll gu mm ed l abe l s
,
n eed l es wi th adjustab l e h an d l es c amel s , ’
,
h ai r b ru sh es b l o tt i ng and filt e r pape r a good r azo r pipett es
,
, ,
( m edici ne d r oppe r s) gl ass r od s a n d t ubes g lass o r p o rcel ai n
-

,
,

d is h es fo r s tai ni ng etc a se t o f sm a ll dissec t i ng i n stru m e n t s , .

,

( s m a ll sca l pe l fo r ceps and s traight poi nt ed scisso rs) a sec tion

, ,
-

,
lif t e r pie es of pi th fo r s ec t io n cu tt i ng thr ead a sh a ll ow
tin pa n
c
,
-

, ,

l i n ed w i th wa x l o ng i n sec t pi ns fo r pi nni ng ou t dissec t ed speci

,

m e ns d ra wi n g m a t e ria l s an d a n o t e book fo r ske tc h es and oth e r

, ,
-

reco r ds .

E ac h tabl e s h ou ld be fu rnish ed wi th a set of s m all r eage nt

bo ttl es a Bu n se n bu rn e r wa sh bo ttl e t est t ubes beake rs an d a
, ,
- -

, , ,
b e ll gl ass f o r p r o t ec t io n f ro m du st
-

Th e rm o m et e r s a ba l a nce .

, ,

m ic r o to m e d ry in g ove n and a paraflin wat e r bath sh ou l d al so be

, ,
-

a cc essib l e .

R E A G EN TS AND T E C H N I C A L ME T H O DS .
*

A lcoh ol .
—
S i n ce bio l ogica l
l abo r at o r ie s be l ongi n g t o incorpo
ra te d i ns ti tu t io n s O b tai n al co h o l du ty f r ee i t s h ou l d be libera ll
y ,

su
pp l ied a n d f r ee l y used A l c o h o l of is “
abso l u t e ”
.
‘
.

”
a l co h o l m ay be pu rc h ased in 1 pou nd bottl es
, S quibb s -

.
’

abso l u t e a l co h o l m ay be ob tai ned O f any drug g is tfi b ut o r di na r y

alco h o l of 9 0—9 5 % a nswe rs n ea rl y eve ry pu r pose “ C ol ogn e .

”
spi r i ts i e al co h o l of a bou t
,
. .
,
m ay be ob tai ned fro m th e
dis till e rs a t 6 0c o r th e r eabou ts pe r gall o n I t m ay th en be
.
, ,
.

B a usc h L o m b O p ti c l C R oc h t r N Y th F rank li n E d uc tion l a o .

, es e , .
:
e a a

Co . H m ilto n P l c B o to n ; or Qu n C C h t nut S tr t Phil d l phi

, a a e, s ee o . es ee ,
a e a .

Ch m i c l n d oth r a ppa rat u m y be obta i n d f ro m E i m r Am nd 205 —

,

e a a e 21 1 s a e e e ,

T h ird Av nu N Y e e, . .

E v ry l bor tory ho ul d be upp l i d w ith so m

e a a f th t nd rd book u pon
s s e e o e s a a s

th i s ubj c t
s
g e S tr b u r
,
g r B t nis
e c h
. e Pr c ti .um J
,
n ; W h it m n as e
’
s o a a c , e a a
'

M th d f Resea rch i n M ic cop ica l A n t my nd E mbry l g

e o s O y B o to n L ros a o a o o ,
s : e e,

The Al icr t mis t Va dc M ec um l t d itio n ; Zi mm er m n Bot nic l Mi

’ ’
o o s as e a s a a c ro
,

tech niq u ( H um p h r y ) H olt N Y

e e , , . .

S ee l W hi t ma n l c p 1 4
a so , . .
, . .
2 22 A P E N DI XP .

di l u t ed t o 8 075 7 056 ,
e t c as n eeded For this pu rpose a n
, .
, .

a l coh o l i m e t e r is ve ry co n ve n ie nt .

A cet i c A ci d —O n e o r t wo pa rts gl acia l ace t ic ac id t o 1 00 pa rt s

.

wa t e r .

A cet i c A cid and Met hy l g r een — T h is is va l uab l e fo r s tain i ng

-
.

n uc l ei in vege t a l t issue s D is so l ve m e thy l gr ee n i n o n e o r two

.
-

pe r ce nt ac e t ic ac id u nt il a ric h deep co l o r is O b ta in ed .

B or ax ca r mine — A dd t o a 4 5‘ aq ueous s o l u t io n of bo rax 2 —

-
. 3 5‘
ca rm i ne a nd h ea t u nt i l th e ca r m i ne d is so l ves A dd a n equa l
,
.

vo l u m e of 7 05‘ a l co h o l and filt e r aft e r 2 4 h ou r s A ft e r s tai nin g

, .

