T HE M AC M I L LAN C OM PAN Y

MAC M I LL AN CC ,
LI M IT E D
L
.

ON DO N B O M B AY C A LC UT T A
M E LB OU R N E
 T HE UNI T ED S T AT E S OF AM R A
E IC

1 91 0 .

B Y T HE M AC M I LLAN C OM P AN Y .

Se t l t
u p a nd e e c ro type d . l
P ub i sh e d N ov e m b e r , 1 910 .

J u ne , 1 91 1 ; M a y De ce m b e r
, . 1 912 ; No v mb
e e r, 1 91 5 ; M a r ch ,
1 916 ;
Se p t e m b e r , 1 9 1 7 ; F e b r u a ry , Oc to b e r , 1 91 9 Nov mb
e e r , 1 9 20 ; Octo b e r ,
No v mb
e e r, 1 92 1 ; J anua ry , 1 9 24 .

N or inoon 13 1 25 5
J . S C us h i ng C o
. . B e r w i ck S m i th C o .

N o r wo o d ,
 P REFAC E

THI S b oo k h as b e e n w ritte n f or th e u se of stude nts ta king the

intro ductory co ur se in Z oo l ogy in Unive r siti e s and C oll e g e s It .

h a s b e e n p re p a r e d e sp e ci a lly f o r th e z o o l o gic a l p a r t o f th e w or k
in Ge n e ra l B io l o gy a t the Unive rs ity o f M ichig a n a nd is e x ,

p e c te d t o suppl e m e nt t h e o ne l e ctu re a n d f o u r h o u r s o f l a b or a

tory w o r k pe r we e k du ring th e fi r st h alf ye a r .

N o te xtb oo k now o n th e m a r k e t c ov e r s th e fi e ld o f the intro

d ucto ry co u r s e in Z o o l o gy a s it i s giv e n a t s e v e ra l o f th e l e a ding
Uni ve r siti e s The se c o urse s de al with inve r te b ra te typ e s o nly
.
,

b e ing fo ll o we d by a co ur s e o n v e r te b ra te typ e s during the s e co nd

h alf ye ar Onl y a f e w a nim als b e l o nging t o th e m ore imp o r ta nt
.

phyl a as vi e we d fro m a n e volutio n a r y s ta ndp o int a r e co n side r e d

, , .

The y a re ho we ve r int e n sive ly studie d in a n e nde a vor to t e a ch

, ,

the fund a m e nt a l p r incipl e s o f Z oo l o gy in a w a y th a t is no t os

p
s ibl e wh e n a s up e r ficia l e xa m in a ti o n o f typ e s fr o m a ll th e phyl a

is m a de F ur the r mo r e m o rph ol o gy i s not sp e ci ally e mph asiz e d

.
, ,

but i s co or di n a te d with physi ol o gy e co l o gy a nd b e h a vi o r a nd

, , ,

se rv e s to illu s t r a t e by a c o mp ara tiv e s tudy th e p r o b a bl e c o u r se o f

e v o luti o n T h e a nim al s a r e no t t r e a t e d a s in e r t o bj e ct s f o r
.

di sse ctio n but as living o r gani sm s wh o se a ctiviti e s ar e o f funda

,

m e nt al imp o r t a n ce N o a rgum e nt s I b e lie v e a re n e c e ssary t o

.
, ,

justify thi s m e tho d of p ro ce du re sinc e th e so c all e d typ e

,
-

co ur s e , de ve l op e d wi th th e p r obl e m s o f or ga ni c e voluti on in
mi nd a nd de aling with dyn a mic a s w e l l a s st a ti c ph e n o m e n a i s
,
,

re c o gniz e d by m o st t e ach e r s a t t h e p r e se nt tim e a s th e b e st

m e thod o f int ro ducing yo ung stude nts to Z ool o gy .

The a uth o r h as t a ught a co u rs e simil a r t o th a t ju s t o utlin e d a t

th e Uni ve r sity o f Wisco nsi n a nd t o thr e e s ucc e s s iv e cl a sse s a t th e

M 41
vi PREFACE

Univ e r sity of M ichiga n and h a s b a se d his p re s e nta ti o n not al on e

,

up o n h i s o wn e xp e ri e nce s but h a s p ro fit e d by the e xp e ri e nc e s of

,

o th e r in s t r uct o r s with wh o m h e w or k e d .

N o o rigin a lity i s cl a im e d f or thi s t e xtb oo k T h e m a t e ri a l

.

co nt aine d in it h as b e e n coll e cte d fro m m a ny s o u rce s a nd an ,

a tt e mpt h a s b e e n m a d e to inc o rp o ra t e th e r e s ult s o f th e l a t e s t

inv e s tiga tio ns T h e m aj o rity o f th e figure s h a ve b e e n b orr o we d

.

fro m o the r t e xtb oo ks a nd fr o m origin al a r ticl e s in s ci e ntific

p e ri o dic als T h e a utho r is inde bte d to th e M a cmill a n C o m
.

p a ny f o r th e m a j o r ity o f th e se illust r a ti o ns He wi sh e s a ls o .
,

to th a nk th e f o ll o wing f o r th e u s e o f ce r t a in figu re s : He n r y
Ho lt a nd C o 1 4 9 1 5 6 ; N e w E r a P r inting C o
.
—
, 26 6 2 63 ; .
, , ,

A I Roo t C o
. . .
, Sci e nc e P r e s s 9 1 1 0 9 1 1 0 1 1 1 1 1 6 ;
, , , , ,

Unite d St a te s D e p a r tm e nt o f Ag r icultu re 1 3 5 1 3 7 1 4 1 , , , ,

1 4 2 ; Wist a r In s titut e o f An a t o my a nd B i o l o gy 1 2 13 25 , , , ,

9 4 . In e v e r y c a s e th e a uth o r s n a m e f o ll’
o ws t h e l e g e nd und e r

t h e figu s r e F igu e s 1 2 3 3 1 4 9 5 1 54 1 2 3 a nd 1 5 7 we re
. r ,
—
, , , , , ,

d ra wn by M r Ge o r ge M C u r ti s unde r th e a utho r s di re ction

. .
’
.

T h e s o u rce s o f info r m a ti o n c o uld no t w e ll b e a ckno wl e dg e d

dire ctly so th e titl e s o f th e mos t imp o r t a nt a nd e a sily a cce ssibl e
,

b ooks a nd o r igin al a r ticl e s ha ve b e e n a rr a nge d a t th e e nd of the

b oo k unde r the he a dings o f th e v ari o us chapte rs Th e titl e s .

a r e numb e r e d a nd are r e f e rr e d t o by numb e r in m a ny pl ace s in .

th e t e xt . It i s hop e d th a t thi s li s t will b e he lpful to te a che rs ,

a nd th a t stud e nt s m ay b e e nc o u r a g e d t o s uppl e m e nt th e i r o wn

o b se r v a ti o n s a nd di s c o v e r i e s by co n s ulting th e origin a l p a p e r s

de a ling with the to pics unde r di s cus sio n .

In a fi r st co u r s e in Z o ol o gy stud e nts e nco unt e r a l a rge ,

numb e r o f sci e ntific t e r ms whi ch ca us e th e m mo re o r l e s s

c o nfu s i on F or th is r e a s o n a gl o ss a r y h a s b e e n incl ud e d
. .

It give s th e m e a ning o f th e mo st imp o r t a nt t e r ms u s e d a ,

k e y t o th e i r p r o nunci a ti o n a nd sh o ws th e i r d e r iva tio n

,
.

In m a king a ckn owl e dgme nt s it i s ne ce ssar y to e xpl ain tha t

th e p r e se nt a uth o r pl a nn e d th e b oo k with Dr A S P e ar s e a nd . . .
,

th a t the l atte r was to have p r e pare d h alf of the manu script

 vii

La ck tim e h o we ve r p re ve nte d him f r o m c ompl e ti ng his share

of , ,

of the w or k C h a pt e r s I II a nd III w e r e w r itt e n by D r P e a r s e ;

.
, ,
.

C h apt e r s I a nd II w e r e th e n r e vi se d j ointly a nd C h apt e r III wa s ,

compl e te d by the p re se nt a uthor a l o n e I wish to e xp re ss my .

gra titude to Dr P e ars e f or a ll o wing m e to u se th e fi r st thr e e

.

chapte r s f o r r e a d ing m os t of th e r e m aini ng ch a pte r s a nd f or

, ,

m any v alua bl e cr itici s ms during th e p r e p a r a ti o n o f th e m a nu

sc ript .

In a n e nd e a v o r to m a k e th e t e xt a s n e a r ly c or r e ct a s pos
sibl e th e a uthor s e nt th e va r i o us ch a pt e r s to sp e ci a lists on
,

th e subj e ct m a tte r t re a t e d th e r e in
-
The s e g e ntl e m e n h ow .
,

e ve r a r e in no w a y r e s p o n s ibl e f or a ny e r r or s th a t m a y h a v e
,

e sc a p e d n o tic e T h e a uth o r is g r a t e f u l t o P r o f e ss o r J a cob

.

R e igh a r d f o r c r iticisms du r ing t h e p r e p a r a ti o n o f th e w o r k

a nd f o r re a ding C h a pt e r X I V t o P r o f e ss o r S J Ho lm e s f o r ,
. .

r e a ding th e e nti r e m a nu s c r ipt t o R e v H F H e gn e r f o r r e a d ,

. . .

i ng C h a pte r s I VIII t o P r o f e s s o r C M C hi l d f o r r e a ding

—
,
. .

C h a pt e r s I III t o P r o f e s s o r F G N o vy a nd D r H H u s f o r
—
, . . . .

re a ding C h a pt e r s I V—VI t o D r 0 P D e lling e r f or r e a ding ,

. . .

C h a pt e r I V to P r o f e s s o r L L Wo o d r u ff f o r r e a ding C h a pte r
, . .

V to P r o fe ss o r T H M o ntg o m e r y J r f o r r e a ding C h a p
, . .
,
.
,

te r VI I to D r E R D o wning f o r r e a ding C h a pt e r VIII to

,
. . .
,

P r o f e ss o r W C C u r ti s f o r r e a ding C h a pte r IX t o P r o fe ss o r
. .
,

J. P e r cy M o o r e f o r r e a ding C h a pt e r X t o Dr A E Or t m a nn ,
. . .

f o r r e a ding C h a pte r XI t o Dr E F P hillips a nd M r M a x , . . . .

P e e t f o r r e a ding C h a pt e r XII a nd to D r A G R uth v e n a nd ,

. . .

Dr H Hu s f o r r e a ding C h a pte r X I V
. . .

ROBER T W . HE GN E R .

[U NE 1 , 1 9 1 0.
 T A B L E OF C O N T E N T S

C HAP TE R I
IN T RODU C T I ON
De fi niti o ns , I C las s ifi cati on, 3 .

C HAPT ER II

P H ENOM E NA OF LI F E .

Or i gi n o f li fe 8 ; C h a r a ct e r i s ti cs o f livi ng o r
, g anis ms , lo ;
P l a nt s a nd a ni m a l s co m pa r e d , 1 7 ; P h s i ca l b a s i s y of li fe : pr o

to plas m , P h i o—c h m ical e xpl a n ti o n o f th e p h e no m e na

a
1
9 ; ys c e

of lif e , 23 .

C HAPTE R III
C EL L A ND C ELL T HE OR Y
Mo rph o l ogy a nd h
p ys i o l o gy o f th e ce ll, 26 ;
29 ; C e ll th e ory , 34 .

C HAPT E R IV

AM EBA

C HAPT E R V
PARAM EC IUM

C HAPT E R VI
OT HE R P R OTOZ OA
C l as s i fi ca ti o n of th e g
P ro t o zo a, 80 ; E u le na, 8 2 ; Plasmo
di u m , 8 8 ; Vo lvo x glo b a t or a nd i ts all i e s ,
9 2 .

C HA P T E R VI I

INT RODU CT ION TO T HE ME TAZ OA

C e llular di ff e r e ntia ti o n : T i ss ue s , l oo ; Ge rm ce lls , 1 02 .

Em br yo l ogy, 1 0 7
x T AB LE OF C ON T ENTS

C H AP T ER VI II

HYD RA A ND C CE LE N TE R A T E S IN G E N E R AL

Hyd ra , 1 16 ; C oe l e nt e ra t e s i n g e ne ra l , 13 9 .

C HAP T E R IX

S P ON G E S, F I AT W O R M S A N D R OU N D W o R M s
,

S po nge s G ra nti a S po ng e s i n ge ne ra l 1 44 ; F l a t Wo r m s ,

P l a na r i a F l a t W o r ms i n ge ne ra l 1 5 3 ; R o u nd W o r m s ,

As car i s — R o u nd Wo r m s i n ge ne r a l 1 60 ,
.

C H AP T E R X
T HE E A R T HWO RM AND A N N E LI D S IN G E N E RA L

Th e e ar th wo r m ,
1 64 ; Anne li d s in ge n l e ra , 1 90 .

C HA P T E R X I

T HE C R A YF I S H AN D A R T H R OP O D S IN G E NE R AL
The cra yfi hs ,
1 93 ; Ar th ro po d s i n g e ne r a l , 225 .

C HA PT E R X I I

T HE HON E Y BEE AN D B E ES IN G E N ERAL

The h o ne yb e e ,
23 3 ; Be es i n ge ne ra l , 2 60 .

C HA P T E R XIII
H I STOR I C A L ZOOL OGY

C H AP T E R X I V

G E N E R A L C ON S I D E R A T I ON S O F Z OOL O G I C A L FA C T S A N D T HEO R I E S 27 5

H e r e d ity a nd v o lu
e ti n 2 7 5 ; T h e s o ci l l i fe Of ni m als
o ,
a a ,

2 97 ; The a ni m al m i nd , 3 05 .
 LI S T O F I L L U S T R A T I O NS

Pa r t s of h owi ng cili a t r i ch o cys ts

a nd
'

P a r a m e e za s

A se cti o n f ro m th e r e g i o n o f th e m o u th o f P a r a m e ci u m S h o wi ng
th e f or m a ti o n o f a f oo d va cu o l e 66
B i na ry fi ss i o n o f P a r a m e ci u m a u r e le a 66
'

D i ag ra m ill us t r a ti ng th e li f e hi s to ry o f th e m a l ar i al fe ve r pa r as it e 89
Volv ox globa tor be tw e e n 9 6 —9 7

Volw x g loéa tor ae tw e e n 96-9 7

E xa m pl e s vario s k i nds o f ti u e
of u ss s 1 00

F ig s ill us t ra t i ng f o ur d i ff
ur e nt k i nd s o f cl e a v g e
e re a

R e ge ne r ti o n a nd g ra f ti ng i n Hyd ra
“
a

P/zys l a t h e
a z , P o r t g e s e m a n Of-wa r
u u -

R e g ne ti o n Of P l n r a m c l t
'

e ra a a z a u a a

Lo ngitu d i n l v e rti cal s e cti o n th r o g h t h e a nt e r i o r

a u
po r t io n of an

e ar th wo r m
S tage s i n th e e mb yol ogy o f th e e a th wor m
r r

R e ge ne ra ti o n i n th e e a thworm r

G ra f ti ng i n t h e e a th wo m
r r

S e m i -d i g ra m m a ti c vi e w o f i nt e r nal or ga ns and a ppe nd ag e s

a of

rig ht S i d e of a m ale cra fi h

y s

S rf a ce of a cr ayfi s h e gg w ith e m b ryo b e gi nni ng to f orm

u

E m bryo o f th e cr ayfi s h i n t h e N aupl i u s s ta g e

Ol d e r e mb yo o f th e c ayfi s h
r r

He ad o f work e r h one ybe e

Le gs of work e r b e e

g of the work e r be e
S ti n
T h e re s pi ratory sys t e m of th e be e
T h e ne rvous sys te m of th e be e
xi i LI ST or ILLUSTRATI ONS

Larvae and u
p pa of h one ybe e i n th e i r ce lls

Inte rnal o rga ns of larval ho ne ybe e

Polli nati on o f an o rch id ( C p l
z e a zu m ) by a bumb e be e
’

p
'

y r

Aris totle , 3 84-3 2 2 B C

. .

Li nnae us at s ixty , 1 7 07 - 1 7 7 8

C u vi e r, 1 7 69- 1 83 2

W illi a m Harve y 1 57 8—1 667

,

K a r l E r ns t vo n Bae r 1 7 92 —1 8 7 6
,

J o h a nne s Mii lle r 1 80 1 —1 8 58

,

C h arl e s Dar wi n 1 809—

, 1 88 2

Varie t ie s of d o m e s ti c pi ge ons
A N I N T R O D U C T I O N T O Z OOLOGY
A N INT RODUC T ION T O Z OOLOG Y

CHAPTER I

INTRODUCTION

1 . DE FI NI TI ONS

THE sci e nc e of B i ology include s th e sum of hum an k n owl e dge

Wl th re g a r d t o o r g a ni s ms . Th a t b ra nch of bi ol o gy whi ch d e als
with anim al s i s kn o wn a s Z oology; th a t whi ch r e l a t e s to pl ants ,

a s B ota ny . Z ool o gy m ay b e d e fin e d a s th e b o dy of f a cts a nd

d o ct rin e s d e rive d f ro m th e sci e ntific study of a nim als Sci e n .

t i fi c study d e m a nd s a ccu r a t e a n d p a in s t a king o b s e r v a ti o n c a rr ie d

o n with s o m e d e fini te e nd in V i e w T h e sci e ntific study o f
.

z ool ogy a tt e mpt s to ga in a n unde r s t a nding o f th e st ructu re a nd

a ctiviti e s o f a nim a l s . It no t o nly d e al s with th e a nim als pe r s e ,

but al so with th e i r r e l a ti o ns to b o th th e o r ga nic a nd in organic

wo rlds .

T h e d e t ail e d inv e s tig a ti o n o f a nim als h a s le d to th e e st abli sh

m e nt o f a numb e r o f sub sidi a ry z ool ogica l s ci e nc e s s e ve ral Of ,

which a r e b ri e fly o utlin e d in th e n e xt f e w p a rag r aphs .

Sys te m a ti c Z oology is c o nc e r n e d with th e d e sc ripti o n of a nim al

sp e ci e s a nd th e i r arra ng e m e nt a ccordi ng t o a l ogic al pl a n of
cl assifica tio n T h e e x a c t m e aning o f the t e r m spe ci e s is a
.

liv e que s ti o n a t th e p rés e nt tim e a nd sys t e m a tic z o ol o gi sts d o

,

no t a g re e a s t o wh a t ch a r a ct e r i s tics Sh o uld b e us e d in s e p a ra ting

one sp e ci e s f r o m a n o th e r One inv e stig a t o r in this fi e ld give s
the f oll o wing d e fi ni ti o n : A sp e ci e s a s c o nc e iv e d by m o st
,

sy s t e m a ti s t s a t t h e p r e s e nt tim e m a y b e d e fin e d a s a g ro up o f
,

int e rb r e e ding in di vidu als whi ch whil e th e y m a y di fle r m a r k e dly

,

a m o ng th e m s e lv e s e t re se mbl e e a ch o th e r m o r e c l o s e ly th a n th e y
, y
d o th o s e of a ny o th e r g r o up ; th e char a ct e r s th a t distinguish the
2 AN I NT R O DUCT I ON T O Z OOLOGY

gro up b e ing conside rabl e not oblit e ra te d by inte rm e di a t e for m s

, ,
”
a nd inh e r it e d f ro m g e n e r a ti o n to g e n e r a ti o n An o th e r p r o mi
.

n e nt zool ogist whe n a ske d f or a de fini ti o n of a sp e ci e s sai d it

, ,
”
wa s s o m e b o dy s opini on
’
He di d no t m e a n th a t sp e ci e s a r e
.

not r e a liti e s in n a tu r e but th a t just h o w m a ny a r e r e p re s e nt e d

,

i n a p a r ticul ar gr o up m ust u s u ally be de t e r mi n e d by c o mp e te nt

a uth or ity .