( 6 — 1 2 h ou r o r ms
o r
,
e fo r l a rge O bjec ts a few m i n u te s fo r sec ,

t io ns) plac e th e objec t i n acidu l at ed a l co h o l ( 1 00c c 3 5 5i a l co h o l .

,

3 —4 d r ops hy d r oc hl oric acid ) a n d l ea ve u n t i l th e co l o r t u r n s f ro m

du ll t o b right r ed ( 1 0—3 0 A ft e r wa r ds r e m ove to 7 05‘
a l coh o l
Ca n a da B a lsa m Mou n t in gih —
.

,
T his i n va l uab l e subs ta n ce m ay .

be ob tai ned i n t h e c rude co n di t io n d ried by p ro l o n ged h ea ting , ,

a n d th e n disso l ved in chl o rofo r m be n zo l e o r t u r pe n t i ne f or , , ,

u se The be nzo1e so l u t ion is pe rh a ps th e be st a n d m ay be o h

.
,

ta ined f r o m m os t of t h e dea l e r s T h e p ri n cip l es o f m ou n t i n g i n

.

bal sa m a re ve ry s i m p l e I t doe s n o t m ix wi th wa te r o r a l co h o l
.
,

bu t m ixe s f ree lv wi t h c l ove oi l c hl o r ofo r m be nzo l e etc -

Oh , , , .

ec ts a r e th e r efo r e ge ne ra lly tr ea t ed fi rs t wi t h ve ry s tr o n g a l co
j ,

h ol 9 5
,
in o r de r t o r e m ove th e wa t e r ; th e n wi th c l ove o il -

c hl o r ofo rm o r t u r pe n t i ne t o r e m ove the a l co h o l and afte rwar ds

, ,

m ou n ted in a d r op o f ba lsa m Th i s sh ou l d u sua lly be p l ace d o n .

th e cove r glas s w h ic h is th e reu po n i n ve rte d ove r th e obj ec t

-

,
.

T h e bal sam g r adua ll y se t s and th e p r epa r a t io n s a re pe rm a n e n t ly

p rese rved
Ca rmi n a —C an n ine m ay be ob ta in ed as a powde r w h ic h
.

w h e n r ubbed u p t h o r oughly wi t h wa t e r in a m o r t a r passes i n t o a

s t at e of ve ry fi ne s ubdivi s io n T h is p r ope rty m ake s i t av ail ab l e .

fo r expe r i me nt s wi t h ci l ia e t c ,
.

I t is m o re oft e n u sed i n so l u t io n as a st ain i n g age n t ( S ee ,

.

B or a x carmi n e
Cell ulose test —S a t u r a t e th e objec t i n iodi ne so l u t io n was h in
.

-
.
,

wat e r and p l ace i t i n s tro n g s u l p h u r ic acid p r epa r ed by ca refully

,

pou r i ng 2 vo l u m es of th e co n ce ntra ted acid i n t o 1 vo l u m e of

wa te r .
 LA B O RA TOR Y ST UDI E S AND DE M ON STRA TI oN S . 22 3

Collodi on a nd Cl ov e oil
-
.
—Used fo r fi xin g sect ion s
th e slide to
i n o r de r to p r eve n t th e di sp l ace m e n t of de l icat e
o r iso l at ed pa rt s

m ba l sa m m ou n t in g M ix one part of eth er co ll odion and thr e e

-

. -

pa rt s o f o il of c l oves In m ou nt i n g va r nish a sli de wi th th e

.
,

m ix t u re b y m ean s of a ca m el s h ai r b rus h l ay o n th e sect ions ’

-

an d p l ace th e s l ide fo r a few m i n u t es

o n th e wat e r ba th i e
( -

i
.

u nt i l th e c l ove oil evapo r a t es) Transfe r th e sl ide t o a wide

-

m ou th ed bo ttl e O f t u r pent i n e (t o disso l ve th e paraflin) rem ove i t ,

a n d d rain o ff th e t u r pe nt i n e p l ace a d r op of C anada bal sam o n

,

th e m idd l e of a cove r g l as s and in ve rt i t ove r th e object

-

Dahlia — D isso l ve i n wa t e r
.

R osin — Di s so l ve i n wa t e r u n t i l a b r ight r ed so l u t ion is oh -

tained . I t s h ou l d be di l u t e d w h e n used
Gl y cer i ne dilu t e —Two pa rt s gly ce r i ne one part dist i ll e d
.

,
,

Hmmat oxylin —A dd
c c of sat u r at ed al coh ol i c 4 . .

so l u t ion of h ae m a t o xy li n t o 1 5 0 c c of str ong aqueous so l u t ion of . .

amm o n ia a l u m ;
-
l e t th e m ix t u re stan d a week or m o r e in th e
lig ht filt e r and add 2 5 c c of gly ce r i ne and 2 5 c c of m ethy l
, ,
. . . .

alco h o l The fl uid i m p r oves g reatly aft e r s ta n di ng som e week s

.

Hiemat oxylin — To a sat u rat ed so l u t ion of cal

c ium c hl o r ide in 7 0 % al coh o l add an excess of p ure al u m ; filte r
aft e r 2 4 h ou r s and add 8 vo l u m es of 7 0% a l co h o l filt e r i ng agai n ,

if n ece s sa r y A dd a sat u ra t ed a l co h o li c sol u t io n of h sematoxylin

.

u n t i l th e l iquid beco m e s pu r p l e bl ue Th e l onge r th e l iquid -

.

s ta nd s be fo r e u si ng th e be tt e r I t sh ou l d be di l u t ed fo r use
,
.

wi th th e al u m ca l ciu m c hl o ride so l u t ion in 7 0% al co h ol

- -
.

Herma nn s Fl ui d — S ee L ee s Va de M ecum
’ ’
.

I odine Sol u t i on — D i ss o l ve po t a ssiu m iodide in a sm all quan t i t y

o f wa t e r add m e ta ll ic iodi n e u nt i l th e m ix t u re assu m es a da r k
,

b r ow n co l o r an d th e n di l u t e t o a da rk s h e rr y co l o r
,
T h e so l u -
.

tio n s h ou l d be kep t fro m th e l ight .

Mag en t a ( A n i li n e R ed) —D i sso l ve i n wa t e r.

.

t
Me hy l G eenr — Used in aqueous o r a l co h o l ic so l u t io n or

N mr a F l i d N r ma l S lt S l u t i n ) —D isso l ve gr am s of
o l u ( o a o o .

sodium c hl o r ide i n 1 l i tr e of di st i ll ed wat e r .

Paraflin .
H ar d ” and
“ soft p araffi ns, i . s .
, th ose of hi gh
2 24 A PPE N DI X .

a nd l ow m e lti ng po in ts s h ou l d be m ix ed i n suc h p r opo rt io ns th at

-

,
°
t h e m e l t in g poi n t lies be t wee n 5 0 an d 5 5 C
°
-
.

Pereny i s Fl ui d —Ten pe r ce n t ni tr ic ac id 4 pa rt s 9 0% a l co
’
- -

h o l 3 pa rt s 5 } aqueous so l u t io n of c hr o m ic acid 3 pa rts N ot

,
7 .

to be u sed u nt il th e m ix t u r e assu m e s a vio l e t h ue L e ave o bjec ts .

in th e fl uid 3 0 m i n u t e s t o a n h ou r t h e n 2 4 h ou rs i n 7 056 alco h o l , ,

a n d fi na ll y p l ace i n 9 0 pe r ce n t a l co h o l
—
.

t M r a t i ng Flui d Disso l ve a gr a m of po tas siu m

’
Sc h ul ze s ace

c hl o r a t e in 5 0 c c of n i tric acid T h e t issue s h ou l d be boil ed

. . .

in t h e m ix t u r e an d af t e rwa rd s th o r oug hl y was h ed in wa t e r .

Sch ul ze s S olut i on — D i ss o l ve zi n c i n pu re hy d r oc hl o ric ac id

'

e vapo r a t e in th e p rese nce o f m e t a ll ic zi nc o n a wa te r bath t o a -

, ,

s m py co ns is t e nc y add as m uc h iodide of po tass iu m as wi ll dis

y ,

s o l ve,
and t h e n sa t u ra t e w i th i odi ne ( W h e n h ea t ed wi th th is .

fl uid ce ll u l ose t u r ns b l ue .

Sect i on cutt in g— Ma ny objec ts ca n be cu t b y h a n d wi th a

-
.

razo r ( wh ic h m u s t be ve ry s h a r p) T h e obj ec t sh ou l d be h e l d in .

th e le f t h a nd w h i l e t h e raz o r is poi nt ed away f r o m th e bo d y a n d ,

a ll owed t o r es t O 11 t h e t ips of th e fi nge r s wi th its edge t u r n ed

t owa rd s t h e l ef t I t is t h e n d rawn ge ntl y toward s th e bod y so

.

a s g radua ll y t o s h ave o ff th e sec t io n Sm all objec ts m ay be h e l d .

’
be t we e n t wo pieces of wa t c h m ake r s pi th p reviou sly soaked in
wa t er . I n ei t h e r case t h e razo r s h ou l d be k ep t we t .