Di s tr i b u ti onal Z oology a tt e mpts to a sc e r t a in th e p a s t a nd

p re se nt h abita ts of a ni m a ls a s w e ll a s th e f a ct o r s whi ch c o nt ro l
,

the i r di stributio n ov e r the s u r fa c e of th e e a r th T h e study o f th e .

dist ributi o n o f r e ce nt a ni m a ls is kn o wn a s Z ooge o gr aph y a nd tha t ,

o f f o s sil f o r ms c o nstitut e s a p ar t o f th e s ci e nc e o f P ale ontolo gy .

Ani m al M orph olo gy i s th e sci e nc e of fo r m a nd s t r uctu r e Th a t .

p ar t of it which p e r t a in s to th e gro ss di sse cti o n of o r g a ns is kn o wn

a s Ana to m y ; th a t which r e l a t e s to th e micr o scopic study of
tis su e s a s Hi s tology If ho w e v e r a s tudy i s co nc e rn e d with th e
,
.
, ,

d e ve l opm e nt o f a ni m al s or ga ns or tissue s it is includ e d in th e

, , ,

S ci e nc e o f E m b ryolo gy .

Ani m d P h ysi ol ogy d e als with the functi o ns of th e p a r ts o f

a nim a ls .

Ani m al E colo gy is th e study o f th e r e l a ti o nshi ps of a ni m a ls

t oo ne a n o th e r a nd t o th e i r e nvi r o nm e nt .

Evolu ti on ar y Z oology is c o nc e r n e d with th e o rigi n a nd d e sc e nt

o f th e di ff e re nt sp e ci e s of a nim al s .

P a le o z oolo gy is th e study Of fo s sil r e m ains o f a nim a ls It is .

intima te ly a ss o ci a t e d with a ll th e o the r b r a nche s o f z oo lo gica l

sci e nc e e xce pt th e study of physiol ogic al p r o c e ss e s T h e physi
,
.

o l o gy o f e xtinct fo r ms ca n o n l y b e infe rre d fr o m th e study o f

th e a ctiviti e s o f r e c e nt a nim a ls Owing to th e fa ct th a t fo ssil s
.

a r e usu a lly e mb e dd e d in st o n e t h e m e ch a ni ca l a ppli a nc e s whi ch

,

a re u s e d in studying th e m a r e n e ce s s a rily quit e di ff e re nt f r o m

those e mpl oy e d in th e inv e stiga ti o n o f m o de r n a nim als .

T h e di f f e r e nt b ra nch e s o f z ool o gic a l sci e nc e a r e cl o se ly r e l a t e d

and mutu a lly int e r d e p e nd e nt ; f o r e x a mpl e t h e s tud e nt s o f e v o
,

lu ti on g a in e vi d e n ce c o nce r ni ng t h e a nce s tr y o f ani m a ls from

4 AN INTRODU CTION TO ZOOLOGY

ti ngui sh the m fro m fo ur o the r cl as se s whi ch m ak e up th e

th e
h l m V r te br a ta r a nim als p osse ssing v e r t e b r a l c o lumns
p y u e o ,
.

T h e sci e ntific n a m e o f a ny a nim a l c o n si s t s o f the t e r m s u se d t o

‘

d e sign a t e th e g e nus a nd sp e cie s ; thi s i s co mm o nly foll o we d by

th e n a m e o f th e z ool o gi s t wh o w r o t e th e fi r s t a uth o r it a tiv e d e
s c r ipti o n o f th a t p ar tic ul a r sp e ci e s T h e s ci e ntific n a m e o f m a n .

i s the r e for e w r itt e n Homo s a pi e ns Linnae u s .

Th e foll o wing c o n ci se syn op si s o f th e chi e f g r o ups o f a nim a l s ,

t og e the r with a b ri e f char a cte r iz a ti on of e a ch of th e fif t e e n phyl a ,

will be o f v alue to the s tude nt f or s ub se que nt r e fe r e nce T h e .

t e r m s us e d in de sc ribing the se sub di vi si o n s will not p e rh ap s be

a lt o g e th e r und e r st oo d by t h e b e ginn e r but the i r m e aning will ,

b e m a d e app a re nt in s u cce e ding ch apt e r s It will suffi ce t o .

say h e r e th a t in a g e n e r a l w ay the cl assific a ti o n o f a nim al s

, ,

de p e nds chi e fly up o n th e ch ara cte ri s tic s whi ch are co nt ras te d

in Tabl e I .

T AB LE I

CHI E F C HAR AC TE R I STI C S U SE D I N SE P AR ATI N G TH E P H L A OF AN I MAL S Y

C ON TR ASTE D

1 .
B od y co m po se d o f o ne m i cr o sco p i c B od y co m p o se d of m any ce ll s .

un it ,
th e ce ll .

2 . B ody f or m e d f ro m t wo p rl m ary B o d y f or m e d f ro m th r e e p r i m a ry ce ll
ce ll l aye rs i e d iplo b l a s ti c
,
. . . l a ye r s i e t r ip l ob l a s ti c
,
. . .

3 . B o d y ra d i a lly sym m e t r i ca l i e ,
. . B o dy b il a t e r a lly sym m e t r i ca l i e ,
. .

pa r t s ( a nti m e r e s ) ra d i a ti ng f ro m p a r t s a r r a nge d i n p a i r s o n e ith e r

a ce nt ra l a xi s S i d e o f a ce nt ra l
.
pl ane .

4 . Di ge s tiv e ca vit y with o nl y o ne Dige s tive ca vit y with t wo Ope ni ngs ,

o pe ni ng . th e m o u th a nd a nu s .

5 . A ca vi ty t h e coe l o m b e tw e e n th e
, No coe l o m p re se nt .

d ige s tive t u b e a nd th e bod y w a ll .

6 . B o d y se gm e nt e d i e co m po se d o f ,
. . Bod y u nse gm e n e d t .

a li ne a l se r i e s o f p ar t s ca ll e d ,

m e t a m e r e s o r som it e s .

figur e s foll o wing the diagno si s o f e ach phylum indic a te

The
t h e numb e r o f r e c e nt sp e ci e s ; th os e in p a r e nt h e se s th e numb e r o f ,

fos sil sp e cie s known to r e p re s e nt it .

 INTR OE UCTI ON

P RIN C IP AL P HYLA OF ANI M AL KI N G D OM I

1 . Singl
P r o to z oa .
e -c e ll e d a nim als ; Ofte n col oni al ; sp e r m
and e gg c e lls usu ally w a nting 44 00 .

2 P o ri f e r a
. Sp o ng e s
. Tripl obl a stic r a di ally sym m e tr i
.

ca l numb e r of a ntim e re s v ari abl e ; b o dy w all p e r m e a t e d by

,

innum e r abl e p or e s a nd usu ally supp or te d by a sk e l e t o n of spicul e s .

6 00

3 . C ce le nte r ata J . p o lyps

e llyfi sh e s ,
a nd c o r a ls D ipl o , .

bl astic ; ra di a lly symm e t r ic al with f o ur o r six a ntime re s ; Singl e,

g ast ro v ascul ar ca vity ; no a nus ; b ody w all co nt ains p e culi ar

-

b o di e s known as n e m a to cysts o r stinging c e lls 3 000 .

4 C te n oph or a
. Se a w a lnuts or c o mb j e lli e s
. Tripl o bl a stic ; .

r a di al c o mbin e d with bil a t e r a l s ymm e t r y ; e ight r a di a lly ar ra ng e d

r o w s o f p a ddle pl a t e s ; a nus p r e s e nt 1 000 ( no f o s s ils ) . .

5 . P la ty h e lmi nth e s F l a t w o r
. ms T r i plo b las t i c b i l a t e r.a lly
symm e trical ; singl e g a stro-va sc ular c avity ; no a nus ; p re s e nc e
o f c oe l o m d o ubtful 1 6 00 ( no fo ssils)
. .

6 N e m a th e lmi nth e s
. R o und w o r ms. Tr ipl obl a stic ; b i .

l a t e r ally symm e t ric al ; p o ss e s s a tubul ar dige stive syste m with an

a nus ; c oe l o m p r e s e nt 1 000 ( no fo ssi l s)
. .

7 . R o ta tori a — Wh e e l a ni m alcul e s T ripl obl astic ; bil a te rally

. .

symme t ric al ; c oe l o m p re s e nt ; a p a i r o f cili a t e d disk s on the a n

te ri o r e nd 35 (
0 .no f o ssil s) .

8 B ryoz oa
. M o ss a nim als Tripl obl a s tic ; bil a t e r a l sym
. .

1
Z l gi ts d n t g
oo o s o o t th n m b
a re e a s f p hyl i nt whi h th
o e u ni m l
er o a o c e a a

k i ngd o m ho u l d be d ivi d e d So m e a u tho r itie s r e cogniz e only e i ght whil e

s .
,

o th e rs m a i nt a i n th a t th e re Sh o u l d b e a s m a ny a s tw e nt o r e v e n m o r e T w o
y ,
.

su b k i ngd o m s a r e ge ne r a ll y re co gni z e d
-
P r otozoa ( P h yl u m 1 ) a nd M e ta zoa
( P h yl a 2 Re ce ntl y m a ny z oo l o gi s t s h a v e co m e to b e li e ve th a t th e
s po nge s ( P h yl u m 2 ) sh o u l d b e s e a r a t e d f r o m o th e r m e t a z oo ns a nd ca ll e d
p
t he P a r a zoa T he a ni m a l s i n P h yl a 1 —1 4 a r e u s u a ll y ca ll e d i nve r te br a te s to
.

d i s ti ngu i sh th e m f ro m th e ve r te br a te s P h yl a 1 4 a nd 1 5 with a f e w a ni ,

m a l s o f so m e wh a t d o u b t f u l re l a ti o nshi p s a r e co m m o nl y p l ace d t o ge th e r
,

u nd e r th e gro u p C hor d a ta a nd a re k no w n a s ch o r d a t e s —
P h yl a 2 4 i ncl u d e
, .

a nim a l s wi th o u t coe l o m s a nd a r e k no w n a s Acce lom a ta i n co nt ras t to ph yl a

5 5 whi ch co nta i n a ni m a l s with a coe l o m a nd a re t e rm e d Cce loma ta

-1
.
,
6 AN INTRODU CTION TO ZOOLOGY

m e try usu ally di stor te d by co l o ni al habits ; ci li a t e d t e nt a cl e s

a b o ut t h e m o uth ; coe l o m a nd a nus p r e s e nt 7 00 .

9 Br ach i opod a
. To ngu e o r l a mp sh e ll s T ripl obl astic ;
. .

bil a t e rall y symm e t rical ; coe l o m p re se nt ; a nus p r e s e nt or abse nt ;

cili a t e d a r ms a r ound th e m o uth ; b ody c o v e r e d by a sh e ll com
p ose d o f dorsa l a nd v e nt r al va lve s 1 00 .

10 .E chi nod e rm a ta Sta r fi sh e s se a cucumb e r s se a u r chins

.
, , ,

se a lili e s Tri plobla sti c ; r a di ally symm e tric al u s ually fi ve

.
,

a ntim e r e s ; c oe l o m w e ll d e v e l op e d ; a nus usu ally p re s e nt ; l o c o

m o ti o n in m a ny Sp e ci e s a cc o mplish e d by ch ara cte ri s tic o rg ans

kno wn a s tub e f e e t ; a spiny sk e l e to n of calca re o us pl a t e s ge ne r
a lly c o v e r s t h e b o dy 2 5 00 .

1 1 . Anne li d a Joint e d wo rms Tripl obl astic ; bil a te rally

. .

symme trical ; c oe l o m w e ll de ve l op e d ; a nus p re se nt ; s e gme nte d ,

s o mite s sim i l ar 2 500 .

1 2 . Ar th r opo d a C r a bs ins e cts spid e r s c e ntip e d e s scor

.
, , , ,

pi o n s ti cks Tripl obl a s tic ; bil a te rally symm e trical ; a nus

,
.

p r e se nt ; coe l o m p oo r ly de ve l op e d ; s e gm e nte d so mite s u s ually ,

m o re o r l e ss di ssimil a r ; p ai re d j ointe d a pp e nd age s p re s e nt on all

o r a p a r t o f th e s o mit e s ; chitin o u s e x o s k e l e t o n .

13 . M ollu s ca C l a ms
. sn ai ls d e vi lfi sh e s Tr ipl o bl astic ;
, ,
.

bil ate rally s ymm e t rical ; a nus a nd c oe l o m p re se nt ; no se gme nta

ti o n ; she ll us u ally p re s e nt ; th e ch ara ct e ristic orga n is a ve nt ral
muscul ar foo t .

14 . T u ni ca ta Se a squi rts
. T ripl obl a stic ; bil a t e rally sym
.

m e tric al ; c oe l o m and a nus p re se nt ; fo r m s a ccul a r o r b arre l

h
S a p e d ; t w o a p e r tu r e s th r o ugh which w a t e r cu rre nt s p a ss in a n d
o u t ; f r e qu e ntly c o l o ni a l ; all p a s s th r o ugh a t a dp o l e lik e l arv a l
-

t
s a e g du ring wh i ch t h e b o dy i s supp o r t e d by a r o d lik e st r uctu re -
,

t h e ch o rd a .
3 00 ( n o f o ss il s) .

1 5 . Ve rte b ra ta B il a t e ra lly symm e t ric a l ; c oe l o m w e ll d e

.

v e lope d ; a nu s p re s e nt ; ch or d a usu a lly m o r e o r l e ss fully r e pl a c e d

by a s e ri e s o f b ony v e r te b rae ; m a ny sys t e ms o f int e r n al org a ns
s e gm e nt e d ; t w o p ai rs o f app e ndage s u su ally p re s e nt ; ce nt ral
n e rv o us syste m always d ors al to dige stive tub e .
 I NT R OD U CT ON I 7

FI G . 1 .

Figu r e 1 is int e nde d to Sh o w the r e l a ti onshi p s o f the di ffe re nt

phyl a of the ani m al ki ngdo m to e ach o the r T h e P r o to zo a are
.

b li ve d to h a ve giv e n ri se t o th e o th e r gr oups T h e ve rti ca l

e e
.

he ight o f e a ch phylum up on the p age re p re se nts the de gre e o f

st ru ctu r al sp e ci aliza tio n whi ch it has a tta in e d .
 C HAP T ER I I

P HEN OM ENA OF LIFE

1 . OR I GI N OF LI FE

THOSE who are inclin e d towa r d the study of living org anisms
will find an a bund a nc e o f m a t e ri a l o n e ve ry ha nd Da rwin .

r a is e d fi v e hund re d a nd thi r ty -s e v e n pl a nts f r o m thr e e sp o o n

fuls o f mud whi ch h e Scoop e d up a t th e e dge o f a p o nd A r e ce nt .

n a tura li s t h as c o ll e ct e d fr o m fo u r squ are fe e t o f me a do w o ve r a

th o usa nd o bj e ct s r e p r e se nting living a nim als or the i r re m ains .

If we go to th e de pth s o f th e se a o r to the tops of th e m o unta ins

, ,

o r e v e n p e n e t r a t e t h e d ar kn e s s o f th e c a v e r ns b e n e a th th e e a r th
,

w e a lw a ys find f o r ms o f a nim a l life w hi ch h a v e b e c o m e a d apt e d

t o a lm o st e v e ry c o nc e iv abl e co nditi o n o f e xist e nce .

T h e g re a t a bund a nce a nd ra pid m ul tiplic a ti o n o f living a nim a ls

le d t h e a nci e nts t o f o r m ul a t e s o m e cu r i o us th e or i e s with r e g a r d
t o th e o r igin o f lif e . E v e n a t this e nlight e n e d d ay it m a y b e
he lpful to us to re vi e w so m e o f th e i r p e culi ar ide a s M any o f .

t h e sch ol a r s o f th a t p e r i o d b e li e v e d th a t e v e r y a nim a l w as o rigi

n a lly cre a t e d by di vin e p r ovide nce A f a vo rit e the ory with
.

th o s e wh o did no t a cc e pt th is id e a o f spe ci al cr e a ti ons wa s th a t o f

,

s ponta ne ou s ge ne r a ti on . Thi s w as a unive r s ally a cce pt e d dogm a

fro m th e tim e o f the Gre e k phil o soph e r Ari s t o tl e (3 84 3 2 2 —

until 1 668 whe n R e di ov e r th re w it with his c are ful obs e rva ti ons .

Anci e nt n a tu ralists b e li e ve d th a t fr ogs and t o a ds a rose f ro m th e

muddy b ottp m o f p o nds unde r th e influe nce of th e sun and tha t ,

ins e cts origin a t e d fr o m d e w Tho se who did not a cce pt th e se

.

a nd o th e r e qu a lly a b s u r d vi e ws w e r e subj e ct t o r idicul e by th e i r

co nt e mp ora r i e s B e for e R e di p e r for m e d his Simpl e but r e m ar k

.

a bly e ff e ctiv e e xp e rim e nts no o ne h a d th o ught o f t e sting t he

,

8
 P HEN OM EN A OF L FE I

ve r ity of the th e or y of sp ont a n e o us ge n e r a ti o n R e di p rove d con

.

clu si v e ly th a t m a gg o ts n e v e r r is e d e novo f r o m d e c a ying fl e sh a s ,

h a d b e e n b e li e v e d up to hi s tim e He e xp o s e d m e a t in t hr e e j a r s
.

unde r dive r s e c o nditi o n s ; o ne w a s l e ft op e n the se co nd w as

,

co ve re d with g a uz e a nd th e t hi r d w a s co v e re d with p a r chm e nt

,
.

Th e m e a t d e ca ye d in a ll o f t h e j a r s but m a ggo ts a pp e a r e d only

,

in the m e a t in t h e one which h a d b e e n l e ft e ntir e ly uncove re d .

F li e s w e re a tt r a ct e d by th e d e ca ying m e a t in t h e se co nd j a r but , ,

b e ing un a bl e t o e nt e r l a id the i r e gg s o n t h e ga uz e whe r e th e y

, ,

we r e found by R e di N 0 e ggs w e r e l aid on th e thi r d j ar f or the

.
,

p a r chm e nt di d no t all o w o dors to e s c ap e By the s e a nd o the r

.

e xp e r im e nts th e th e o r y o f sp o nt a n e o us g e n e r a ti o n f e ll int o dis

r e put e ,
but m a ny of i ts a dh e re nts still m a inta in e d tha t e ve n
tho ugh so m e a nim al s might no t o rigin a te sp o nta n e o usly th a t ,

fa ct di d no t p r e clude th e p o ssibility o f o the r s do ing SO .

Su ch dis se nt e r s m a d e th e i r l as t st a nd wh e n th e mic ro sc op e
r e v e a l e d th e sw a r m s o f minut e a nim a ls whi ch h a d b e e n p r e v i o usly

unkn own In one o f th e i r e xp e r im e nt s h ay w a s b o il e d in w a t e r

.

until a ll the life h a d b e e n kill e d ; the liquid w a s the n filte re d OE

a nd pl a c e d in a tightly s topp e re d b o ttl e T h e s ubs e qu e nt
.

a pp e a r a nc e o f a scum o f living o r g a nism s in such infusi o ns co n

vi nce d th e m th a t lif e h a d a ri se n sp o nt a n e o u sly In 1 7 7 5

.
,

Sp all a nz a ni p r o v e d th a t wh e n p r op e r p re ca uti o ns w e r e o bs e r v e d
no life d e v e l o p e d in a cul tu r e o f thi s kind . He p r e v e nte d th e
e nt ra n ce o f a i r int o th e v e s s e ls c o nt a ining t h e nut r i e nt sohi ti o n

by h e r m e tically s e a ling th e sl e nde r n e ck s of hi s fl a sks in fl a m e .