Ma ny O bj ec t s h oweve r requi re m o r e ca r efu l tr ea t m e n t b y

, ,

o ne of t h e fo ll owi ng m e t h ods :

A P a rcm j ll etho tl A f t e r h a r de n i g a nd s t ai n i ng th e
’

. zn i n
.
-

o bjec t is soaked in s t r o ng a l co h o l ( 9 5 % o r m o r e u nt i l th e wa te r
)
is th o r oug hly e x t rac t e d ( 2 — 1 2 h ou r s c h a ngi ng th e a l co h o l a t ,

l east o nce) th e n in c hl o r ofo r m u n t i l t h e a l co h o l is ex trac t ed

,

( 2 — 1 2 ) h ou r s
) , (
a nd th e n in m e lt ed pa raffi n n o t wa r m e r th a n 5 5
°

C ) O 11 a wa t er ba th fo r 1 5 t o 3 0 m i nu t e s ( to o h ig h a t e m pe r a
.
-

t u re o r t oo l o ng a ba t h cause s exc es sive sh ri n kage) S o m e of t h e .

pa raffi n is th e n pou r ed i nt o a s m a ll pap r box o r i n to adju s tab l e e -

m e ta l f r a m e s Th e O bj ec t is tr a ns fe rr ed t o i t and aft e r t h e m ass

.

h as begu n to s et i t is p l aced in co l d wa t e r u n t i l qui t e h a r d It .

is th e n ce m e n t ed ( by pa r affi n) t o a squa r e piece of co rk a n d

p l aced i n t h e s ec t io n cu t t e r o r m ic ro t o m e
-

T h e sec t io n s m ay be cu t s in gl y wi th th e ob l ique k nife o r by

 LA B ORA TO R Y ST UDI E S AND DE M ON STRA TI ON S . 2 25

th e r i bb o n m e th od th e kn ife be in g kep t d ry i n ei th e r case In

-

,
*
.

m ou nt i ng th ey sh ou l d be fi xed by th e coll odion m eth od ( S ee -

Collodi on and Clov e oil ) -

B
. Cello ioli u M ethod — T h i s is especiall y app l icab l e to de l i
c a t e v ege ta l tissues A f te r de hy drat i ng th e objec t th o roughly in
.

a l co h o l soak i t 2 4 h ou r s in a m ix t u r e of equal part s of al coh o l

,

a n d e th e r M ake a th ick so l u t ion of ce ll oidi n in th e sam e m ix

.

t ure an d soak th e objec t fo r so m e h ou r s in i t I t m ay th e n be .

im bedded as fo ll ows : Dip th e sm all e r en d of a tape ri ng co r k

in th e ce ll oidi n so l u t io n a ll ow i t t o d ry fo r a m o m e nt b l owi ng
, (
an d th e n bu il d upo n i t a m a ss of ce ll oidi n
o n i t if n ecessa r y
) , ,

all owin g i t to d ry a m o m e nt aft e r eac h addi t ion Tr ansfe r th e .

objec t to th e co r k and cove r i t th o roughl y wi th th e cell oidi n .

T h e nfloa t th e co rk in 8 2 —8 5 56 s
p gr ) a l co h o l u nt i l t h e . .

m ass has a fir m co nsis t ency ( 2 4 I t m ay th e n be cu t in th e

m ic r o to m e wi th th e ob lique k n ife w h ic h m u st be kep t d rippi ng
,

wi th 8 2 8 5 % a l co h o l K eep th e sect io ns in 8 2—8 5 % al coh o l u nt i l

-
.

r ead y to m ou nt th e m th e n soak th e m fo r a m in u te in str o ng

,

al co h ol tra n sfe r to a s l ide pou r o n c hl o r ofo r m u n t i l th e al coh o l

, ,

is r e m oved d r ain O ff th e l iquid quick ly add a d r op of bal sa m

, , ,

a n d cove r ( S ee al so W h i t m a n 1 c p
.
,
. .
, .

S ee Wh it m an l
, . c. p . 71 .
A b so rp tion 48 5 2 , , , 1 01 , 1 65 . B iology 1 6 7 8 , ,

cc re tio n 1 66 , , .

A , .
B i x u l 7 3 13 0
se a , ,

c hro m a ti n 23
.

A , .
B l to por 80 85
as e,
c ti n o p h ry s 1 66
, .

A . . B l to ph r 8 5
as s e e, .

A d p ta tio n 97 98
a , , , 144 . B l t ul 80 9 0
as a, , .

A d ve ntitio us b uds , 1 30 . B loo d 15 1 6 9 0 1 02

, , , , .

B loo d v l 5 4 -
es s e s , .

zE tio lo gy, 6 . B l u gr n l g 183 1 92

e -
ee a a
e, , .

A mo g e nes is 7 3 , 1 3 0 1 63 . , . Bod y 1 9 24 84 1 07 1 5 6
, , , , , .

b um ino us es 36 bod i . . Bod y c v ity 4 7 -

a , .

A l i m t tio
en a n, 10 5 . Bo n 1 6 e, .

lm t r c
A i e n a y a nal 82 92 , , . B ot ny 6 7
a , , .

Al im t r t m
en a y sys e , 49 . B r nc h 1 1 1 122 1 3 0
a es , , , .

A llotobop ho ra 4 1 , . B r nc hi 62
a ae, .

lt r tio
A e na n of e ne rat io ns, 1 30 . B udd i ng 1 86 , .