T h e fl a sks w e r e th e n pl a c e d in b oiling w a t e r f or th r e e qu ar t e r s
o f a n h o u r a ll g e r ms c o nt a in e d in th e m b e ing d e s t r oy e d
,
No .

life app e a r e d in n u t ri e nt fluids thus t re a t e d D e spit e this

.

d e m o n s t r a ti o n th e qu e s tio n c o ntinu e d t o b e a git a te d f or m a ny

y e ar s and it wa s not until th e tim e of P a ste u r ( 1 8 64) a nd Tynd all
,

m
( 1 8 76) th a t th e old th e o ry o f sp nt n us g n ti n
o a e o e e ra o w a s co

p l e t e ly o v e r th r o
. wn
At th e p r e s e nt tim e w e a r e a bl e t o a fli r m th a t no livi ng or g a n
ism is kno wn to o rigin a te e xc e pt f ro m so m e o the r p re e xi s ting
IO AN INTRODU CTION TO ZOOLOGY

or ga nism a nd Pr e y e r s a pho r ism all life f ro m life i s unive r

,
’

s ally a cce pte d B u t tho ugh in th e light o f ou r p re s e nt kn o wl e dge

.

a ll a r e willing t o a dmit t h e t r uth o f t hi s a ph o ri s m th e re ar e th ose ,

wh o h op e th a t th e a dv a nce o f sci e nti fic kn o wl e dg e m a y s o m e tim e

in the futu re e n abl e us to cl e a r up the p robl e m o f the b e ginnings
o f life whi ch s till re m a ins uns o lv e d .

C HARAC TERI STI C S OF LI V I N G OR GANI SMS

All th e bj e cts tha t c om e withi n th e ra nge o f hum a n e xpe ri

o

e nc e m a y b e g ro up e d int o tw o cl a s s s e living a nd no n-living

, .

Alth o ugh o ur ide a s c o n ce r ning th e o rigin o f living or g a ni s ms

c a nn o t go b e y ond th e a s se r ti o n tha t the y a lwa ys co me fro m o th e r
living b odi e s ye t th e cha ra ct e ri s tics tha t se p a ra t e th e tw o cl a sse s
,

ca n b e s t a t e d in a v e r y d e fini t e w a y Living or g a ni s ms a r e
.

ch a ra cte riz e d by ( 1 ) d e finite size ( 2) de finite fo rm (3 ) de fini te

, ,

e l e m e nt a ry c o mp o siti o n ( 4) d e fini t e or g a niz a ti o n ( ) g ro wth by

, 5 ,

intussu s ce pti o n ( 6) re p r o ducti on and ( 7) i rrit abili ty

, , .

( 1 ) S iz e — I f o ne we r e a sk e d th e qu e s tio n Are o rg ani sms

‘

.
,

unifor m in size ? he w ould d o ubtl e ss a nswe r N o "but o n m ore ,

c are ful c onside ra tio n it w o uld b e co m e app ar e nt tha t a ltho ugh a ,

wide ra ng e o f v ari a tion e xis ts ye t th e di ffe r e nc e s in Size a re con

,

fine d withi n ra the r de finite limits C e r t a in p ar asite s fo und in

.

hum a n bl oo d which can o nly b e s e e n with th e highe s t p o we r s of

,

th e micr o sc op e r e p r e s e nt th e sm a ll e st a nim a l s kn o wn ; th e wh a l e
,

is the l arg e s t Altho ugh th e di s c re p a ncy b e tw e e n the se a nim a ls

.

is e n orm o us ye t th e ra ng e of v a r i a ti o n illu s t ra te d by the m h a s a

,

finite limit On the o th e r h a nd th e siz e v ari a ti o ns oi inorg ani c

,

subst a nc e s a re infinite ; f o r e x a mpl e wa te r i s a lwa ys re cognizabl e

,

as such wh e th e r it e xists in th e f o r m o f minut e p ar ticl e s o f v a p o r

or r e a ch e s th e dim e nsi o ns Of th e P a cific Oc e a n Alth o ugh a .

wide r a ng e of v a ri a tio n o ccurs a m ong a ni m als in ge n e ra l the in di ,

vi d u als o f any giv e n sp e ci e s ar e p ra ctic all y e qu a l in siz e In .

m any inst anc e s it i s obvi o us th a t the size o f a c e r t ain sp e ci e s

fits it f or th e conditi o ns unde r which it live s T he whal e could .

no t ch a nge pl ac e s with one of th e mic ro-o rg ani sms fo und in d e

1 2 AN INTR ODU CTION TO Z OOLOGY

T he l m e nt s con s tituting living substa nce for m c o mpl e x

e e ,

o r g a ni c c o mp o und s whi ch e xhi bit th r e e s t r iking p e c ul i a r iti e s :

( 1 ) th e y a r e in m o s t c a s e s l abil e s ub s t a nc e s i e th e y a r e u n ,
. .

s t a bl e a nd m a y b e b r o k e n up int o simpl e c o mp o und s or synth e ,

siz e d into m ore co mpl e x m ol e cule s ( 2) ca rb o n is a lways a n i m

,

p or t a nt c onstitu e nt a nd ( 3 ) the i r imp or t a nt e l e m e nt s ar e all

,

no n-m e t a llic .

T h e ch e mic a l c o mp ositi o n o f living m a tt e r is hi ghly v a ri abl e .

A c o mpl e t e a n a ly sis o f r e p re se nt a tiv e s o f a ny tw o sp e cie s w o uld

n e v e r giv e ide nti ca l r e s ults ; a nd e v e n y o ung a nd o ld in di viduals
o f th e sa m e sp e ci e s w o ul d no t a g re e N e v e r th e l e ss a ll a ni m a l s
.
,

Sh o w a re m ar k a bl e unif or mity in ch e mic a l c o mp o siti o n So m e .

a nim a ls might b e c o n side re d e xce ptio n s to thi s ge n e ra l r ul e ;

f o r e x a mpl e a cl a m up o n a n a ly si s w o uld Sh o w a l arg e p e r c e nt a ge
,

o f c a lcium c a r b o n a t e o wing t o th e p r e se nc e o f th a t s a lt in its

she ll T h e she ll how e v e r i s r e a lly no t a p a r t o f the living sub

.
, ,

s t a n ce ,
but re p re se nts an inor g a nic a c cr e ti on th a t h as form e d
a ro und th e b o dy .

(4 ) O r ga niz a ti o n .
— A lth o ugh living o r g a ni sms e xhibit g r e at

dive rsity in s t r ucture a c a r e fu l e x ami n a tio n of all th e p ar ts

,

p rove s th a t th e fund a m e nt a l e l e m e nt s a r e e sse nti ally th e s a m e

in e v e ry individu a l All pl ant s a nd anim als a r e comp o se d o f
.

s i m il ar mi cro s c o pic unit s c a ll e d ce lls which a r e a r ra ng e d in a

, ,

ch ara ct e r i stic m a nn e r f o r e ach sp e ci e s Thu s th e b o dy o f a ve ry

.

simpl e org a nism m ay co n si s t o f o nly a s ingl e c e ll wh e r e a s tha t ,

o f a c o mpl e x or g a ni s m m a y co nt a in billi o ns C e lls m ay b e .

c o mp a r e d with b ricks which a r e u se d to build ho us e s B ricks .

m ay Sh o w v a r i a ti o ns in f o r m col o r a nd c o mp ositi o n but h av e

, , ,

c e r ta in ch ara ct e ri s tics which m a k e the m e a si ly r e c ogniz abl e

wh e the r the y are se p a ra t e o r built into wa lls In a simil a r w ay .
,

tho ugh individu al ce ll s Show min o r di ffe r e nce s the y a gr e e in ,

fund am e nt al C h ara ct e ristics and m ay r e a dily b e ide ntifi e d e ve n

, ,

whe n constitu e nt s o f a m a ny c e ll e d ani m al - .

Whe the r an o rg ani sm i s u ni ce llul ar o r multic e llul a r it a lways ,

co n sis t s o f a numb e r of unlike p ar ts whi ch a re incap abl e o f inde

 P H ENOM EN A OF L FE I I 3

p e nde nt e xiste nce Wh e n t he e ss e nti al p ar ts o f a li ving a nim al

.

a r e s e p a r a t e d f ro m o ne a n o th e r the y s oo n l ose the i r ide ntity

,

a nd ultim a t e ly di sint e g r a t e int o simpl e in org a nic co mp o unds .

N on living b o di e s h a v e no co mm o n unit o f st r u ctu re whi ch is

-

co mp arabl e to the c e ll and the i r p a r ts Sho w no int e r de p e nde nce

,
.

5 ) G r o w th in living or g a ni s ms is usu a lly by i nt u ss u s ce

pti on ,

i e by th e a dditi o n o f ne w p a r ticl e s a m ong th o s e a l r e a dy p re s e nt ;

. .

thi s re sults in a swe lling o f e v e r y p ar t and d oe s no t n e ce s sit a te

,

a ch a ng e in for m . N o n living b odi e s usu ally g ro w by accr e ti on

-
,

i e th e a ddi ti o n o f succe ssiv e l a ye r s o n t he o ut sid e

. . .

Gro wth in a ny living thing inv o lve s a co mpl e x se r i e s o f ch a ng e s .

The che mica l c o mp o unds whi ch m a k e up t h e b o di e s o f a nim a ls

a r e e xt r e m e ly l a bil e ; the y a re co nst a ntly b re a king do wn into
S impl e r subst a nce s o r b e co ming m o r e c o mpl e x by th e a dditi o n

o f ne w m a t e r i a ls . The re is no tim e during the life o f a ny in di

vidual whe n e l abo ra te che mica l r e a cti o ns a re no t t a king pl a c e .

M e tab oli s m is th e t e r m us e d to includ e t hi s g re a t c o mpl e x o f

ince ss a nt ch a nge s Those p r o ce ss e s which u se e n e rgy t o build
.

up co mp ounds a re s aid to b e a na boli c; th ose which de st roy sub

st a nc e to p r oduc e e n e rgy are t e r me d ka taboli c .

Anim a ls a r e p r im ar ily k a t a b o lic o r g a ni sms . The y ca nno t

m a ke organic comp o unds fro m simpl e ino rga nic s ubst a nce s ; in
thi s r e sp e ct the y di ffe r fro m pl a nt s which m a nufa ctu r e st ar ch
fro m ca rb o n di o xide a nd wa te r Since a ni m al s must h a v e o rg a nic
.

f oo d it foll ows tha t pl ant p ro ducts a re n e c e ss ary B e fo r e a nim a l

, .

gr o wth is p ossibl e foo d mu s t b e co nve r te d into living sub s t a nce

,
.

B y th e p ro ce ss o f dig e sti o n foo d i s p r e p a r e d f o r a b so rpti o n . So m e

sub s ta nc e s th a t ca nno t be di ge s te d a re p a sse d ou t of th e b o dy a s
f ce ce s A
. ft e r a bs o r pti o n f oo
,
d is c ar r i e d t o s o m e p a r t o f t h e b o dy
whe r e it is n e e de d ; he re it is tra nsform e d int o living substa nce
by the p ro c e s s o f a ssi m i la ti on .

In or de r th a t me ta b olic a ctivity m ay go on witho ut ce a sing a ,

co n st ant s upply of e n e r gy is n e ce ss ary This e n e r gy is i n p a r t

.

fu rni she d by oxid a ti on i e th e che mical uni o n o f o xyg e n with th e

,
. .

li ving substa nce in a m ann e r which m ay be co mp are d to t h e

14 AN INTR ODU C TION T O Z OOLOG Y

cha nge s ta king pl a ce during co mbustio n Re spi r ati on supplie s .

t he o xyg e n f or su ch m e t a b o lic a ctiviti e s a nd a ls o e li m in a t e s ce r

,

ta in gase o u s e xcre t ory p roducts T he was te p ro ducts for me d .

by the b re a king do wn o f li ving sub s ta nce a re ca st off a s e xcr e ti ons .

The se sho uld no t b e co nfus e d with fa c e s which have n e ve r a ctu ,

a lly b e c o m e c o n s titu e nts o f living m a tt e r .

F igu r e 2 i s int e nd e d to indi ca t e th e r e l a tio ns o f the v a ri o us

m e tab olic a cti vitie s It Shows th a t an a nimal r e quir e s f oo d and
.

OX YG EN EXC RET IONB

FAEC EB

FI G . 2 . D i agram s h owi ng t he l ti o ns of the v ar i o us

re a m e t abo l ic ac tiviti e s
o f a ni m a l s .

o xy ge n ; tha t p ar t o f the f ood i s di ge ste d and a ssimil a te d whil e ,

th e r e st is c as t ou t as f a ce s ; a nd th a t t h e o xid a ti o n o f t he living
substance r e s ul ts in the p ro ductio n of c e r ta in e xcre tio ns whi ch
a r e e limin a t e d Ur e a is th e m o s t imp or ta nt of th e se e xcr e t or y
.

p ro ducts .

Gr o wth h a s b e e n s aid to re p r e se nt th e e xce ss o f a n ab o li sm

o v e r k a t a b olism T h e p o w e r o f gr o wth b e co m e s gra du ally l e s s
.

thr ougho ut th e life of a ny individu al a nd all a ni m als p ass ,

thro ugh c e r t ain w e ll -m arke d s tage s yo uth m a tu rity a nd old a ge

, , ,
.

You th i s g e n e r ally ch a ra ct e r iz e d by gr e a t vigo r a nd a cti vity a s

w e ll a s r apid g ro wth At ma tu ri ty a n ab o li sm a nd ka ta b oli sm
‘

are a b o ut e qu a lly b al a n ce d D u ring thi s a dult p e r i o d littl e

.

cha ng e i s ob se rv abl e in e xt e r n a l for m or gross w e ight M a tu rity .

g ra dua lly give s way to old age a pe riod of de clin e whe n the b o dy
, ,
 PHENOM ENA OF LI FE 1 5

l wly waste s a wa y and the vita l p ro c e sse s fin ally ce a s e altoge the r

S o .

The s e thr e e p e r i o ds yo uth m a tu rity a nd old age m a ke up th e

, , ,

lif e cycle o f e v e r y individu al .

( 6) R e pr od u cti on . P e r ha ps t h e m o s t r e m a r k abl e cha r a c

te ri s ti c o f a living o r g a ni sm i s its ability t o p ro duc e ne w indi
vi d uals lik e it s e lf a t s o m e ti m e in th e lif e cycl e This r e p ro ductive
.

p o we r usu ally m a ni fe sts its e lf during m a tu rity Am ong a nim al s .

a n o ff sp r ing m ay o r igin a t e fr o m its p a r e nt by a s e xu a l or s e xu a l

m e tho ds R e p r o ductio n m ay t ak e pl a ce by a Sp litting of th e

.

e nti r e b o dy int o t w o or m o r e p ar ts e a ch o f whi ch t a k e s up a n

,

inde p e nde nt e xist e nc e This m e thod o f multiplica ti o n is o fte n

.

call e d as e xu al r e pr oducti on; if th e ne w indivi du als a r e fo r me d by a

divisio n of th e or igin al or p are nt b ody int o a pp ro xi m a te ly e qu al
p ar ts re p roducti o n is s aid to b e by fi ssi on; if o n the o the r ha nd
, , ,

o nly a sm a ll p o r ti o n o f th e p a r e nt indi vidu a l b e c o m e s s e p a r a t e d

a s a distinct o r g a ni s m r e p ro ducti o n i s by bu ddi ng a nd th e o ff

, ,

sp ring are call e d b uds So m e tim e s an a nimal p a ss e s thr o ugh a

.

re sting st a g e du ring whi ch th e b o dy f ra gm e nt s int o a l a r g e num

b e r o f sm all p a r t s which subs e qu e ntly b e co m e f r e e indivi du al s .

Thi s a se xua l m e th o d o f r e p roductio n i s kno wn as sporu lati on .

S e xu al r e pr odu cti on inv olv e s th e uni o n of sub s t a nc e s fro m two

di fl t nim al Wh e n thi s p ro ce s s o ccu r s in ce t in ce ll e d
'

e r e n a s . r a o ne -

a nim als tw o individu a ls c o m e t og e th e r a nd e xch a ng e a p or ti o n

,

o f th e i r living sub s t a nc e ( Fi g in m any ce ll e d a nimal s the re

.
-

is a uni o n of se xu al ce lls e gg a nd spe r m whi ch o rigin a te fro m

, ,

t w o s e p ar a t e individu a l s Ase xu a l r e p r o ducti o n inc re a s e s th e

.

numb e r of indi vidual s witho ut alte ring the n a tu re of the i r

sub s t a nce ; s e xu a l r e p r o ductio n origin a t e s ne w individual s by
mi xing p o r tio ns o f two p r e e xisting o r ga ni sm s .

Ase xu al r e pr odu cti on m ay b e illu s t ra t e d by P ar a me ci u m ( p .

a singl e c e ll e d a nim a l w hich di vid e s twic e du r ing e v e ry tw e nty

-

fo u r ho u rs whe n co nditi o n s are fa vor abl e I ts m e ta b oli s m is s o

.

r a pid th a t it ca n gr o w to i t s m a ximum s iz e in h a lf th a t pe r i o d o f

time T h e ho n e y b e e ( C ha p XII ) fu r nishe s a n e xce ll e nt illus

. .

t r a ti on o f se xu al r e pr oducti on Thr e e sor ts of b e e s e xi s t : the

.
1 6 AN INTRODU CTION TO ZOOLOGY

wo rk e r s whi ch a re ste ril e fe m al e s and t ake no p ar t in re p ro

,

ducti o n ; qu e e n s o r se xu ally m a tu re fe male s ; a nd dro n e s o r

, ,

male s . N e w indi vidu a ls a r e fo r m e d by th e fu si o n of r e prod u c

tiv e or ge rm ce lls T h e dr o n e inj e cts th e sp e r m ce lls into th e
.

que e n whe r e the y are stor e d up to uni te with the e gg ce ll s a s

,

the y rip e n within h e r b o dy .

( 7) I rr i ta b ili ty o r r e a cti ve ne s s is a ch a r a ct e r istic p r op e r ty o f

o rg an i sms It giv e s the m th e p o w e r to r e sp o nd to ch a nge s in
.

the i r e nvi ro nm e nt in su ch a w ay tha t the y usu ally a tt ain f a vorabl e

conditi o n s f or e xi s t e nce Any ch ange in co nditi o n s t ha t pro
.

duc e s th e r e a cti o n s of a n ani m al i s t e r m e d a s ti m u lu s Stimuli .

m ay b e e xt e r n a l o r int e r n al ; th e f or m e r includ e s ch a nge s in

t e mp e r a ture or li ght in th e c o mp o siti o n o f the su rro un di ng Ine
,

di u m e t c hunge r a nd fa tigu e m ay s e rv e a s e x a mpl e s o f i nte r n a l

,
.

stimuli .

In m a ny inst ance s a nim al s show v e ry unifor m r e sp o ns e s t o

stimuli An e xce ll e nt illu s t ra tio n o f thi s f a ct i s fur ni she d by th e
.

k a tydid Th e re l a ti o n b e twe e n th e te mp e r a tu r e and th e numb e r

.

o f c a ll s pe r minut e i s s o c o n s t a nt in t hi s in se ct th a t t h e e x a ct

a tm o sph e ric t e mp e r a tu r e m ay b e c o mput e d f ro m th e foll o wing

fo rmul a .

no Of c a lls pe r minut e
Te mp e r a tu re
.

60
3
Th e ad a p pti ve owe r o f a nim als
fu r th e r illust
m ay b e
r a t e d by
th e kidn e y s o f m a n In m an as in all v e r te b ra t e s th e se ar e
.
, ,

p ai re d or g ans o ccupying a p ositio n n e a r th e d or s al side o f th e

b o dy w all Unde r n o r m al c o nditi o ns b o th e limi n a t e an approxi
.

m a te ly e qua l a m o unt Of e xc r e to r y m a tte r How e v e r no p e rm a .

,

n e nt inj u ry n e c e ssar ily r e sults if o ne is r e m ov e d f o r th e kidne y o n ,

t he u ni nj u r e d s id e o f th e b o dy e nl ar g e s t o such a n e xt e nt th a t
it is a bl e t o ca rry o n th e functio n s p re vi o u sly p e r fo rm e d by b o th .

In s t a n ce s o f thi s a d aptiv e p o w e r might e a sily b e multipli e d f o r ,

the y are co mm o n e v e r ywh e re In a b ro a d se n se e ve n th e i n

.
,

t e lli ge nce o f m a n a nd th e d e g r e e o f s ucc e s s h e m a y a tt a in d e p e nd

up o n h o w s ucce ssfully hi s r e a cti o n s m e e t sur ro unding co nditi o ns .
 P HEN OM EN A OF L F E I

3 . P LAN TS AN D AN I M ALS C OMP ARE D

is e a sy to ch oo s e ch ara ct e ristics th a t will s e rv e to di stingui sh

It
a t r e e f r o m a m a n but th e s e p a r a ti o n of th e Simpl e st a nim a l s
,

fro m the simpl e s t pl a nt s i s a m ore di ffi cult p r obl e m In f a ct .

,

the re a re a t th e p re se nt tim e a numb e r o f o r g ani s ms th a t a r e

cl a im e d by b o th b o t a nist s a nd z oo l o gists The r e i s no singl e .

p e culi arity whi ch can b e u se d in a ll ca se s to discr imin a te b e twe e n

th e se g r o ups of o r g a ni sms T h e vie w now g e n e r ally a cc e pte d
.

i s th a t pl a nt s a nd a nim al s o r igin a t e d t o g e th e r but h a v e d e v e l op e d

a l o ng di v e rg e nt lin e s Ho w e ve r ce r t a in g e n e r a l f e a tu r e s ca n
.
,

b e indi c a t e d in whi ch th e tw o kingdo m s di ff e r The se a r e giv e n .

in th e foll o wing t abl e ; but th e r e a de r sh o uld b e ar in mi nd th a t

th e re ar e e xc e pti o ns to e v e ry o ne o f the se crite ri a .

T AB LE II

C HI E F C HAR ACTE RI STI C S OF P L AN T S AND AN I M ALS C ON TR ASTE D

P L AN TS AN I M AL S

1 . St ru ct ur e Fo r m of bo dy ra th e r v a r i Fo r m of bo d y usua ll y i nva
ab el ; ne w o r ga ns a dd e d r i a b le
; o r ga ns co m pa c t
e x t e r na ll y . tl i t
a nd m o s y n e rna l .

2 . Lo co m o ti o n U su ally no ne i n a d u lt co n U su a ll y w e ll d e v e l ope d .

d iti o n .

I r r it ab ility R e sp o nd to ti m u li sl o wl y ;
s R e spo nd to ti m u li q u i ck
s

no ne r vo u s sys t e m . ly ne r v o u s sys t e m p r e s
e nt i n hi gh e r f o r m s .

4 M e tab o li sm P o sse ss c hl o rophyll ; m a nu N o chl o ro p hyll re q u i r e

f a ct u r e o r ga ni c f oo d f r o m o rga ni c f ood .

C 02 a nd H 20 i n t h e p r e s
e nce of light .

Wa s te p ro d u cts Oxyge n , ca r b o n d i o xi d e , C a rbo n d i o xi d e , wa te r ,

u re a ,

wa te r . f a ce s .

p r e se nce o f chlorophyll in g r e e n pl a nts h as a n imp o r ta nt

Th e
b e a r ing o n m e t ab o lis m f o r it e n abl e s pl a nts to m a nufactur e
,

sta r ch whe n light i s a v a il abl e Thi s simpl e o r g a nic c o mp ound i s

.

c
1 8 AN I NT R OD U C TION T O ZOOLOGY

bui lt fro m th e ino rg ani c s ub s t a nc e s c arb o n di oxide a nd w a te r

, .

With st a r ch a s a b a sis m o r e co mpl e x o r g a nic co mp o unds a r e

e l a b ora t e d by th e living s ub s t a nc e s N o w th e living s ub s t a nc e
.

o f pl a nt s i s p r a ctic a lly th e s a m e as th a t o f a nim a l s a nd it s e e m s ,

s t r a ng e a t fi rs t th o ught th a t pl a nt s s h o uld giv e o ff o xyg e n a s a

w as t e p ro duct f or thi s ga s i s one o f th e co n s ta nt n e e d s o f li ving

,

sub s t a nce If h o w e v e r w e k e e p in mind th e fa ct th a t chl oro

.
, ,

phyll i s c arrying on o ne se t o f p r o ce sse s ( Fi g 3 ) which r e qui r e .

C AR BON DIOXIDE OX YG EN

“
AT R E S T AR CH

FI G 3 . . D i agr am S h o wi ng t h e ac tivi ti e s o f chl oro phyll .

c arb o n di o xide and lib e r a te o xyg e n as a wa st e p r o duct whil e ,

th e living s ub s t a n ce i s c a rr ying o n a n o th e r se t whi ch u se o xyg e n

a nd r e l e ase c a r b o n di o xid e ( Fi g th e m a tt e r b e co m e s o ne o f
.

S impl e a dditi o n a nd s ubt racti o n ; o ne g r o up o f p r o c e sse s u se s th e

w as t e p ro du cts o f th e o th e r .

T h e qu aliti e s th a t a r e u s u a lly cit e d a s b e ing p e cu li a rly ch a ra c

t e ri s ti c o f a nim al s a r e locom oti on a nd ne rvou s acti vi ty With th e .

e xc e pti o n o f a f e w e xt r e m e ly se n s itiv e sp e ci e s o f whi ch th e co m

m o n se n sitiv e pl a nt M i m osa pu di ca i s th e m os t f a mili a r e x

, ,

a mpl e ,
pl a nts r e sp o nd v e ry sl owly to e xt e rn a l s timu li and th e i r
p o w e r o f tr a n s mitting impul se s i s p oo rly de ve l op e d Lo co mo tio n .

i s ab se nt e x ce pt in a f e w simpl e for m s a nd f r e e swimming r e p ro

d ucti v e ce lls .
20 AN INTRODU CTION TO Z OOLOGY

str ucture of p r o t opl a sm . Of the s e th e th r e e foll owing m ay b e

m e ntio n e d .

(1 ) T h e R e ti cu la r The ory m aint ains th a t p ro t opl a sm c o nsists

o f a living n e tw o r k o f a n a s t o m o s ing fib e r s ( Fi g 4 B ) ; b e tw e e n .
,

th e se a v a ri e ty of no n living sub st a nce s s uch a s w a t e r a nd f a t

-
, ,

m a y b e p r e s e nt .

( 2) T h e Alve ola r The ory ( B u tsch li 1 8 9 2) supp o s e s th a t pr o ,

t Opla s m h a s a f o a m lik e st r u ctu r e so m e wh a t Simil ar to a m ass o f

-

m inut e d ropl e t s such a s e xi s t in a n e mul si o n o f o i l a nd w a t e r

( Fi g 4 A) T h e o r igin a t or .
,
.

o f thi s th e or y w a s a bl e to

m a k e a r tifici a l e mul s i o ns

whi ch sh owe d a s t riking

r e se mbl a nc e t o living sub

s t a n ce He m a in ta in e d th a t .

th e fib ro us n e two r k d e ,

s cr ib e d by th e a dhe r e nt s o f
B
th e re tic u l a r th e o r y r e pr e
F I G 4 Di g m ill t ti ng (A) th
. . a ra s lv
us ra e a e ’

l
o ar ,n d ( ) th
a B ti l th y f e s e
re nt e d
cu a rt h e w a ll
e or s Of th e 3 1" 6
o
‘

p t
ro o
p l m i t t
as ( F m D hl o la r b o di e s whi ch h a d b e e n
c s r u c u re ; ro a

gr n nd K p n )
e a e er .

cu t across .

( 3 ) T h e Gr a nula r The or y (Altm a n 1 8 9 2) a sse r t s th a t p r o t o ,

pl a sm i s co mp o se d o f innum e r abl e living g r a nul e s a rr a nge d e ithe r

a l o ng fib e r s o r a m o ng a lv e o li ; n o thi ng i s e sse nti a l h ow e v e r e x , ,

c e pt th e g ra nul e s .

N o n e o f th e se th e o ri e s h a s univ e r sal app ro v al but th e fi rs t tw o ,

h av e b e e n m o r e wide ly a cce pt e d th a n th e l ast M a ny r e ce nt .

a uth o r iti e s b e li e v e th a t p ro t opl a s m i s highly v a r i a bl e in s t r u ctu r e .

A fou r th vi e w h e ld by m a ny Ob s e r v e r s is th a t th e v a r i o u s typ e s
d e s crib e d a b ove a r e conn e ct e d by int e r m e dia t e g r a d a ti on s a nd
m ay b e t ra n s fo r m e d o ne int o a n o th e r ”
( 9
1 p ,
.

P r o t opl a sm i s a mixtu r e o f e xt r e m e ly co mpl e x ch e mi ca l co m

p o unds ; the se ar e a t l e a s t in p a r t e xt r e m e ly l abil e a nd the r e fo r e
, ,

a r e a pt t o v a r y a t di fl e r e nt tim e s It i s imp o ssibl e t o s tudy.

livi ng substa nc e Sin ce e v e n the m os t ca r e ful m e th o d s o f a n aly si s

,
 P HE N OM EN A OF L F E I 21

imm e di a te ly kill p r o t opl a s m He nc e id e a s up o n t he che mi s t ry

.

Of th e living o bj e ct ca n b e Obt a in e d o nly by d e ducti o n s f r o m

”
che m ic a l di s co ve ri e s in t h e d e a d o bj e ct ( 1 7 p N e ve r
,
.

t h e le ss w e h a v e g a in e d a co n sid e r a bl e a m o unt o f kn o wl e dg e a s t o
,

t h e che mi ca l n a tu r e o f p r o t o pl a s m N in e ty se ve n pe r c e nt o f
.
-

t h e o rg a ni c a nd in o r g a nic co n s titu e nt s o f p r o t o pl as m a re m a d e
up of fo ur e l e m e nt s o ccu r r ing in th e foll o wing p ro p or tio ns
,

Oxyg e n 6 5 pe r ce nt
C a rb o n p e r c e nt
Hyd r o g e n 1 1 p e r c e nt
N it r og e n p e r c e nt
Th e se a nd o the r l e ss imp o r t a nt e l e m e nt s for m c e r t a in ra th e r
de finite g ro up s o f c o mp o und s w h i ch w e will no w consid e r in so m e
,

d e t ail .

Wa te r i s th e m o s t imp o r ta nt o f th e in o rg a nic co n s titu e nts ;

it co mp r i se s m o r e th a n 5 0 pe r c e nt o f th e we ight o f m o s t
a nim a l s a nd in so m e m a r in e f o r m s r e a ch e s a s high a s 9 9 pe r
,

ce nt . It se e m s r e m a r k a bl e th a t m a n i s a bl e to e xi s t whe n 59
p e r ce nt o f hi s b o dy i s m a d e up o f w a t e r It n o t o nly
. o c c u r s in
c o mbin a ti o n s but a l so a cts a s a so lve nt f o r v a r i o us s ub s t a nc e s
,

th a t a r e fo und in p r o topl a s m .

As ide f r o m w a t e r th e m os t imp o r t a nt in o rg a nic sub s t a nc e s a re

v ari o u s s a lts a nd di sso lve d ga s e s Of th e salt s t h e chl o ride s
.
,

ca rb o n a t e s a nd ph o sph a t e s Of t h e c o mm o n e r a lk a li a nd a lk a li
,

e a r th m e t a l s d m in t T h p incip l g c b n di
p r e o a e .e r a a se s a re a r o

o xid e a nd o xyg e n .

T h e o r g a ni c co mp o und s f o und in p r o t opl a s m a r e divid e d int o

th re e ge n e ra l cl asse s : p r o te ids ca rb ohyd ra t e s a nd fa ts
, ,
Of .

the se o nly th e p r o t e id s a nd the i r d e riva tiv e s h a ve b e e n de m o n

s t r a t e d with ce r t a inty a s co mm o n t o a ll c e ll s ; h e nc e th e y mu s t ,

b e se t a p a r t a m o ng th e o r g a ni c c o n s titu e nt s o f living m a tt e r a s
th e e s s e nti a l o r g e n e r a l s ub s t a n ce s in c o nt ras t t o a ll sp e ci a l
,

s ub s t a n ce s (1 7 p ,
.

P rote id s a r e a lw a y s co mp o s e d o f th e fi v e e l e m e nt s carb o n , ,
22 AN INTRODU CTION TO ZOOLOGY

o xyge n hyd rog e n a nd ni trog e n and a small a m ou nt of sulphu r

, , .

In thi s r e sp e ct the y di ffe r f r o m th e o the r two cl a ss e s o f o rg a ni c

co mp o unds whi ch l a ck the nit r og e n a nd s ulphur T h e p ro t e id .

m ol e cul e s a re e xt r e me ly l a rg e o ne Oft e n co nt aining m o re tha n

,

a th o u sa nd a t o ms The y a r e no t re a dily di ffusibl e th ro ugh

.

a nim a l m e mb r a n e s t hi s b e ing d u e in p a r t t o t h e siz e o f t h e m o l e

,

cul e s a nd a l so to th e fa ct tha t such c o mpl e x c o mp o unds d o no t

di sso lve c o mpl e t e ly unl e ss the y b r e a k do wn into simpl e s u b
st a nce s In ste a d o f di ssolving p ro te ids a b so r b e no r mo u s
.
,

qu a ntiti e s o f wa te r s we lling up like a spo nge ; the y a r e c all e d

,

colloid s t o di s tingui s h th e m f r o m cr ys ta lloid s lik e s ug a r w h i ch , ,

a r e e a sily so lubl e C oa gu la ti on o r cl o tting is a n o th e r p e cu li a r ity

.
, ,

Of p r o t e id s .

C a r bohydr a te s a re c o mp o unds o f c a r b o n hyd ro g e n a nd o xyg e n , , ,

the l as t t wo a lw a y s o c cu r ring in t h e s a m e p rop or ti o n s in whi ch

the y a re fo und in wa te r ( H O) T h e m os t fa mili a r e x a mpl e s g .

o f thi s cl as s Of s ub s t a nc e s a r e t h e s t a r ch e s a nd s ug a r s whi ch a r e ,

ch a ra cte ri s tic o f pl a nt s ra the r tha n a nim a l s th o ugh fo und in b o th ,

.

C a r b o hyd ra t e s a r e co mp a r a tive ly simpl e co mp o unds wh e n

co nt ra s te d with th e p ro t e ids a nd a r e r e a di ly o xidi zabl e thus
, ,

p ro ducing e ne r gy So me va r ie tie s of living s ub sta nce app are ntly

.

c o nta in no c arb ohyd ra t e s .

F a ts lik e wi s e a r e no t inva ri a bl e c o n s titu e nt s o f p ro t opl as m

, , .

Tho ugh o f wide sp r e a d o ccurre n ce fa tty co mp o unds a r e p a r tie n ,

larly ch a r a ct e r i s ti c Of a nim a l s The y m ay b e said to co n si s t o f

.

a n a lc o h o l (glyc e r in) whi ch h a s l os t so m e w a t e r a nd c o mbin e d

with a fa tty a cid The y a r e a ll light e r th an wa t e r a nd d o no t

.
,

unit e with it .

P r o t opl a sm c o n si s ts th e n o f c o mpl e x a nd v a ri abl e mixtu re s o f

, ,

p ro te ids c arb o hydr a te s a nd fa t s toge th e r with w a te r s alt s a nd

, , , , ,

di s solve d g as e s N o unifo r m che mica l formul a can b e give n f or

.

it ; in f a ct o ne Of i ts chi e f chara cte ri s tics lie s in th e unce asing

,

ch ange s th a t it unde r goe s It h as th e p1 Ope r ti e s of m e t ab olic

.
°

a ctivity i rr it a bility
,
c o nt ra ctility r e p r o ducti on a nd all o th e r s
, , ,

p e culi ar to living or g an i sm s (p It s e e m s s tr a ng e tha t s uch

.
 P HEN OM EN A OF L F EI 23

a ch a ng e abl e sub s t a nc e sho uld p o s se s s g re a t sp e cificity ye t such ,

i s th e ca se T h e p r o t opl a s m o f e a ch sp e ci e s o f o r g a nism is p e r
.

f e ctly di s tin c t f r o m th a t o f e v e r y o th e r sp e ci e s A t r e e of a
.

c e r t ain v a r i e ty a lw a ys giv e s r is e to o the r t re e s simil a r to it ;

f o r e x a mpl e a p e ar s e e d n e ve r d e v e l op s into a p e a ch t r e e
, ,

but a lw a ys int o a p e ar t r e e o f th e s a m e kind tha t p ro duc e d it ;

like wi se th e e gg o f a pure b r e d P lym o uth R o ck h e n n e ve r giv e s
-

r is e t o a nything but a chick e n o f t h e P lym o uth R o ck v a r i e ty .

N e ve r th e l e s s b o th th e tr e e a nd t h e h e n t a k e th e i r o r igin f r o m
,

a n a pp are ntly s impl e mic r o s c o pic bit o f p r o t opl a s m th e f e r tiliz e d

,

g e r m c e ll .

5.A P HY S I C O C
- H E MI A
C L E X P L AN ATI ON OF THE P H E N OME N A
OF LI F E

The e ffo r ts of l h e mi s t o f Old w h o w ro ught in th e se clu si o n

th e a c

o f hi s l a b o r a t o ry w e r e di re ct e d t o w a r d t h e so luti o n o f t w o p r o b

l e m s : th e p ro du cti o n o f a n e lixi r o f life a nd th e m a nufa cture o f

,

gold Hi s tim e h a s p a sse d a nd hi s p r o bl e m s w e r e n e v e r s olve d

.
,
.