A moeba 2 7 , 1 216
, . Bu rs i 1 7 6 ar a, .

m boi d c ll
A oe e s 64 , .

A mph i t r
as e 84 , . C l c i f ro u gl nd 5 1
a e s a s, .

Am p h m i x i
i s 1 68 , . CA K I N S G N 1 7 1
L , . .
, .

A nabol i ms 3 3 10 0, 149 , 1 64 , , . C p i ll ri 5 4
a a es , .

A nac hmi s , 29 . C p ul f gg 7 8
a s es o e s, .

C p ulog no u gl nd 4 6
a s e s a s, .

A n to m y 7
a , . C rboh y d r t 3 7 1 01
a a es , , .

A ni m l c ul 1 5 8
a e, , 1 99 . Ca rch esi um 1 76 ,
.

A nn ul u 1 32 s, . C rnivor 1 7 7 203
a a, , .

A n us , 46 , 8 2 1 65 , .
C rtil g 1 5 1 6
a a e, , .

An e th ri d i
a , 1 35 .
C ti ng 4 2 5 3
as s, , .

A orti c rc h
a es , 5 4 , 5 5 . C ll 1 2 20
e , ,
.

A pic l b d
a u s , 1 1 1 , 1 1 6, 1 23 . C l l d ivi ion 24 8
e -
s , ,

p c l c ll
A i a e 1 23 , .
C ll th ry 20
e -
eo , .

p g m
A o a y 14 3 .
C ll ulo 3 7
e s e, .

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,

A p o pory 1 43
s .
e -
a , , .

C ntr om 79 83 84
,

A c lla
r e e os e, , , .

A rc h g o n i
e 1 37
,

a, .
C r br l g ngl i 65 69
e e a a a, , .

A rc h n t ro n 80 8 2 8 5
e e , , .
Ch l k 1 66
a , .

Ch
,

A r c h pori um 1 3 1
es .
24
a ra , .

C h m iot x i 1 3 9
,

A r c ho p l m 7 9 80 as , .
e a s, .

A rth ro po 19 5
s re .
,
Chlo c cc u 1 78
ro o s, .

C hlor gog u c ll 5 2 61 9 3
.

A s co por 187 a e e s,
-

s e, , , .

C hloro p hy ll 1 2 6 1 5 1 2 1 5
.

A sex u l r p ro d uc tion 7 3
a e , . ,
.

,
C hloro ph y l l bod i 1 79 2 15
.

A i m il tion 1 8 2 es ,
-
.
ss a ,

Ch oc u 1 83
.
,

t r
'

s,
A s e 79 84 ro oc c .

C hrom ti n 23 83
.
, ,

a , , .

C hrom to p hor 1 4 7 1 79a es , , .

C hrom o om 83 84 s es , ,
.

C il i 3 1 63 7 4 1 3 7 1 92
a, , , , , .

C i rcul tion 4 8 5 3 101 1 65

a , , , , .

C L A P A R E DE , 96 .
22 8 I N DE X .

Clas i fi ca tion 7
s ,
. En d o pore 187 1 94s , , .

Cl it ll um 4 6 7 7 7 8
e , , , , 88, 92 . En d o pori um 1 34 s , .

C oa g ul tio n 36 39 a , , . E n r gy 32 99 1 4 6
e , , , , 15 1 .

C occ i 19 2 ,
. E ntobl as t 8 1 , .

Coel e nte ra ta 88 , . E n to p‘l as

m 15 8 , .

( m lo m 4 7 8 2
‘
, , . E n i ro n e nt 97 1 03 , 1 44 , 15 1
v m , , .

C l m i fl ui d 5 3
cE o c , . pd m
E i e r a l sys te 1 14 m , .

CO R N , 2L pd m
E i e r i s , 1 14 1 1 6 , .

C ol d tor g 1 99 s a e, .
fi ning / I ts 1 7 6
'

, .

Col loi d l 36 a , . p
E ith e l i u , 9 0 m .

C lo ny 1 7 6
o , . E ug le na 1 7 6 , .

C mm i ur 65
o ss es , . E xc
retio n 48 5 3 , 5 9 , 100, 1 65 . , .

Co nj ug tio n 1 7 1 181 a , , . x p
E o s ori u 1 34 m , .

Co nn c tiv ti u 7 0 90
e e ss e, , . p
E y e s ot , 1 7 6
-

Co nsc io u n 69 7 0 s es s , , .

Co nt r c til ity 62 164

a , , . Faeces 5 3 , .

C O din t i n 48 64 67 1 64
o r a o , , , , . FAR L O W 143 , .

C p ul tio n 7 7
o a , . F ts 1 7 3 7 1 01
a , , , .

C ro f rtili z tion 74
ss -
e a , . F th rs 1 8
ea e . .

C ry t l 17
s a s, . F r n 1 05
e s, .

C u hio n 1 3 5
s , . F rm n t 5 2
e e , .