Hi s succ e sso r o f m o d e r n tim e s th e bi o ch e mi s t h a s lik e wi se

, ,

ch o se n p r o bl e m s Of fund a m e nt a l imp o r t a nc e a nd g r e a t di ffi culty ;

h e h a s a tt e mpt e d t o giv e a n e xpl a n a ti o n o f a ll vit a l ph e n o m e n a

by m e a ns o f physi ca l a nd ch e mic al l a w s T h e sci e nti fic w o rld
.

now c o nt a ins m a ny s ch ol a r s w h o m a int a in th a t living o r g a ni s m s

a r e r e ally m achi ne s a nd a r e o pp os e d t o th e id e a o f vi ta li s m whi ch
, ,

p re supp o se s th e p r e se nce o f so m e vit a l p r in cipl e One o f th e .

l e a di ng inve stiga to r s in thi s fi e ld cl a im s th a t livi ng o r ga ni sm s a r e

to b e co nsid e r e d a s
“ ch e mi ca l m a chin e s co n si s ting e s se nti a lly
,

o f c oll o id a l m a t e r i a l whi ch p osse ss th e p e culi a r itie s o f a ut o

,
”
m a ti ca lly d e ve l oping p r e se r ving a nd r e p r o ducing th e m se lve s
, ,

( 5 p An o th e r p r o min e nt z ool o gi s t m a k e s th e fo ll o wing

“
1 , .

sta te m e nt : I s a y a g a in with a ll p o s sibl e e mph asi s th a t th e

, ,

m e ch ani s tic hyp o th e sis o r m a chin e th e o ry o f living b e ings i s no t

fully e s t abli sh e d th a t it m ay no t b e a de qu a t e o r e ve n t r u e ; ye t I
,

ca n o nly b e li e v e th a t until e v e r y o th e r p oss ibility h a s r e a lly b e e n

24 AN INT R ODU CTION TO Z OOLOGY

e xh a uste d s ci e nti fic biol ogi s ts sho uld h old f ast t o th e wo r king

,

p rog r a m tha t h as cr e a t e d th e scie nc e s o f bi o l ogy T h e vita li st i c .

hyp o the si s m ay b e he ld and is h e ld a s a m a tt e r o f faith ; but w e

, ,
”
c a nno t c all it sci e n ce wi tho ut mi s use o f th e wo r d ( 20 p ,
.

A g re a t m a ny s ci e nti s t s h a v e re c e ntly c o m e f or w a r d t o c o mb a t
t h e o ld id e a o f vit a li s m a nd th o ugh s o m e o f th e m h av e m a d e
, ,

e rrors th e i r influ e nc e as a wh ol e h as b e e n highl y stimul a ting t o

,

z ool o gic al tho ught .

Until co mp ara tiv e ly r e ce nt time s it w a s b e lie ve d th a t o rg a nic

co mp o unds cou ld b e built up o nly by living s ub s t a n ce E ve n .

su ch a n abl e s ci e nti s t a s Li e big s upp ose d th a t hi s b e e f e xt ra ct

c o nt ain e d s o m e vit ali s tic p rin cipl e th a t w a s imp a r te d t o tho s e
wh o a t e it Hi s id e a s w e re o v e r th r o wn by a s impl e e xp e r im e nt
. .

Si x puppi e s f r o m th e sa m e litt e r w e r e di vid e d int o tw o l o t s ;

thr e e we re f e d b e e f e xt r act a nd th e o the r s w e re all ow e d to r e m a in
witho ut a ny foo d wh a t e v e r Tho se f e d with th e b e e f e xt ra ct
.

die d fir s t thu s de m o n s tr a ting th a t th e e xtr a ct w a s m ore a sti m u

,

l a nt th an a foo d T h e b e li e f th a t o rg a nic c o mp o unds co uld

.

on l y c o m e f r o m living o r g a ni s ms w as p r o v e n e r ro n e o u s as e a rly
a s 1 8 2 8 wh e n Wo e hle r m a d e u r e a by a s ynth e ti c p r o c e ss in h i s
,

l a b o r a to r y It wa s to co mb a t such vi tali sti c id e a s th a t th e

.

m e cha ni s ti c the or y fi rs t a r ose a nd i ts a dh e r e nt s h a v e m e t

,

with a co n side ra bl e d e g r e e o f s ucce s s E xp e rime nts fro m m a ny

.

s o u r c e s mi ght b e cit e d whi ch supp o r t th e m e ch a ni s tic vi e w but ,

o nly a f e w se l e ct e d in s t a nc e s ca n b e giv e n .

M a ny o r g a n s will ca rr y o n the i r n o r m a l a ctiviti e s wh e n r e m o v e d

f r o m a n a nim a l s b o dy A tu r tl e s h e a r t will re m ain a live and
’
.
’

b e a t f or a co upl e o f d ays if it i s pl a ce d in a dish co nta ining a

p rop e r mixtu re o f min e r al salts in w a te r F u r th e r m o r e th e ra t e
.
,

o f th e b e a ts m a y b e in cr e a s e d o r d e c r e a s e d a t th e will o f t h e e xp e r i

m e nte r by v a r ying th e p r op o r tio n a te a m o unts o f th e s alt s in

s oluti o n .

Org a n s f r o m o ne a nim al m ay b e g r a ft e d up o n a n o th e r Th e r e
.

i s a gl a nd th e thyr oid in th e n e ck o f hum a n b e ing s which wh e n

, , , ,

di se ase d c a use s a m al a dy kn o wn a s g o it e r T h e r e m ova l o f th e

,
.
 P HE N OM E N A OF L F E I 25

thy roid i s foll o we d by d e a th It w a s di s c o ve r e d so m e time a go

.

th a t thi s gl a nd might b e r e m o ve d with impu ni ty if a pie c e o f a

h e althy gl a nd f r o m a n o th e r a nim a l s u ch a s a sh e e p w e re g r a fte d
, ,

und e r th e s kin in a ny p a r t o f th e b o dy T h e fore ign pie ce o f

.

ti ssu e app rop r i a te s a bl o o d s upply a nd b e c o m e s functi o n a l in its

ne w p o s iti o n . Thi s i s o ne o f th e m a ny c a se s p r oving tha t a
d e finite p o siti o n a l r e l a ti o n o f o r g a n s is no t a lwa ys n e c e ss a r y if
th e i r se cr e ti o n s a r e p r e s e nt
.

Th e Opp o n e nt s o f vit a li s m h a v e a l s o b ro ught f or w a r d m a n y

a r gum e nt s a g a in s t th a t th e o r y f r o m a s tudy o f e nzym e s o r f e r

me nts a cl ass o f s ub s t a n ce s w hi ch c a u se ch e mi ca l re acti o n s with

,

o u t und e r g o ing a ny ch a ng e th e m s e lv e s An e x a mpl e i s pty a lin

.
,

whi ch i s p r e se nt in hum a n s a liva a nd h a s f o r i ts functio n th e

ch a ng e o f s t a rch t o s ug a r . A l a rg e numb e r o f s ub s t a nc e s o f thi s
sor t are kn o wn to b e p r e se nt in living o r g a ni s m s . T h e e xpl a n a
ti o n o f the i r a cti o ns m ak e s cl e a r m a ny p r o ce sse s which w e r e
fo r m e rly s upp o se d to b e d ue to ce r ta in vitali s tic p rin cipl e s R e .

se a r ch in thi s fi e ld i s o nly in i t s inf a ncy .

The se in s t a nc e s a r e s uffi ci e nt t o sh o w th a t th e m e ch a ni s tic

po int o f vi e w is a mu ch m o r e p r o g r e ssiv e o ne th a n th e vit alis
ti c
. Thr o ugh i ts influ e n ce sci e nti s ts h av e b e e n a bl e to p ro ve
th a t a l a rg e numb e r Of th e a ctiviti e s p re se nt in living m a tte r a re
s ubj e ct t o d e fini t e l a ws m a ny o f whi ch h a v e b e e n kn o wn f o r a
,

l o ng ti m e t o phy si ci s ts a nd ch e mi s ts Ho we ve r thi s m e tho d h a s

.
,

no t e xpl a in e d a ll vit a l ph e n o m e n a a nd p e r h a p s n e v e r will

, .
 C HAP T ER III

TH E C ELL AN D TH E C ELL TH E ORY

1 . M ORP H OLOG Y AN D P H Y SI OL OG Y OF C E LL S

T HE p ro topl as m co nt ain e d in th e b o dy of a n a ni m al i s no t o ne
co ntinuo us m ass but i s s e p a r a t e d by m e mb r a n e s a nd o the r
,

str uctu r e s into a g r e a t numb e r o f minute b o d i e s c all e d ce lls .

Th e se ce lls tho ugh v a r ying g r e a tly in size a nd sh ap e all h a v e

, ,

th e s a m e fund a m e nt a l pl a n o f s t r u ctu re T h e simpl e s t a nim a l s

.

co n si s t o f a singl e c e ll but the b o di e s Of th e m ore co mpl e x fo r ms

,

a r e m a d e up o f milli o n s o f th e s e tiny bit s o f p r o t opl a s m On .

a cc o unt o f i t s univ e r s a l p r e se n ce in a ni m a l s a nd pl a nt s th e ce ll ,

h a s b e e n c a ll e d th e u ni t of lif e It i s n e ce ssar ily o f v a s t i m

.

p o r ta nce in th e s oluti o n o f bi ol o gi ca l p ro bl e m s a nd h a s b e e n ,

much s tudi e d by inv e s tig a t or s du ri ng th e p a s t fo r ty y e a r s .

In o r de r t o h a ve th e s tr uctu r e o f a typi cal ce ll cl e a rly in mind

w e will now tu r n ou r a tt e nti o n t o th e f o ll o wing d e scr ipti o n o f its
p a r ts R e fe r e nc e to F igur e 5 w ill m a ke th e wr itt e n de s cripti o n
.

cl e a r e r Sinc e th e c e ll c o nt ain s p r o t opl a sm whi ch a cts lik e a

.
,

fluid it n a tura lly t e nds to h av e a sph e ric al fo r m Thi s co nditi o n

,
.

i s s e ld o m r e ali ze d in n a tu r e f or v a r i o u s f a ct o r s such as un e qu a l
, ,

g rowth and p re ss u r e fr o m o the r c e ll s m o dify th e sh ap e T he ,

.

fa ct th a t th e c e ll sho wn in th e fi gu re h a s a n Obl ong o utlin e is ,

the r e fo r e o f no Signifi can ce

,
.

T h e c o nt e nt s o f m a ny ce ll s includ e s no t o nly th e a ctiv e p r o t o

pl asm but a l so v ari o u s kind s o f p assiv e b o di e s s uch a s f a t gl o b
,

ul e s The se a re kn own a s m e taplasm T h e pr otoplasm of a ny

. .

c e ll i s m a de up o f tw o p rincip al kind s o f s ub s t a n ce th e cytoplasm ,

a nd th e k a r yoplas m T h e nu cle u s c o nt a i ns a ll o f th e l a tt e r
. It .

is th e c e ntral b o dy a nd i s f or v a r i o u s r e a so n s r e g ar de d a s th e
, , ,

26
28 AN IN T R ODU CTION TO Z OOLOGY

a rr a ng e d in co nc e ntr ic z o n e s e a ch co nt a i ni ng a di ff e r e nt kind o f
,

p ro t opl a sm ; o r it m ay b e m o difi e d to fo r m h a i r lik e p r oj e cti o n s -

c all e d ci li a a nd o the r c e llul a r o rg a n s T h e st r u ctu r e of cyto

,
.

pl a s m i s h o w e v e r r a th e r u ni for m in th e sam e ce ll but hi ghly

, , ,

v a ri a bl e in di ff e r e nt ce ll s a nd so m e tim e s e v e n in diffe r e nt s t a g e s
,

o f a n i ndividu a l c e ll A ca r e ful e x a min a ti o n o f ce r t a in ce ll s

.

a pp e a r s t o l e a v e no d o ubt but th a t a r e ticul a r n e tw o r k i s p re s e nt

in the cytopl asm Oth e r ce ll s sho w no sign o f s u ch a n a rra ng e

.

m e nt o f m a t e r i a l s T h e a lve ol a r co nditi o n e xhibit e d by m a ny

.

c e ll s i s p e rh ap s th e m o s t typica l .

Alth o ugh a d e fi nit e ce ll wa ll i s m o r e o ft e n f o und in pl a nt s th a n

in a nim al s th e cyto pl a s m o f th e l a tte r m ay s e cre te a de lica te
,

m e mb r a n e o r e v e n a w a ll o f co n side ra bl e t hi ckn e ss a s in th e ca se
, ,

Of c a r til a g e ( Fi g 4 8 C ) .
,
.

E mb e dd e d in th e cyt o p la sm u s u a lly n e a r th e nu cl e u s i s a , ,

mi nute but imp or t a nt b o dy th e ce ntr os om e o r a p a i r cl o se to , ,

g e the r T he ce nt r o so m e s a r e o ft e n s u rr o unde d by a cl e a r sp a ce
.
,

t h e ce ntr osphe r e Am o ng t h e o th e r b o di e s s u sp e nd e d in th e cyt o

.

pl asm m ay b e m e nti o n e d Oil a nd wa te r vacu ole s crysta ls a nd ce r , ,

t a in o rg a n s kn o wn a s pla s ti ds P l a stid s m ay b e c ol o r e d ( ch ro m o
.

p l a s tid s) or whit e ( l e uc o pl a s tid s) .

The r e i s a de finit e di vi si o n o f l ab or a m o ng th e p a r t s o f a c e ll .

T h e p a r ti cul a r functi o n o f th e nu cle u s a sid e fro m i ts imp o r t a nt ,

r e l a ti o n t o c e ll divi s i o n t o b e d e scr ib e d l a t e r s e e m s t o b e t h e
, ,

co nt r o l o f th e a ctiviti e s by w hi ch t h e p r o t o pl a s m is e l a b o r a t e d .

T h e cytopla s m f ro m its di r e ct r e l a ti o n to th e o ut s id e w o rld i s

, ,

th e s e a t o f s u ch fun cti o ns a s i r r it a bility a b s orpti o n di g e s ti o n , , ,

e x cr e ti o n a nd re spi r a ti o n
,
Th e ce ntr os ome i s o f imp o r t a n ce
.

du ring c e ll divi sio n T h e ce ll cove r i ng m ay b e se cre t e d f or pro

.

t e cti on o r supp o r t o r m ay b e e xt r e m e ly d e lic a t e a nd h a v e si g

,

ni fi ca nce o nly as it h e lp s t o c o nt r ol th e a b sorpti o n o f ce r t a in

fluid s P las tid s m ay r e p r e se nt s to r e d foo d o r w a st e p r o du ct s ;
.

s o m e o f th e m h o w e v e r h a v e o th e r fun cti o n s e g th e chloropla s ts

, , ,
. .
, ,

whi ch c a rr y on photosynthe si s in m a ny pl a nt a nd a f e w a ni m a l
c e lls .
 T HE C ELL AN D T HE C ELL T HEOR Y

As h as l
dy b e e n n o t e d th e s i ze a nd for m o f c e ll s a r e v ari abl e
a re a ,
.

B act e r i a a r e a m o ng th e s m a ll e s t kn o wn so m e Of th e m m e a s u r i ng
,

no t m o r e th a n o ne hund r e d th o u sa ndth o f a m e t e r i n l e ngth .

On th e o th e r h a nd th e e ggs o f a l ar g e bi r d a r e s ingl e c e ll s ; th e y
,

a r e e xc e e d e d in l e ngth by ce r t a in n e r v e c e ll s wh o se fib e r s e xt e nd

o u t a lm os t a m e t e r T h e limit in s iz e i s p ro b ably d e t e r min e d

.

by th e nucl e u s sinc e it r e gul a te s th e m e t ab olic a ctiviti e s o f th e

,

c e ll
.

So m e id e a o f th e wid e v a r i a ti o n in t h e f or m o f c e ll s m ay a l so
b e g a in e d fr o m Fi gu r e 48 w hi ch Sh o w s sp e cim e n s o f v a r i o u s typ e s
,
.

All o f th e lif e p r o c e sse s o f th e Simpl e s t a nim a l s th e P r o t o z o a , ,

a r e ca r r ie d o n by a s i ngl e c e ll T h e ce ll s o f a multic e llul a r a ni

.

m al a M e t a z o o n a r e mutu a lly d e p e nd e nt a nd ca nn o t e xi s t if
, , ,

i so l a te d In m a ny c a se s th e r e i s a ctu a l c o ntinuity b e tw e e n th e
.

p ro topl a sm o f di ff e r e nt c e ll s a nd so m e z oo l ogi s t s h av e g o n e SO
,

f a r a s t o m a int a in th a t th e b o dy Of a c o mpl e x a nim a l Sh o uld no t

b e c o n sid e re d a n a s se mbl a g e Of s e p ara t e ce ll s but a co ntinu o u s ,

m a s s o f p r o t opl a s m with nucl e i s c a tt e r e d thro ugh it Oft e n .

this co nn e ctio n b e twe e n ce ll s i s b ro ught a b o ut by sl e nde r p r o t o

pl asmi c b r idge s ( Fi g 4 6 C ) ; in o th e r in s t a nc e s ce ll s a r e so indi s
.
,

t i ngu i sh a b ly bl e nd e d th a t no b o und a r i e s ca n b e di s ce r n e d A
“
.

ce ll i s th e r e fo r e no t a lw a y s a t r u e unit but a n e ce ssa r y p a r t

,

in a co mpl e x who l e T h e numb e r o f ce ll s a sso ci a t e d t o fo r m

.

a singl e o r g a n i s o ft e n e n o r m o u s ; th e numb e r o f n e rv e c e ll s in
t h e g r a y m a tt e r o f th e hum a n b r a in h a s b e e n e s tim a t e d a t
a ll o f th e se t o g e th e r w o uld o ccupy o nly a t r ifl e

m o re tha n a cubic in ch o f sp a c e .

2 . C E LL D I VI S I ON

C e ll smultiply e ithe r by di r e ct o r in di re ct divisio n I ndi r e ct .

ce ll di vi s i on a lw a y s inv o lv e s a d e finit e se ri e s o f s t a g e s whi ch

fo ll o w e a ch o th e r in a r e gul a r se qu e n ce th e wh ol e p ro ce ss b e ing
,

kn o wn a s m i tosi s o r mito ti c ce ll divi si o n Di r e ct di vi s i on o r

, .

a mi tosi s i s a m o r e s impl e p ro ce ss h a ving no co mpl e x s e r i e s o f

,

s t a ge s .
3 0 AN IN T R ODU CT I ON TO Z OOLOGY

( )
I M i tos i s — . ll th a t i s no t unde r going divi si o n i s s aid
A ce

to be in a r e s ting c o n di ti o n ( F i g 6 A) Whe n it di vide s by the

.
,
.

indi re ct m e tho d th e se ,

r i e s o f ch a ng e s m a y co n

v e ni e ntly b e a rr a ng e d in

f ou r s tage s kno wn a s th e ,

p p
ro h a s e ,
m e ta phas e ,
a na

hase , a nd te lophase T h
p e .

m odifi ca ti o n s o c cu rr ing
during mit osi s a r e p rim a
r ily c o n ce r n e d with t h e

nu cl e u s a nd c e nt r oso m e s ,

th e cyt opl a s m r e m a ini ng

co mp a r a tiv e ly p a ss iv e .

T h e pr ophase i s ch a r
a cte ri z e d by p r e p a r a t o r y

cha nge s Th e chro m a tin .

g ra nul e s th a t a r e s ca t
t e r e d th r o ugh th e nucl e u s
in th e r e s ting ce ll b e com e
a r r a ng e d in t h e f o r m Of

a l o ng th r e a d o r s pi r e m e

( Fi g 6 B ) At the sa m e .
,
.

tim e the c e nt r o so m e s
m o v e a p a r t fin a lly r e a ch ,

ing o pp o site sid e s o f th e

nu cl e u s (B C D E) , , ,
.

T h e ra di a ting lin e s th a t
a pp e a r a b o ut th e m ( B )

l a t e r give rise to a s pi ndle

D
( ) W hil e thi s i s g o i.ng
F G 6
I .D i g m ill t ti ng m it ti
. a ra s ll o n th e nu cl e a r m e mb ra ne
u s ra o c ce

d ivi i n ( F m Wil n )
s o . ro
g e n e ra lly disint e g ra t e s
so .

a nd th e s pi r e m e s e gm e nt s int o a numb e r Of b o di e s ca ll e d chr a

m os om e s ( D) ; the se t a k e a p o siti o n a t t h e e qu a t o r o f th e spindl e ,

 T HE C ELL AN D T H E C ELL T HEOR Y

h alfw a y b e tw e e n t h e ce nt ro som e s ( E) T h e s t a ge s hown in

.