C uti c l 7 1 9 1 e, , . F rm nt tio n 1 9 1 197

e e a , , .

C y a no p h y c eaa 1 83 , 192 , 1 99 , . F rtil i z tio n 7 3 78 1 89

e a , , , .

C c li c l c h g
y a a n e , 5 , 7 2 , 89 . F ibr v c ul r t m 1 14
o
-
as a sy s e , .

C to p l m
y as 22 , , F ibro vas c ul r bund l es 142
-
a , .

F iltr tio n 200a , .

DAR W I N 4 2 5 1 7 0 99 1 0 3 , , , , , . F i io n 1 63
ss , .

De a th 1 5 2 , . F l g ll um 1 7 6 1 92
a e , , .

DE B A B Y 1 1 5 1 4 3 , , . FO L , 79 .

De f ze c a lio n 5 3 1 65 , , . F ood 1 46 s, .

De s m i d s 1 7 8 1 83 , , . For m i ni f r
a e a, 1 66 .

Di a l y s i s 3 6 2 10 , , . For gut 86
e -

, .

Di a s t a ti c f e rm e nt 5 2 , . Fo s rs n MI C H
'

, A EL , 1 5 3 , 1 63 .
Di a to m s 1 7 8 18 3 , . . FR E D E R K Q 5 2 '

, .

Di c ho g a m y 1 38 . . F d
ro n 1 25 , .

Di ff e re nti a t io n l l 84 1 4 1 , , , . F c
un tio ns 9 , .

Di ff e re n ti a tio n a nt e ro p os te rior, ,
~
F und m nta l a e sy s e t m , 1 14 .

1 10 . F ung i 1 4 7 , .

Di ff e renti a tio n dors o ven tra l , 43 1 10 ,

-

, .

Di ff e renti a tio n o f t h e ti ss ues, 25 G am t 1 8 1 e e,

m
. .

Di e/g ut 1 66 , . G m og n i 7 3 130 1 68
a e es s , , , .

Di g es tio n 4 8 4 9 5 2, 1 01 , 1 65 , , , . G ng lio n 64 9 4
a , ,

Diplococc us 194 . . G tr ul 8 0
as a, .

Di s e ase g en us 1 92, 1 97 -

, . G tr ul tio n 84
as a , .

Di s i n f ec tio n 2 00 , . G r m c ll
e 24 7 3 90 1 30
-
e s, . , , .

Di s se p i m ents 4 7 9 4 , , . G rm i n tion 1 3 4
e a , .

Di s trib ut io n 7 , . G rm l y r 8 1 84 85
e -
a e s, , , .

Di v i s io n o f l a bor , 1 1 , 26, 1 5 6, 1 65 . G r m l y r th ory 88

e -
a e e , .

Dors a l p ore 4 8 ,
. G rm pl m 89 1 5 2
e -
as , , .

Dors a l v es s e l 5 4 , . G rm i n l pot 74
e a s , .

DUJAR D I N , 2 1 . G rm i n l v i c l 7 4
e a es e, .

G i nt fi b
a 94 -
res , .

E a rth w or m , 41 . G ill 62 s, .

E c tobl as t 8 1 , . G i rd l 7 8 e, .

E c to p l as m 1 5 8 , . G i zz rd 5 1 7 1
a , , .

Eg g 24 . .
G lazoca p sa . 1 7 8 , 1 83 .

Eg gl y i ng 7 7 a , . Glu o c s e, 52 .

Eg gnuc l us 7 9
-
e G l y co g e n, 3 7

g
, .
.

Eg g tri n ; 7 4
-
s . G r ega ri na 64 .

E m bry o
,

, Gro w th 1 65 , .

E mbry ology 7 7 2 , , , 78 . Gua r ells dc

1 28 -

, .