Figu r e 6 F i s kn o wn a s th e a m phi a s te r ; a t thi s tim e a ll o f

, ,

the m a chi n e r y co nc e r n e d in mit os i s i s p r e s e nt The r e a re tw

. o

as te r s e a ch c o n s i s ting Of a c e nt r o s o m e su r r o und e d by a numb e r

,

Of ra di a ting a s tr a l r a ys a nd a Spindl e whi ch li e s b e tw e e n th e m

,
.

T h e ch r o m o s o m e s li e in t h e e qu a tor i a l pla te (e p) .

D u r ing t h e s e co nd s t a g e t h e m e ta phase t h e ch ro m o s o m e s split

, ,

in such a w a y th a t e a ch o f the i r p a r t s co nt a in s a n e qu a l a m o unt

o f ch r o m a tin ( G) AS w e sh all se e l a t e r thi s i s o ne Of th e m o s t
.
,

S ignific a nt e v e nt s th a t t a k e s pl a ce dur ing mit o si s Oft e n the .

ch ro m o s o m e s s plit b e f o r e th e y h a v e a s s um e d a n e qu a t or i a l p o si

ti o n ( Fi g 6 E ) ; o the r min o r v a r i a ti o n s a l so o ccu r

.
,
.

Du ring t h e a na pha s e ( Fi g 6 H) th e ch r o m os o m e s f orm e d

.
, ,

by splitting m o ve a l o ng th e spindl e fib e r s to th e c e nt r oso m e s .

AS a r e s ult e v e r y ch ro m oso m e p r e s e nt a t t h e e nd o f t h e p ro ph a se
( )
F s e nd s h a lf o f i t s ch ro m a tin t o e ith e r e nd o f t h e spindl e T he .

m e ch a nis m th a t b ring s ab o ut this mig ra ti o n i s a s ye t so m e wha t

in qu e s tio n . F ib e r s a r e u s u a lly l e ft b e tw e e n t h e s e p a r a ting
ch ro mo so m e s ; the se a r e kn o wn a s i nte r zona l fi be r s ( H if ) -
,
.

T h e te lopha se ( F i g 6 I J) i s a s t a g e o f r e co n s t r u cti o n fro m

.
, ,

whi ch the nucl e i e m e r g e in a r e s ting co nditi o n ; th e ch ro m a tin

b e co m e s s c a tte r e d th r o ugh o ut t h e nu cl e u s whi ch i s a gain e nv e l
,

Op e d by a d e finit e m e mb r a n e ( J ) ; t h e ce nt ro s o m e divid e s a nd ,

with th e ce nt ro sphe r e t a k e s a p ositi o n n e a r t h e nucl e u s F in ally

,
.

th e cy cl e i s co mpl e t e d by t h e co n s t r i cti o n o f th e ce ll int o t w o

d a ughte r c e ll s ( I J ) e a ch o f whi ch i s in co nditi o n to c a r r y o n the
,

r e gula r m e t ab o li c p ro c e s s e s until it i n tu r n b e c o m e s re a dy to

di vide .

T h e or i gi n of the s tr u ctu r e s th a t t a k e a n a ctiv e p ar t in mit o s i s

i s no t d e finit e ly kn o wn Only t h e ce ntr os om e s a r e r e p r e se nt e d
.

in the r e s ting c e ll ; the se u su ally a r i se fr o m th e di vi si o n o f pr e

e xisting ce nt roso m e s but in c e r t a in case s th e y a r e wh o lly a b s e nt
,

(hi gh e r pl a nt s) o r th e i r e xi s t e nce i s qu e s ti o n a bl e ( P r o t o z oa )
, .

Th e chr om os om e s a r e fo r m e d di r e ctly by a co nd e n s a ti o n o f ch ro
m a tin The origin o f th e s pi ndle fi be r s i s v a ri abl e So m e tim e s
. .
3 2 AN INTRODU CTION TO ZOOLOGY

the y a re form e d f ro m the cytopl asm a nd s o m e tim e s th e y a ris e

,

fro m th e linin n e tw ork in t h e nucl e u s Linin do e s no t h o we v e r

.
, ,

di ffe r g re a tly in ch e mi c a l c o mp ositi o n f r o m th e cyt opl as mi c

s ub s t a nc e a nd th e spindl e fib e r s a r e th e r e f or e
, ,
a lw a y s c o mp o s e d
,

o f achr om a ti c m a t e r i a l T h e nu cle olu s i s a pp a r e ntly o f no imp o r

.

t a n ce in mit o sis ; it d e g e n e ra te s during th e e arly stage s ( Fi g 6 .

,

G H n) a nd i s re fo r m e d du r ing th e t e l oph a s e ( J)
, ,
T h e t wo ce ll s
.

tha t r e sult fr o m mit osi s m ay b e e qu a l (J) or un e qu al in size T h e .

divi si o n of th e cyt opl asm app e a r s t o b e of littl e imp or ta n ce but ,

with ra re e xc e pti o n s the r e se e ms t o b e an e qu al divisio n o f

chr o m a tin ; thi s i s app a r e ntly th e e sse nti al p r o ce s s .

Unde r or din a ry co nditi o n s e v e r y a nim al d e ve l ops fro m a singl e

ce ll a nd s inc e th e ch r o m a tin p e r si s t s f ro m o ne c e ll g e n e r a ti o n to
, ,

a n o th e r th e ch ro m o so m e s a r e co n sid e re d by m o s t z o ol o gi s t s t o
,

be th e be ar e r s of he r e di ta ry qu a li ti e s f ro m p a r e nt t o Off sp ring .

E v e r y sp e ci e s o f a nim a l h a s a d e finit e numb e r o f ch r o m oso m e s

tha t app e a r wh e n th e ce ll s o f i ts b o dy unde rgo mitosi s Thus .

sixte e n a re chara ct e r i sti c Of th e c e ll s of o x e n guin e a pig s a nd m a n;

, ,

th e g r assh opp e r h a s tw e lv e ; th e b r in e s h r imp (A r te mi a ) o ne ,

hund r e d a nd sixty e ight ; th e r o und w o r m (As ca r i s) fo u r or tWO ,

.

T h e l a st e x a mpl e illu s t ra t e s o ne o f th e unu su a l case s in which t wo

in di vidu a ls th a t ar e mutu ally f e r til e h ave a di ffe re nt numb e r o f
ch ro m oso m e s An e ve n numb e r o f ch r o m oso m e s i s ch a ra cte ri s tic
.

o f m os t a nim a l s but r e ce nt r e se a r ch e s h a v e d e m o n s t ra t e d th a t
,

s o m e form s p a r ticul arly th e m a l e s o f in se cts h av e a n o dd

, ,

numb e r .

( 2) Ami tos i s — Alth o ugh mit osi s i s c o n side re d th e typic al

.

m e tho d o f c e ll di vi s io n th e multiplic a ti o n o f nu cl e i by di re ct
,

divi sio n or a mi tosi s i s no t unco mmo n Thi s p r o ce ss ( Fi g 7)

. .

d o e s no t h a v e su ch a de fini t e se r i e s o f s ta ge s a s mito si s a nd is ,

much simpl e r T h e nucl e u s e l o ng a t e s a nd th e nu cl e o lus di

.
,

vide s o ne h alf p as sing t o e ith e r e nd o f th e nu cl e u s So m e tim e s

, .

th e nu cl e olu s r e m a in s a t o ne e nd a ne w o ne b e ing f o r m e d a t th e
,

o pp os it e e nd T h e nu cl e u s divid e s by o ne o f th re e m e th o d s :
.

( )
a o ft e n th e nucl e us i s pinch e d in t w o in th e midd e or ( )
b a
l
,
 T HE C ELL AN D T HE C ELL TH EOR Y

pl a te is for m e d a t the pl an e of di visi o n which l a te r b e co m e s

do ubl e and th e n th e two pl a t e s se p ar a t e o r (c) tw o nucl e ar m e m
, ,

b ra ne s a r e built up i n side o f th e old m e mb r a n e whi ch the n b r e aks ,

do wn allo wi ng th e d a ughte r nucl e i to e s cap e

, .

Th e ce ll b o dy is th e l a st t o di vid e in a mit o sis a nd in m a ny , ,

p e r haps the m aj ority of c ase s it doe s not do so a t all ( 2 1 p

, ,
.

Am it o ti c nu cle ar d ivi s i o n i n th e f o lli cl e ll s o f a cr i ck e t s e gg ( From

’
FI G 7
. . ce .

K pn )
Da hlgr e n a nd e er .

The amit o tic p r o c e ss is u s u ally a sign of se ne sce nce o r d ec ad e nce in

a c e ll . M a ny ce lls divid e by mit o si s so l o ng a s th e r e i s a ctu a l
divi si o n o f th e cytopl asm as we ll a s o f the nucl e us but if th e c e ll ,

b o dy fa il s to divide th e nucl e us a l o n e m ay c o ntinu e t o multiply

,

a mit o ti ca lly p r o ducing a m ul tinucl e a t e c e ll

,
T h e c o mm o nly .

r e c e iv e d id e a a t p r e se nt c o nc e r ning a mit os is i s th a t it i s a t e r mi
, ,

na l p r o ce ss in th e c e ll s lif e a ctiviti e s a nd is a m e th o d o f s e cu ring

’
,

m o r e nu cl e a r su rfa c e f o r u se in for c e d m e t ab olism o r s e c re ti o n

3 9)
I n co n cluding thi s cco unt o f c e ll divisi o n two p oint s ar e w or thy
a

o f Sp e ci a l e mph a si s F i rs t with r e g ar d to t h e conti nu i ty of the

.
,

chr om a ti n it m a b e s id th t t h ch m tin is c ntinu o us

, y a a e ro a o

f ro m o ne ce ll g e n e ra ti o n t o a n o the r T he c e ll s r e s ulting f r om
.

mito si s m ay diff e r g r e a tly in siz e but th e ch ro m a tin s e e m s to

,

b e di vid e d e qu a lly b e tw e e n th e m with g r e a t e x a ctn e s s Se c o nd

'

.
,

ce lls a r e ne ve r k nown to a r i s e e xce t fr om

p p r e e xi s ti n ce lls
g Th e se .

t w o f act s a r e p e r h a p s th e m o s t imp o r t a nt f o r u s t o k e e p in mind

a s we
g o o n t o s tudy t h e m o r e c o mpl e x p r o bl e m s o f fe r tiliz a ti o n
a nd c e ll divi si o n in t h e m a ny ce ll e d a n i m a l s f o r g r o wth in e v e ry
-
,

M e t a zo o n i s re a lly n o thi ng m o re th a n ce ll divi si o n and ce ll

gr owth .
 AN IN T R ODU CTION TO Z OOLOGY

3 . T H E C E LL THE OR Y
Owing to th e imp or ta nt pl ace th a t c e ll s h a ve o ccupi e d in th e
de ve l opm e nt of o u r funda m e nt al biol ogic al ide as it se e m s p ro p e r ,

t o co n sid e r b r i e fly th e o r igin a nd g rowth o f th e ce ll th e o r y C e ll s .

w e re fi r s t de scr ib e d by Hoo ke a n E ngli shm a n in 1 6 6 5 He

, , .

publish e d a figure sh o wi ng th e minute st r uctu re o f co r k ( Fi g .

FI G 8
. . Fa csi m il e of a fi gur e by Hoo k e r e p r e se ti
n ng ce ll s o f cork .
( Fr o m
F a r m e r i n La nk e s t e r
’
s Z l oo o gy .
)

T he gul ar a r r ange m e nt of th e c omp a r tm e nt s in this tis su e

re

r e mind e d him o f t h e ce ll s o f t h e m o nk s in a m o n a s t e r y F or thi s .

”
r e a s o n th e y w e r e giv e n t h e n a m e c e ll whi ch the y b e ar t o thi s
,

d ay . Se v e r a l l a t e r inv e s tig a t or s ob s e r v e d ce llul ar s t r uctu r e s in

th e ti s s u e s o f pl a nt s a nd a nim a l s with o ut r e a lizing th e i r imp o r
t a nc e .

In 1 8 3 3 B ro wn a n E ngli sh b o t an is t de s c rib e d th e nu cl e u s a s a
, , ,

co n s t a nt e l e m e nt o f t h e ce ll It w a s no t h o w e v e r until th e tim e
.
, ,

o f Schl e id e n a b o t a ni s t a nd Schw a nn a z o o l o gi s t th a t th e ce ll
, , , ,

the o ry w a s e s t abli sh e d Schl e id e n in 1 8 3 8 publi she d a sm all

.
, ,

p amphl e t in whi ch h e a dva n ce d th e ide a th a t a ll pl a nt s a r e co m

p o se d o f ce ll s A y e a r l a te r h i s coll e a gu e Schwa nn b ro ught
.
, ,

fo r th a m o r e p r e t e ntio u s w o r k in which h e m a de th e sam e ge n e r ali

3 6 AN INT RODU CTION TO ZOOLOGY

th a t Ko lli k e r , R e m ak , N a ge li , a nd Ho fme i s te r p e n e d th e w ay
O

t o a n und e rs t a ndi ng o f th e n a tu re o f e mb ry ol o gic al d e v e l op

m e nt and th e law o f g e n e tic co ntinuity lying a t th e b asis o f
,

inh e r ita nc e It w as th e c e ll the o ry ag ain whi ch in th e h ands

.
-
,

o f Go o d si r Vi r ch o w a nd M a x Sch ultz e
, ,
in augur a t e d a ne w e ra
,

in th e hi s to ry o f physi ol ogy a nd p a th o l o gy by sho wing th a t all

,

th e v a r i o us functi o n s o f th e b o dy in h e a lth a nd in di s e ase a re

, ,

but the o utwa r d e xp r e ssio ns of c e ll a ctiviti e s And a t a s till .

l a t e r d ay it w as th r o ugh th e c e ll -the or y th a t He r twig F ol Va n , ,

B e n e d e n a nd St r a sb u r g e r s o lv e d th e l o ng s t a nding riddle o f th e
,
-

fe r ti liz a ti o n of th e e gg and th e m e ch a ni s m of h e r e di t ary t ran s

mi ssio n N O o th e r bi o l o gica l g e n e ra liz a ti on s a v e o nly th e the o ry
.
,

o f o r g a ni c e v o luti o n,
h as b r o ught so m a ny app a r e ntly div e r se
ph e n o m e n a unde r a co mm o n p o int o f vi e w o r h a s a cco mpli she d
,

m o r e f o r th e un fic ti
i a o o n f kn o wl e dg e .T h e c e ll -th e o ry mus t
the r e fo r e be pl a ce d b e side th e e volutio n the o ry a s one of the
-

f o und a ti o n sto n e s o f m o de r n biol o gy ( 2 5 p ,

.
 C HAP T E R I V

AM EBA

( Ame ba pr ote u s Le idy)

Ame ba pr ote us ( g 9)
F i . is a o n e -ce ll e d a nim a l a b o ut 2 5 mm . .

( fi m inch) in di a m e te r , a nd is , the r e for e , invi sibl e to th e n a k e d e ye

l
.

Unde r th e co mp o und micr o scop e it app e ar s i

a s a n r re gul ar co l or

FI G 9 . Ame ba pr ote u s
. . I ,
l e u s ; 2 cont ra ctil e v acuo l e ;
nu c , 3 , p s e u d o po d i
a ,

d o tt e d li ne l e a d s t o e cto pl a sm ; 4 f oo d va cu o l e s ; 5
, , gr a ns i of sa nd .

( From Shipl e y a nd M a cB r id e a f te r Gr ub e r )
.

l e ss p ar ticl e of a nim a t e d j e ll y which is co n st a ntly ch a nging its

s ha p e by th r u s ting o u t fin e r -lik e p r o c e s se s ( Fi
g g 9 .
,

37
3 8 AN I N T RODU CTION TO ZOOLOGY

H a b i ta t and C o ll e cting Am e ba pr ote u s live s in f re sh -w a t e r

.

p o nd s a nd s t re a m s It . m a y b e Obt a in e d f o r l a b o ra t o r y u se f r o m
a v ar i e ty o f pl a c e s s uch a s th e o r g a ni c o o z e f r o m d e c a ying v e e
, g
ta ti o n o r th e l o w e r su r f ace o f li ly p a d s P e r h a p s th e m o s t ce r .

t a in m e tho d i s tha t s ugge s t e d by P ro f e sso r J e n ning s Ab o ut

t wo w e e ks b e f o r e th e sp e cim e n s a r e n e e d e d a m a ss o f p o nd w e e d ,

( a tophyllu m i s th e b e st ) sho uld b e g a the re d p la ce d in fla t

C e r ,

di she s a nd imm e rse d in wa t e r T h e ve ge ta ti o n soo n de cays a nd

,
.
,

a b ro w n s cum a pp e a rs o n th e su r f a c e In thi s s cum Am e b a s .

m a y b e f o und .

G e n e r a l An a to my — T wo re gi o n s a r e d i s tingui sh a bl e in th e
.

b o dy o f Ame ba a n o ut e r co l o rl e s s l a y e r o f cl e a r cyt opl a sm th e

, ,

e ctos a r c a nd a c o mp a r a tiv e ly l a r g e c e nt r a l m a ss o f g r a nul a r cyt o

,

pl a s m th e e nd os a r c A singl e cl e a r Sphe r ica l b o dy u s ually lying

,
.
,

n e ar the e nd o f the a ni m a l awa y f ro m t h e di re ctio n o f m o tio n a nd ,

di sapp e a ring a t m ore o r l e s s r e gul a r int e r va l s i s th e contr acti le o r ,

p u l s a ti ng v a cu ole ( g 9
F i S u s p e nd e.d in ,
t h e e nd o s a r c i s a

nu cle u s ( Fi g 9 I ) u s u a lly o ne o r m o r e f ood vacu ole s ( Fi g 9

.
, ,
.
,

m a te ri a l r e a dy f o r e x cre ti o n fo r e ign s ub st a n ce s s u ch as g ra ins o f

,

sa nd ( Fi g 9 .
,
a nd undig e s t e d p a r ticl e s t h e a m o u nt o f th e ,

l a tte r de p e nding up o n th e fe e ding a ctivity of th e Sp e cime n a t th e

tim e whe n e x a min e d .

F ro m thi s d e s c ripti o n it will b e n o t e d th a t A m e ba pr ote u s

co nta in s a ll Of th e e sse nti a l co ns titue nt s Of a ce ll It i s m o re ov e r .
, ,

Simpl e in s t r u ctu r e s h o w s a numb e r o f phy s i o l o gi ca l a ctiviti e s

,

in the i r simpl e s t fo r m i s o ne o f the mo s t p rimitiv e o f all a nim a ls

, ,

a nd is e a s ily o bt a in e d F o r th e se re a s o n s it h a s b e e n a nd s till is
.
, ,

a f a v o r it e s ubj e ct f o r s tudy .

D e tai le d Ana to my T h e e ctopla sm ( Fi g 9 3 ) is e a sily d i s

.
-
.
,

ti ngui sh e d f r o m th e e nd op la s m b e ca us e o f t h e a bs e n ce within it
o f g r a nul e s . T h e e ct opl as m i s fi r m e r th a n th e e ndopl a s m p r o b ,

a bly b e ca u s e t h e o ut e r p r o t o pl a sm t e nd s t o s ti f f e n und e r t h e

influe nce o f s u rface t e n si o n .