Haema tococcus
H m og lobi n
ae
H i r 18
a , .
Hay inf us io n 2 01
H rbi vora 1 7 6
e 203
H e red ity 84
H r m p hrod ite
e a
H E RT W I G 7 9
H i nd gut 8 6
-
H i s tolo gy 7
H OOK E R O B ER T 20
, ,
H OO K E R S I R W J 106 , .
H O PP E S E Y L E R 35
H UXL E Y 2 4 , ,
H y p od e r m i s 92
I mp g
re na tio n, 7 3 , 1 39
dvd
I n i i ua l ,
d
I n us i u , 1 3 1
m N itro en, 1 4 7 g
I nf us io n s 1 68
I n f us ori a , 1 68 2 1 7
I nh e ri ta n e 8 , 84
c p
I n t uss us e tio n, 4 , 165
I rri ta i lity , 1 64
b .
JO HN S O N , 3 5
K a ta bol is
L a t e ra l rid es , 1 1 1, 1 14
L ea f 1 1 , 1 5
L a xn o s s fi x 9 5
L umbric m
Ma c ro ga m e te 1 7 5
Mac ro n uc l eus 1 7 0 1 71
Ma l i c ac i d 1 3 9
MA UPA S 1 7 0
, .
M e ri s te m 1 23
M es ob las t 8 1
, .
, .
M es o p h y ll 1 26
Me ta boli s m 3 3 1 0
0 1 01
M e ta m e ri s m 4 5
M ETC H N I K O FF 5 3
M i c ro ga m e te 1 7 5
M i c ro n uc l eus 1 7 0 1 7 1
229
178 .
Moth er of v i negar 1 94
54
,
, .
M oth er c ell s 1 34 1 3
-
-
, ,
M otion 4 8
7
-
, , .
, .
, .
Motor sy s t em 62
Mouth 4 9 80 8 5 1 65
, .
, , .
, ,
Mus c l es 14 2 6 2 7 62
, , .
, .
7 3 , 1 30 90 , ,
MU L D E R 3 5
, , , .
, .
, .
Mycoderma 1 94 202
, .
My x ob ac t eri a 1 9 9
, .
, .
My x omy c et es 1 9 9
,
, .
,
.
N a t ur a l s el e tion 9 9 c
.
, N e hri i a p 59 d , .
, .
N e r es , 64 9 0
v
, .
.
N e r e ell s , 9 4
v c
, .
-
, .
N e r e entr e, 68
v -
c
N er e i v
.
-
mp
ul s es 67
N er o us s y s t e
v
.
,
.
, 64 , 82, 94
, 10
2 m
N itella 2 8
.
, .
.
, .
N ucleo lus 2 3
, .
uc leo pla s m 22
0
, .
c , .
uc leus 1 6 2 3 1 86
, .
.
N utrition, 99 , 1 4 6
, , , .
.
(E s op h g
a us 1 8 , .
.
O ld ag e 7 2, 1 5 2 1 66,
, .
OOph o re , 1 3 0
m .
, 38 99 , 1 4 9, 1 64 , . O b s ph ere, 7 3 , 1 38 .
O as po re, 1 3 9 .
1 86, 1 95 . O rg i m
an s s, 9 .
O rg og
an en , 85
y
O rg a ns , 9 .
Ov ri
2 a es , 74
.
.
.
Ov i d ct
u 75 , .
, .
Ov mu , 7 3 7 4 , 89 , .
P a m mazcium 1 68
, .
, 41 .
Pa ras it es 1 9 2 , .
P c m
a r en h
y a 1 16 , .
PA E R
ST U 188 , .
P t uri z tion 200
as e a , .
P t ur fl ui d 1 89 197
as e
’
s , , .
, .
P thog n i c 200
a e
, .
, , . P thology 6 7
a , , .
, .
P p ti c f rm nt 5 2
e e e
, .
P p ton 5 2 1 01
e e,
. .
P ri t lti c ction 5 1 5 4 5 5
e s a a s, , , .
P FEFFE R 1 39
, .
, .
P h gocy t 5 3 61 64 1 5 8
a es . , , , .
, , , P h ry ng l g ng li 67
a ea a a,
.
, . P h ry nx 4 9
a
.
,
, .
P hy iolog i c l p p rt es of proto
s a ro e 1
, .
p l m 163 1 82 1 83
as
, , , .
, , . P h y iology 6 7 166
s , , , .
P hy iology f th nervous system 67
s o e
so
, .
. P ol r c ll 7 9
a e s, .
Pol c ll 82
e -
e s, .
Poi o n 3 9
s s, .
Pl m 5 3
as a, .
P leuroooocus , 1 78 .
2 30 I N DE X .

P rim ordia l utri c l 29 e, . S m ll 4 2 69

e , , .

P roc tod um 8 2 86 ae , , . Soc iolog y 7 8 , , .

P ro nuc l u 7 9 e s, . So m ti c c l l 7 3
a e s, .

P ro nc h ym 1 1 6
se a, . So m ti c l ay r 85
a e , .

P ro to m i um 45
s , . S om t p l ur 82 86
a o e e, , .

P rot c tio n 7 1
e , . So m it 45 es , .

P rot i d 3 33 5 2
e s, , . S P E N C ER H E R B E RT , ,
3 , 99 , 1 46 .

P rot u m m l ule 2 7 1 5 8
e s a a c , , . S p r m ri
e 74 75 a es , , .

P roth ll i um 1 3 0 1 35 2 14
a , , , . S permat o s ph ere , 7 7 .

P ro toco cc ua 1 7 8 , . Sp r m to z oi d
e a , 13 7 .

P ro to nema, 1 34 . S pe rma to zo b n 7 3 , 74 ,

P p m
roto l as 1 6 20, 207 , 208 , , . p rm d ct
S e u 76 -

, .

P z
roto oa , 1 5 8 . p rm c l
S e n u eus 7 9 -

, .

P d pd
se u o o i a 2 7 1 5 8 , ,
°
Sp i d r ort
e w 29 , .