T h e e nd opla s m o ccupi e s th e ce nt ra l p o r ti o n o f th e b o dy Be .

ing l e ss de n se tha n th e e ctop l a sm it c o nt ain s withi n it a ll o f th e ,

 AM E B A
39

l arg e gra nul e s N 0 fix e d lin e

. of s e p a ra tio n b e twe e n it a nd th e
e ct o pl a sm i s p o s s ible .

T h e nu cle u s )( g 9,
F i . Iil y
i s no t
se e n in living
e as sp e cim e n s .

In a nim al s th a t h av e b e e n p r op e r ly kill e d and s t a in e d it app e ars a s

a sph e r ic a l b o dy lying in th e e nd opl a s m I t s p o siti o n i s no t
.

d e finite but ch a ng e s dur ing th e m o v e m e nts o f th e Am e ba It h as

,
.

a fi r m m e mb r a n e a nd c o nt a ins a g r e a t m a ny sph e r ic a l p a r ti cl e s o f

chro m a tin s c a tt e r e d ab o ut in th e nu cl e a r s a p D u r ing th e life .

o f A m e ba b e fore t h e p e r i o d o f r e p r o du cti o n t h e nucl e us pl a y s

, ,

a n imp o r t a nt r O le in th e m e t a b o li c a ctivity o f th e c e ll Thi s h as .

b e e n p rove d by e xp e rim e nts in whi ch th e a nim al w a s cut in two .

Inv ari ably th e vit a l p r oce sse s w e r e di s tu r b e d in t h e e nucl e a t e d

f ra gm e nt and d e a th r e s ult e d P r o fo und ch a ng e s in th e nucl e us
,
.

t ak e pl ace dur ing r e p r o ducti o n ; the se will b e d e sc r ib e d in d e t ail

l ate r.

T h e contr a cti le va cu ole ( Fi g 9 2 ) i s a cl e ar s p a c e fill e d with a

.
,

fluid l e ss de n se th a n th e s u r r o unding p r o t opl a sm It d e r iv e s i ts .

n a m e f ro m th e f act th a t a t m o r e o r l e s s r e gul a r int e rv a ls it su d

d e nly di s a pp e a rs i ts w a ll s h a ving co nt ra ct e d thu s f orcing o u t
, ,

th e c o nt e nt s Th a t th e v a cu o l e disch a r g e s to t h e o ut sid e o f th e
.

b o dy h a s no t b e e n de finit e ly Ob se r v e d in A me ba no d o ubt b e ca u se ,

t h e fluid i s u s u a lly e xp e ll e d o n th e upp e r s u r f a ce o f th e b o dy a nd

the re fo r e c a nn o t b e s e e n “ At fi rs t th e v a cu o l e li e s n e ar
th e nucl e u s but a s it g r o w s it b e co m e s s e p a r a t e d fr o m th e l a tt e r
, , ,

a nd a t t h e tim e o f i t s c o nt r a cti o n li e s a t t h e e nd o f th e b o dy f a r

t h e s t f r o m th e a dv a n cing p s e ud o p o di a a t wh a t i s s o m e tim e s c all e d

,

t h e p o s t e r i o r e nd I ts r e a pp e a r a n ce i s a lw a y s so m e wh e r e n e a r
.

i t s p o int o f d i sa ppe a r a nc Whil e s till sm all it i s ca r ri e d al o ng

by th e st r e a ming p ro t o pl a s m b ack to a p o s itio n n e a r t h e nucl e us ,

whe re it co mpl e t e s its d e ve l o pm e nt ( 2 9 p 8 8 Fi g ,

.
,
.

T h e fu ncti o ns o f t h e c o nt r a ctil e v a cu o l e a r e e x cr e t o r y a nd r e s

p ra t o ry a s i s e xpl a in e d o n p a g e 5 0
i , Se v e r a l inv e s tig a t o r s h a v e
.

r e ce ntly r e g a r d e d it a s a hyd r os t a ti c o r g a n r e gul a ti ng t h e qu a ntity

,

o f w a t e r co nt a in e d in t h e b o dy o f t h e a nim a l a nd SO i t s w e ight ,
.

Thi s wo uld a l so a ffo r d a me a n s o f g e tting ri d Of the wa t e r t a ke n

 AN INTRODU CTION TO Z OOLOGY

fl -
’ p

G 1 1
FI G . 10 . Am e ba pr ote u s A, B , C , D , E , F , s h o w s u cce ss iv e s t age s i n th e
.

gr o wth o f th e co nt r a ctil e v a cuo l e (v) ; G , H , I , a nd J , Sh o w f o u r s t age s

in t h e t
co n r a c tio n o f th e v a cuo l e .
( Fro m C a l k i ns )
.
 AM EBA 41

in by the p rot opl a sm th ro ugh the b o dy wa ll a nd co nse qu e ntly

r e gul a ting t h e t e n si o n b e tw e e n t h e p r o t o pl a s m a nd th e su rr o und

i ng w a t e r . D e a th by d ifli u e nce i s thus p r e v e nt e d .

F ood vacu ole s ( Fi g 9 4) a r e p ro duc e d wh e n fo o d is t a ke n int o

.
,

t he b o dy . Such a v a cu o l e h a s a s i ts nucl e u s a p a r ticl e o f nut r i

tive m a t e ri al and is a s or t of t e mp ora r y s to m a ch .

Lo com o ti on Alth o ugh e xt re m e ly simpl e in st r uctu re Ame ba

.
,

c arri e s o n p ra ctic ally all o f the vit a l a ctiviti e s chara ct e ri s tic o f

the hi gh e r a nim a ls It is cap a bl e o f a ut o m a tic m o ve m e nt o f
.
,

r e a cting t o v a r i o us stimuli o f c a r r ying o n m e t a b o lic p ro c e ss e s

, ,

o f g ro wth a nd Of r e p ro ducti o n
,
Th e s e a re all fund a m e nt a l
.

p rop e r ti e s o f p ro topl a s m ( C h ap II p 1 0) a nd are he re e x.

,
.

hi bi te d in th e i r simpl e st fo r m .

T he locomoti ve fu ncti on is l o c a t e d in th e e ct o pl a sm T he .

m o til e orga ns a re fi nge r like p ro t r usio n s o f cyto pl a sm c all e d

-

p s e u d op o d i a A. p se ud o p o dium is f o r m e d in t h e f o ll o w ing m a n

ne r . T he e ct opl a s m bulg e s o u t a nd e nl a r ge s until a blunt pro

j e c t i o n i s p r o duc e d ; t h e e nd o pl a sm th e n fl o w s int o it T h e .

r e s ult is a m o v e m e nt o f t h e e nti re a nim a l in t h e di r e cti o n o f th e

p se udop odi um If m ore th a n o ne i s fo r m e d a t th e sa m e time

.
,

the re o ccur s a s tr uggl e f o r sup r e m a cy until fin ally o ne s u r vive s

whi l e the o the r s flow b a ck a nd g r a dua lly di sa pp e ar Ame ba .

m ove s the r e fore by thr usting o u t p se ud op o di a a nd th e n fl o wing

, ,

into the m It i s n e ce ssa r y t o di s tinguis h b e twe e n p se ud op o di a

.

th a t a dhe re t o the su r fa c e o f so m e Obj e ct a nd th o s e tha t a re

th r us t o u t f re e ly into th e s u r ro unding fluid Since a n hypo the sis ,

tha t e xpl a ins the form a ti o n o f o ne do e s no t a lwa ys a cc o unt f o r

th e o th e r .

The re are th re e p rincip al the orie s whi ch a tte mpt to e xpl ain
the f o r m a ti o n o f ps e ud op o di a ; th e s e m a y b e o utlin e d b r i e fly as
foll o ws :
( 1 ) Th e Adh e r e n ce Th e ory It w as l ong a go disc o ve re d th a t
.

if a drop of in or gani c fluid whi ch i s sp r e a ding o n a s olid su r fa c e

i s m a d e t o a dh e re o n o ne Sid e m o re s t ro ngly th a n th e o th e r it ,

Will m ov e t o war d the fo r m e r sid e Thi s fa ct sugge ste d th a t the

.
4 2 AN INTR ODU CTION TO ZOOLOGY

b o dy o f the Ame ba m ay move in a c e r tain di re cti o n f or the s ame

re aso n Thi s the or y ho w e ve r a cco unts only f o r the
, ,

ps e udop odi a tha t a re a tt a che d a nd doe s no t e xpl ain fre e ps e ud o

,

p odi a .

( 2) Th e S ur f ace T e n s i on Th e ory Accor ding to th e s u r fa c e

.

te n si o n th e o r y l o c a l ch a ng e s in th e su r f a c e t e n sio n p ro du ce
,

cu rr e nt s which m ove for wa r d in th e c e ntr al a xi s a nd b a ckwar d

a l o ng th e su r f a c e ( 2 8 It h as b e e n sho wn ho w e ve r th a t
, , ,

th e cu rr e nts d o no t r u n in thi s m a nn e r

(3 ) T h e C o n tr ac ti l e Th e or y M a ny e ar ly inv e s tiga t o r s he ld
.

th e Vi e w th a t th e c o nt r a ctility o f th e p r o t opl as m a c co unt s f o r

ch a ng e s which t a ke pl a c e in a m o ving Am e ba Thi s the o r y was .

l a te r give n up by m os t z ool o gi s t s b e c a use the cu rre nts in the pro

topla s m b e g a n a t th e p o int of a dva nc e a nd e xt e nd e d b a ckwa r d

FI G . 1 1 . D i agr a m of th e m o v e m e nt s o f a pa r ti cl e a tta ch e d to th e o u t e r sur

f a ce oi Ame ba ve r ru cosa i n s i d e V i e w
, (Fr om J e nni ngs )
. .

inst e a d o f c o mme ncing a t the hinde r e nd or in th e c e nte r Two .

re ce nt a uth or s h a v e a g a in b r o ught f o r th e vid e n ce which t e nd s

t o s h o w th a t a ft e r all th e c o nt r a cti o n th e o r y i s th e co r r e ct int e r

re t a t i o n J e nni ng s ( 3 9) by mi xing s o o t in w a t e r w a s a bl e t o
p .

Sh o w th a t Am e ba ve r r ucosa r e se mbl e s a n e l as tic sac fill e d with a

fluid Sp e cim e n s w e r e pl a c e d in wa t e r c o nt aining so me fi ne s oo t

. .

T h e p ar ticl e s b e c a m e a tt a ch e d t o th e sur fa c e o f th e a nim a l a nd

th e cu rr e nt s in th e e ct o pl a sm c o uld e a sily b e d e t e r min e d by
w a tching th e i r m ove m e nt s F ro m the s e ob se rva tio n s th e foll o w
.

ing co ncl usi ons we re r e a che d In an a dvan ci ng Am oe b a sub

.

sta nc e fl ows fo r w ar d o n th e upp e r su r fa c e rolls o v e r a t th e ant e ,

r i o r e dg e co ming in c o nt a ct with th e s ub s t r a tum th e n r e m a ins

, ,

quie t until the b ody of the Amoe b a has passe d o ve r i t I t the n .

44 AN IN T R ODU CT ION TO ZOOLOGY

In a dv a ncing th e Ame bce e xt e nd the a nte ri or e nd fre e in th e

w a t e r a nd a tta ch it a t or n e ar th e tip a nd the n co ntr a ct . At th e

FI G . 1 3 . P h o t o gr ap h s of Am e ba pr ote u s i n s i d e
vi e w t o s h o w m e th o d o f l o co

m o ti o n A B a nd C S h o w a spe ci m e n a tt a ch e d a t t wo po i nt s a a nd b; th e
.
, ,

p se u do po d pr o j e cti ng f ro m o ne e nd b e nd s d o w n t o th e su b s t ra t u m a s
s ho w n i n B a t d ; C s h o w s t wo m o r e a tt a ch e d p se u do p o d s a t c a nd f ,

and se v e ra l f re e pse u d o po d s a t e .
 AM EB A 45

FI G . I 3 conti nue d . D a nd E ho w the e xt e ns io n a nd a tt a ch m e nt o f a l o ng

S

thi n pse u d opo d .

( Fro m De lli nge r i n J ou r n E xp Z ool )
. . .

s a m e ti me th e p os t e rio r e nd i s co ntr a cting a nd the s ubst a nc e

thu s push e d a nd pull e d fo r wa r d g o e s to fo r m th e ne w a nte rio r
e nd Thi s c o ntinu e s a s l o ng a s th e Am e ba a dv a nc e s ( Fi g 1 3
. .
,

A B C)
, ,
Oft e n t h e a nt e r i o r e nd i s pu s h e d a l o ng t h e s ub s t r a tum
.
,

but no a tt achm e nt s f o r m e xc e pt a t d e finit e p oint s .

In o the r c ase s th e a nt e r i or e nd i s lifte d f r e e and the n cu rve s

do wn to th e sub s t r a tum a nd a tt a ch e s fo r ming a l o ng l oop T h e ,
.

p oste r i o r e nd i s the n r e l e a s e d a nd th e s ub s t a nc e fl o ws o ve r to
,

t he a nt e r i or e nd At th e sa m e tim e a n o th e r a nt e r io r e nd i s e x
.

t e nd e d (3 4 p 3 5 1 F i g I 3 D E) Whe n cr e e ping o n th e c e i l
,
.
,
.
, ,
.

ing th e m ov e m e nts a r e th e s a m e a s whe n c r e e ping o n th e fl o or .

Wh e n b e ginni ng to m o v e p se ud op o di a a r e e xte nd e d in all di rce

,

tio n s until o ne b e co m e s a tt a ch e d whe n th e a nim al a dva n ce s in ,

th e di re cti o n o f th e p o int o f a tt a chm e nt Th a t a co nt r a ctil e .

subst a nc e i s p r e se nt in Am e ba a nd a cco unt s f o r i ts m ove m e nts

se e m s t o b e p ro v e n by D e lli n e r s Ob se r v a ti o n s
’

g .

M a ny a tt e mpt s h a v e b e e n m a d e t o i mi tate the move me nts of

45 AN IN T RODU C TION TO Z OOLOGY

Ame ba by m e a ns o f in or g a nic sub s t ance s in a n e nde av or to di s ,

c ov e r th e physi ca l o r ch e mi c a l n a tur e o f i ts l o co m o ti o n
E v e n wh e n a n app a r e ntly re li abl e i mi t a ti o n is p r o du ce d w e can ,

no t b e ce r t a in th a t th e f o r c e s a t w o r k a r e a ctu a lly th o se whi ch

c a u se th e m o ve m e nt s of Ame ba P r a cti ca lly all of th e imit a ti o ns
.

thu s f ar r e p or te d de p e nd o n s urf a c e t e n sio n .

One m e th o d o f p ro du cing a m e b o id m o v e m e nt s i s a s f oll o w s

A l ar g e d r op o f m e r cu r y i s pl a ce d in a fla t—b o tt o m e d w a tch
gla ss a nd c o ve re d with 1 0 pe r ce nt nitr ic a cid A pi e c e of po .

t a ssi u m bich r o m a t e wh e n pl a c e d n e ar th e m e r cu r y p r o du ce s a
s o luti on whi ch ca u se s l o c al l o w e r ing o f th e s u r f a c e t e n sio n o f
t h e d r op a nd re s ult s in t h e f or m a ti o n o f p r o j e cti o n s a nd m ov e
,

m e nt of th e m e r cur y in v ar io u s di r e cti o ns .

M e ta b o l i s m . T h e v ar i o u s m e t ab oli c a ctiviti e s o f a nim al s

w e r e di s cu sse d in C h apt e r II p ag e s 1 3 t o 1 5 In A m e ba th e se
,
.

p ro ce sse s a re s e e n in th e i r simpl e s t f o r m a nd will the r e fore b e , , ,

c o n side r e d in so m e de t a il in spite o f th e d a ng e r o f r e p e titi on .

T h e e nti r e s e r i e s o f p r oce ss e s c o nn e ct e d with th e m a nufa ctu r e

a nd d e s t r ucti o n o f p r o t o pl a s m a r e t h e ing e s ti o n o f f o o d dig e s ,

ti o n e g e s tio n a b so r pti o n ci rcul a ti o n a s simil a ti o n di ssimil a

, , , , ,

ti o n s e c r e ti o n e xc r e ti o n a nd r e spi ra ti o n
, , ,
.

F oo d — T h e foo d o f A m e ba co n s i s t s o f v e r y sm a ll a qu a tic
pl a nt s s uch a s Osci lla r i a a nd d i a t o m s P ro to z o a b a cte r i a a nd
, , , ,

o th e r a nim a l a nd v e g e t a bl e m a tt e r A c e r t a in a m o unt o f ch oic e

.

o f f o o d i s e x e r ci s e d o r i t s b o dy w o u ld b e c o m e o v e r l o a d e d with
,

p a r ti cl e s o f sa nd a nd o th e r indi ge s tibl e m a t e ri al am o ng whi ch it

liv e s F u r th e r m o r e it s e e m s to e vin ce a p r e fe r e n ce f or di a to m s
.
,

w hi ch o ne w o uld think t oo l a rg e f o r e a s y co n s umpti o n .

Thi s .

a pp ar e nt ch o i ce o f f o o d m a y b e d ue t o o r din a r y phy sic a l l a w s o f

fluid s .

In ge s ti on . T h e ing e sti o n o r t a king in o f f o o d o ccu r s with

, ,

o u t th e a i d o f a m o uth F o o d m ay b e e ngulf e d a t a ny p oint o n

.

th e s u r f a ce o f th e b o dy but it i s u s u ally t a k e n in a t wh a t m a y
,

b e ca ll e d t h e t e mp o r a r y a nt e ri o r e nd th a t i s th e p a r t o f th e b o dy
, ,

t o w a r d th e di re ctio n of l oco mo tio n Je nni ng s de scr ib e s inge s

.
 AM EB A 47

tio n a s foll o ws The A me ba flo ws a g a in s t th e f o o d p a r ti cl e

.
,

whi ch doe s no t a dhe re but t e nds to b e pu she d fo r wa rd a wa y

f ro m the a nim a l ( Fi g 1 4 I ) Th a t p a r t o f th e b o dy di re ctly
.
,
.

b a ck o f th e foo d ce a s e s i t s fo r wa r d m o ve m e nts w hi l e o n e ithe r ,

s id e a nd a b o v e p se udop o di a a r e e xt e nde d which g r a du a lly f o r m

,

a co n ca vity in which t h e f o o d li e s ( F i g 1 4 2 ) a nd fin a lly b e nd .

,

a ro und th e p a r ti cl e ( F i g 1 4 3 ) until th e i r e nds m e e t a nd fu se

.
,

FI G . 14 . A me ba i nge s ti ng a E u gle na cys t . I ,

2, 3 , 4, s u cce ss ive s t age s i n the
p r o ce s s .
( Fr o m Je nni ngs ) .