P c g
s y holo y 7 , 8 , . Sp irilla 1 92 , .

P ul 5 4 se , . Sp l anc h no p l ure 82 86 e , , .

P utr f c tio n 1 9 7
e a . ,
201 . Sp o nt n o u g neratio n 33
a e s e , .

P UR K IN JE 2 1 , . S p o r ng i a130 a, .

Spor 24 1 3 0 194
es , , , ,

Ra d iol ri 1 66 a a, . Spor p hor 1 30o e, .

R c pt c l
e e 131 a e, . Stap hy l c cc u 1 94 o o s. .

R c p ta c ul um
e e orum 7 5 ov , . S t rc h 1 7 3 7 1 4 6
a , , , .

R fl x ac tio n 67
e e , . St nt 1 7 6
e or, .

R g n r tio n 7 3
e e e a , . S t ril i z tio n 1 99
e a , .

R p rod uc tio n 48 7 2 1 11 1 30
e , , , , ,
S ti m ul u 6 7 s, .

R esp ir tio n 6 1 1 5 0 165

a , , , . S ti pe 1 25 , .

R E TZI U S 9 5 , . S to m c h i nt ti ne 5 1
a -
es , .

R h i zoi d 1 34 s, . S to m t 1 26 1 28
a a, , .

R hi z om 1 1 1 14 0 e, , . S to m od um 82 86 ae , . .

R hi z o pod 1 66 a, . St r ept cocc us 1 94 o , .

R i gor c lori 39 a s, . S tr uggl f x i t ence 203 e or e s , .

R i g or m orti 209 s, . Sigh 11011111 1 7 0

/ 11 1 , .

R oots , 1 22 . S ug r 3 7 a , .

S un ni m l c ul
-
a 1 66 a e, .

S ccha r my ces
a o , 1 84 . S u rv i v l f t h fi ttes t 99 a o e , .

S AC H 1 15 S , . Sy m bio i 1 7 7 s s, .

S l iv r g l nd
a a y a s, 51 .
S y mm try bil t r l 44 1 1 0
e ,
a e a , , .

S a p, 1 4 . S y mm try ri l 45 e , se a , .

S p ro p h yt
a es , 1 92 . Sy m p th ti c y s te m 67
a e s , .

S rc i n
a 1 94 a, .

Sc hi z o m y c t e es , 19 2 . T te 4 2 69 7 0
as , , , .

S o n nm ns n 20 . . T x o no m y 7
a , .

S c ri uurz s M A X , 2 1 , . Te m p ra t ur 38 199 e e, , ,
2 10 .

A
S C H W N N , 20 . Te t es 7 4 7 5
s , , .

Sc i nc biolog i c l 1 6
e es , a , , . T i ues l l 1 3
ss , , .

S c i nce p h ic l 1es , y s a , . T o uc h 42 69 7 0 , , , .

S gm nt tio n 24 8 0
e e a , , . T o x i c ology 3 9 , .

S gm nt tio n c v ity 84
e e a a , , 85 . T r c he 1 16
a ae , .

S m i n l r c pt c l 7 7
e a e e a e, . T r c h ei d 1 1 6
a s, .

S mi n l v icl 7 6
e a es e, . Tra des m n tia 2 9 , .

S n tion 4 8
e sa , . T p
ra ns ira tio n 1 4 6 . .

S n or g n 4 2 69
e se a s, , . T c c
ri ho y s t s 1 68 , .

S e ns e s 4 2 69 , , . T p c
ry ti f e r ent , 5 2 m .

Sen iti v y t m 69
s e s s e , . T w i ns 88 , .

S t 4 6 63
e ae , , . T p
y hlos ol e 5 1 9 1 , ,
.

S et i g ro u g l nd
e 63 7 7 s a s, , .

S x u l r p rod uc tio n 7 3
e a e , . Un i c ll ul r ni m l 1 5 8
e a a a s, .

S i v t ub
e e 1 16
-
es , . Uni c ll ul r org n i m 1 5 6
e a a s s, , 1 77 .

S i g ht 4 2 69 7 0
, , . Un i c ll ul r p l nt 1 7 8
e a a s, .

Sk in , 1 28 .

S li ppe r a ni m a l c ul e , 1 68 . Va c uol es , 24 , 1 62 1 7 0 , .
I N DE X .
23 1

W hite blood cell s 64

W hirl pool 2
, .

W I N O G RADS K Y 1 9 7
, .

, .

Y eas t , 1 7 8 .

Y e as t , botto 19 0 m , .

Y eas t red, 1 9 1
, .

Y eas t to, p 190

, .

d
Y ea s t , w il , 1 9 0 .

o glo
Z o ea , 1 9 4 , 1 9 5 .

ZoOids , 1 7 6 .

Zo Olog y, 6 7 , .

Zo Os pores , 1 81 .

Zoé tk a mmon, 1 7 6
'