( Fi g . 1 4, m all a m o unt o f w a t e r is t ak e n in with the foo d

A s ,

so th a t th e r e i s f o r m e d a v a cu o l e wh o se s id e s w e r e f or m e r ly t h e

o ut sid e o f t h e b o dy a nd wh o se c o nt e nt s c o n s i s t o f a p ar ticl e o f
,

nut ritive m a t e ri a l su sp e nde d in w a t e r T h e whol e p ro c e ss o f .

food ta king o ccupi e s o ne or m or e minute s de p e n di ng o n th e

-
,

ch a r act e r o f t h e f o o d A m e ba i s no t a lw a y s s u cc e ss f u l in a c
.

co mpli shi ng wh a t it und e r t a k e s but wh e n it d o e s no t c a ptu r e i t s

,

p re y a t o n ce it se e m s to sho w a p e rs is te nc e u su ally o nly a ttribut e d

,

t o highe r o rg a ni s m s N O d o ubt th e r e a cti o n s in f oo d ta king

.
-

de p e nd up o n b o th m e ch a ni ca l a nd che mic a l s timuli

Imit ati on s o f th e e ngulfing o f f oo d by Am e ba h av e b e e n d e
vi se d b a se d o u th e th e o r y tha t i nge s ti o n d e p e nd s o n th e physical
,

a dh e si o n b e tw e e n t h e liquid p ro topl a s m a nd t h e so lid fo o d .

Drop s o f w a t e r glyce r in whit e o f e gg e tc will dr a w into co n

, , ,
.
,
48 AN IN TR ODU C TION TO Z OOLOGY

t a ct a nd e ngulf solid p ar ti cl e s o f v a r io us kinds One o f th e m o s t .

ing e ni o us o f th e se imita ti o n s i s th a t r e p or te d by Rh u m ble r of

th e ing e s ti o n o f a fila m e nt o f Osci lla r i a A th r e a d Of sh e ll a c
is b r o ught int o co nt a ct with a d r op o f chl o r o fo r m in a w a tch
gl a ss of w a t e r T h e d rop a dhe r e s to th e fila m e nt l e ngthe ns
.
,

a l o ng it a nd b e ca u se o f i t s t e nd e n cy t o a g a in b e c o m e sph e r ic a l
, , ,

s ucc e e d s in b e nding th e no w so ft e n e d th r e a d T h e t e n si o n o f t h e .

s u r f a c e film g r a du a lly d ra ws in m or e of th e fil a m e nt unt i l fin a lly ,

FI G . 1 5 . Ame ba ve r r ucosa d e vo ur i ng a fil a m e nt of Os ci llar i a .

( Fr o m R h um
b l e r i n Ar chiv f E nt wi ck -m cch
. . .
)

th e wh o l e thr e a d i s e mb e dd e d in a co mpli ca t e d co il within th e

d rop T h e ing e s ti o n of Osci lla ri a by Am e ba i s quit e simil a r
.

( Fi g .

C h oi c e foo d m ay al so b e imit a t e d with in or g a nic subst an ce s

of .

F o r e x a mpl e a d ro p o f chl o r o f o r m in a w a t ch gl a ss o f w a t e r will

,

t a k e in Sh e ll a c p ar affin a nd o the r s ub st ance s a nd will r e j e ct

, ,

sa nd w o o d gl a ss e t c
, ,
T h e s ub st a n ce s a cce pt e d a r e th o s e
,
.

whi ch a dh e r e t o th e d ro p o f chlo ro fo r m Si nce in th e m a j or ity .

Of ca s e s foo d p a r ti cl e s d o no t s ti ck t o th e s u r fa c e o f the
 AM E B A 49

Ame ba , w e c a nn o t e xpl a in ing e s ti o n by m e a ns of the s e physical

imit a tio n s .

Di ge s ti on . D ig e sti o n t a k e s pl a c e with o ut t h e a i d o f a s t o m a ch .

Aft e r a f oo d v a cu o l e h a s b e c o m e e mb e dd e d in t h e e nd o p la sm
i t s w a lls p o u r int o it a s e c r e ti o n o f s o m e m in e r a l a cid p r ob a bly
HC l .
“ It i s p rob a bl e th a t th e minute p a r ticl e s o f nucle opro
,

t e i d s th a t a r e co n s t a ntly a r i sing in t h e n e ighb or h oo d o f th e

nucl e u s co nt ain ce r t a in di ge stive f e r m e nts whi ch stimul a t e th e
fo r m a ti o n o f t h e mi n e r a l a cid in t h e vicinity o f th e g a st ric
v a cuol e ( 3 2 p ,
T h e dig e s tiv e flui d s e e m s t o di sso lve o nly
.

p r o t e id sub s t a nc e s h a ving no e ffe ct up o n fa t s a nd c a r b o hydr a t e s

, .

Th a t th e nucl e u s pl a ys a n imp o r t a nt r Ole in dig e stive p ro ce s s e s

wa s sh o wn by Ho f e r wh o f o und th a t if a w e ll f e d A m e ba wa s-

cut in t w o th e non nucl e a t e d p or ti o n w a s un abl e to dig e s t foo d

,
- .

Ege s ti o n . Indige s tibl e p a r ti cl e s a r e e ge s t e d a t a ny p o int

o n t h e s u r f a ce o f t h e A m e ba the r e b e ing no sp e ci al op e ning to
,

t h e e xt e ri o r f o r thi s w a s t e m a tt e r U su a lly s uch p ar ti cl e s a re

.

he avi e r th a n th e p ro t o pl a sm a nd a s th e a nim al m o ve s fo r wa rd
,

the y lag b e hi nd fin a lly p a ssing o u t a t th e e nd a w a y f ro m t h e di re c

,

ti o n o f mo ve m e nt ; th a t is Ame ba flo ws a w a y l e a ving th e indi

, ,

ge st i ble s o lid s b e hi nd .

As s i m ila ti on — T h e p e pt o n e s d e r ive d f ro m t h e di g e s ti o n o f
.
,

p ro t e id s ub s t a nc e s t o g e the r with th e w a t e r a nd min e ra l m a tt e r

,

t a ke n in wh e n the g a st ri c v a cu o l e w as fo r m e d a re ab so r b e d by ,

t h e s u r r o unding p r o t o pl a s m a nd p a ss int o th e b o dy s ub s t a nc e o f
t h e a nim al no ci r cul a t o r y s y s t e m b e ing p r e se nt s o f a r a s w e kn o w
,
.

Th e se p a r ti cl e s o f o r g a ni c a nd in o rg a ni c m a tt e r a r e th e n a ssimi
l a t e d ; th a t i s th e y a r e r e a rr a nge d t o fo r m ne w p a r ticl e s o f li v
,

ing p ro to pl a s m whi ch a r e d e p o sit e d a m o ng th e p r e vi o u sly e xi s ting

,

p a r ticl e s T h e a bility t o thu s m a nuf a ctu r e p ro t o pl a s m fro m

.

uno r g a niz e d m a tt e r it will b e r e m e mb e r e d i s o ne o f th e fund a

, ,

m e nt a l p ro p e r ti e s o f living s ub s t a nc e (p .

Di s s i m i l a ti on T h e e n e r gy f o r th e w o r k d o n e by Am e ba co m e s
.

f ro m th e b r e a ki ng d o wn o f co mpl e x m o l e cul e s by o xid a ti o n o r

“ p hy si ol o gi ca l bu r ning
”
Thi s i s kn o wn a s di ssi m i l a ti o n o r
.

E
5 0 AN INTRODU C T ION TO Z OOLOGY

k a tab oli sm T h e p ro duct s o f this sl ow co mbu s tio n ar e the e n e r gy

.

o f m o v e m e nt h e a t a nd r e sidu al m a tt e r Or din a rily this con

, ,
.

si s t s o f s o lids a nd fluids m a inly wa t e r so m e min e r a l subst ance s

, , ,

u re a and C0 Se cr e ti o n s e xcr e ti o n s and th e p r o duct s o f r e spi

2 .
, ,

ra ti o n are includ e d in t hi s list .

S e cr e ti on We h av e a l r e a dy n o t e d th a t an a cid is p o u re d
.

int o th e g a s t ri c v a cuo l e by th e su rro unding p r o topl as m Such a .

p ro du ct of di ssi m il a tio n whi ch is o f u se in th e e co n omy of the

,

a nim a l i s kn o wn as a s e c r e ti o n
,
.

Excr e ti on M a t e r i a l s r e p r e se nting th e fin a l r e ducti o n o f

.

sub s t a nce s in th e p r o c e ss of k a t ab olis m a r e call e d e xcre ti o n s .

Th e se a r e de p osite d e ithe r within or o ut side o f th e b o dy A l ar ge .

p a r t of th e e xcr e t ory m a tt e r includi ng u r e a and C 0 p a ss e s , 2,

through th e g e n e r al surf a ce o f th e b o dy T h e fluid co nt e nt o f .

th e c o nt ra ctil e v a cu ol e i s kno wn t o co nt a in u r e a the r e f or e t hi s ,

o rg a n i s e xc re t o r y in functi o n It i s a l so r e spi r a tory since C 0

.
, 2

p rob ably m a k e s i ts way to th e e xte ri or by w ay o f thi s o rg a n .

Oxyg e n di sso lve d in w a t e r i s t a k e n in th r o ugh th e su r f a c e o f th e

b o dy Thi s gas is n e ce ss a r y f o r t h e life of th e a ni m al ; if r e pl a c e d
.

by hyd rog e n m o ve m e nts ce ase a ft e r twe nty f o u r ho u r s ; if ai r i s

,
-

the n int ro duc e d m o ve me nts b e gin a g ain ; if no t d e a th e n sue s

, ,

G r o w th If fo o d i s pl e ntiful m or e s ub s ta n ce is a dde d to th e
.
,

living p r o topl as m o f th e Am e ba th a n i s u se d up in i ts v ar i o us
physic al a ctiviti e s T h e r e s ult i s an in cr e a s e in th e v olum e of
.

th e a nim a l This i s g r o wth and as in all o the r living or g ani s m s

.
, , ,

g rowth by th e a dditi o n of ne w p a r ticl e s a m ong th e p re e xisting

p ar ticl e s i e g r o wth by i ntu ss u sce pti on
,
. . .

R e pr o d u cti on Th e r e i s h o w e v e r a limit with re g ar d to th e

.
, ,

siz e th a t m a y b e a tt a in e d by A m e ba t it ly x d
pro e u s a s ra r e e ce e s ,

. 2 5 mm ( T ir . inch) in di a m e t e r Wh e n thi s limit i s r e a che d the

e .
,

a nim a l divid e s int o t w o p a r t s Why sh ould the re b e s u ch a .

limit T h e fo ll o wing e xpl a n a ti o n i s giv e n by He rb e r t Sp e nc e r

?

a nd o th e rs T h e v olum e o f a n o r g a ni s m v a r i e s a s th e cub e of
.

i ts di a m e t e r th e s u r fa c e a s th e s qu a r e
,
Thu s a s a n a nim al .
,

g rows th e ra ti o b e tw e e n su r f a ce and vo lum e de cr e as e s ; a nd since

, ,
5 2 AN INTR ODU CT I ON TO Z OOLOGY

( 1 ) Du ring nu cl e u s app a r e ntly divide s a mi

bi na r y d i vi s i on th e
t o ti cally a lth o ugh indi ca ti o n s o f mit os i s h a v e b e e n r e p o r t e d
, A .

de fini te mito tic figur e is fo r m e d in c e r tain sp e ci e s o f A m e ba ,

e g A m e ba bi nu cle a ta ( Fi g
. .
Whil e the nucl e us i s e l ong a ting
.

a c o n s t r i cti o n a pp e a r s a r o und th e mid dl e o f th e c e ll t h e nucl us

; e

th e n s e p a r a te s int o two app a r e ntly e qu al p ar ts whi ch m ove

towa r d th e e nd s o f th e a nim al M e a nwhi l e th e c o n s tr i cti o n o f .

th e b o dy h a s g r o wn d e e p e r a nd fin a lly ,

s e v e r s th e co nn e cti o n b e tw e e n th e t w o

e nd s a nd two d a ught e r Am e b a r e s ul t
‘

(4 7 Fi g
,

( 2) Sch e e l (4 6) se e ms to b e th e o nl y
p e rso n wh o h as e v e r witn e sse d sporu lati on
in Am e ba pr ote u s Th e whol e p r o c e s s .

l a st e d fr om tw o a nd o ne h alf to th re e
m o nth s T h e ps e ud op o di a w e re fi r st .

d rawn in and th e a nim al b e c a m e sphe r

ical A th re e l ay e re d cys t wa s th e n .
-

s e cr e te d T h e Ame ba r o t a t e d within thi s .

f o r s e v e r al d a y s a ft e r whi ch a ll m o ve ,

m e nt s c e ase d T h e nucl e u s divid e d until

d l“
FI G .
7 Am b
1 .bi n e a u .

ea b gi nni ng t di e
the re w e r e tw e nty o r thi r ty p re se nt
o ‘

,
V i d B th n‘d i h v a r ra nge d n e a r th e sur f ace
e O
f
e a e
C o ntinu e d
‘

f m d m t t fig
. .

or e i o ic ure s
d i v i si o n r e s ult e d I n a n I n cr e a se o f nu cle I
.

( F m L ngro ft a a er .

S h d i nn )
c au t o f r o m fi v e hund re d t o six hund r e d
.
.

T h e c e nt r al p ar t o f th e b o dy h a d
n on e C e ll w all s now app e a r e d a t th e p e r iph e ry cutting
.
,

o ff th e nucl e i e a ch with a s m a ll a m o unt o f th e s u rr o unding

,

cytopl asm T h e w all o f the cyst b e ca m e so ft b r o k e a nd

.
, ,

a ll o w e d th e sm a ll Am e b a to e s c ap e Hund r e ds o f th e s e .

Ame bu lae o r ps e u dopodi ospor e s a s th e y a r e so me tim e s ca ll e d

, , ,

b r o ke o u t a t o ne tim e Th e y b e cam e r e cogniz abl e a s Ame ba .

p r o te u s in f r o m t w o a n d a h a lf t o th r e e w e e k s N o r e a s on .

c o uld b e di sc o v e r e d f or sp o r ul a ti o n altho ugh e xp e rim e nts we re ,

c onduct e d in whi ch sp e cim e ns we r e s ta rv e d we re give n a n e xce ss ,

 AM E BA 53

of foo d we re all o we d to de sicc a t e a nd w e r e tra nsfe rre d to wa te r

, ,

f ro m diff e re nt l o c a lit ie s ; n o n e o f the s e re sulte d in e ncystm e nt and

sp o r ul a tio n .

T h e d e ve l o p me nt o f Ame ba is simply a m a tt e r o f g ro wth ;

b o th th e sp o re s a nd the da ughte r c e ll s r e s ulting f ro m bin ary di vi
s i o n b e c o m e f ull g r o wn sp e cim e ns by m e a ns o f a g r a du a l inc re a s e
-

in volume .

B e h avi or of Am e b a . T h e s u m t o t a l o f a ll th e v a r i o us m o ve
m e nts of an a nim al co n s titute wha t i s kn o wn a s its b e h avi o r .

In Ame ba th e se mo ve me nts m a v b e s e p a ra t e d into tho s e co n

ne cte d with l o c o m o ti o n a nd th o se r e s ulting f r o m e xt e r n a l stimuli .

We h av e a l re a dy give n an a cc o unt of th e l o c o m o tio n o f Ame ba ,

a nd so sh a ll c o nfin e o u rse lv e s now to a discussi o n o f its r e sp o n se s

t o di ffe re nt kind s o f s timuli . T h e re a cti o ns Of A me ba to s timuli

ha ve b e e n g ro up e d by J e nnings ( 3 9 4 1 ) int o p ositiv e n e g a tive
, , ,

a nd fo o d -t a king . T h e l a s t n a m e d w e r e di s cuss e d o n p a ge 4 7 .

The fo ll o wing a cco unt the n will d e a l with p ositive a nd n e g a tive

, ,

r e a cti o ns o f Ame ba t xt n l stimul ti n

p r o e u s t o e e r a a o .

De fi ni ti ons of T e rm s . F i r s t it wil l b e w e ll t o d e fin e a f e w of
,

th e t e r m s u se d in d e sc r ibing th e m o t or r e sp o ns e s o f a ni m a ls to
s ti muli A r e a cti o n r e s ulting f r o m a ch a nge in a n a nim a l s ’
.

e nvi ro nm e nt ,
e g a n inc r e a s e in th e int e nsity o f th e light
. . is
,
”
kn o wn as t ropi s m o r t a xi s . Th e te r m t ropis m m e a ns
to tu r n it i s us e d inst e a d o f s a ying th a t an a nim a l lik e s
or di slike s c e r t ain stimuli b e c a use w e ca n de te r min e by e x
,

i m n t whi ch w a y a n individu a l wi ll tu r n und e r v a ri o us c o ndi

p
“
e r e

ti o n s but are no t in a p o siti o n to d e cide whe the r like s o r

,

di sl i ke s e nte r into th e phe n o m e n a T h e kind o f s timulus

.

e mpl o y e d is indic a t e d by a p r e fi x . T h e p r incip al kinds o f tro

p sm s are a s fo ll o ws :
i
( 1 ) Thigmo t ropi sm r e a cti o n to c o nta ct .

( 2) C he mo t r opi s m re a cti o n to a ch e mic al .

( 3 ) The r m o t ropi sm r e a cti o n t o h e a t .

(4) P ho to t ropis m r e a cti o n t o light .

5) El e ctro t r opi s m re ac ti o n t o e l e c t r ic c u rre nt .

54 AN INT R ODU CTION TO Z OOLOGY

( 6) Ge o t ropi s m cti o n to g ra vity

re a .

( 7) C hr o m o t ropi s m r e a cti o n to c o l o r .

( )
8 R he o t ropism r e a cti o n to cu r re nt .

Ta xi s i s Ofte n e mpl oy e d in s te a d o f tro pi s m whe n the te r ms

” ”
re a d thigm o ta xis che m o ta xi s e tc If the a n im al re a cts
, ,
.

by a m o v e m e nt to wa r d th e s ti m
ulu s s uch as light it i s sa id to
, ,

b e p ositive ly ph o t o t r opi c ph o t o ,

t a ctic e tc ; if aw a y fr o m th e
,
.

stimulu s n e g a tiv e ly ph o to t ropic

,

o r ph o t o t a ctic e tc Am e ba h as ,
.

81 Ame ba
. ti m l ts d b
u a e e e n f o und t o re s p o nd t o c o nt a c t
g l ass d ro . with so lids t o ch e mi c a l s t o h e a t , , ,

to light to co l ors a nd to e l e c, ,

t ri ci ty Aft e r d e s c r ibing the re

.

a cti o ns o f A m e ba to e a ch o f th e

a b o v e n a m e d s timuli th e g e n e r a l
-
,

signifi ca nc e o f its b e h avi or will b e

co n side r e d .

Thi n tropi s m — I f a m o ving

FI G Am b m v lw ys °

.
91 . e a o es a a

f m m h ni l
ro ec a tim l
ca s
A m e ba IS t o uch e d a t a ny p o int
u u s.

( F mro J n n i
e ng ) s. with a so li d O bj e ct e g a gl as s ,
. .

ro d th e p a r t a ff e ct e d co nt racts
,

a nd m o v e s a w a y If the a nte rior e dge is s timula te d in this w ay

.
,

t h e p a r t t o uch e d s t ops a nd c o nt ra ct s whil e a ne w p se udo p o dium

i s pu sh e d o u t in s o m e o th e r pl a ce a nd th e a nim a l m o ve s a w a y in
a n o th e r d i r e cti o n ( Fi g This is cl e a rly a n e g a tive re a ctio n
.
,

a nd t h e a ni m a l i s sa id t o b e n e g a tiv e ly thigm o t ro pic The r e .

are ,
ho we ve r c e r ta in co nditio n s which c all fo r th p ositiv e r e
,

sp o nse s t o c o nt act with s o lid s F or e x a mpl e if th e Am e ba i s

.
,

fl oa ti ng fre e ly in th e w a te r and a p se udop odi um c o me s in c o nta ct

with th e s ub s t ra tum ( Fi g 20) th e a nim al m o v e s in th e di re ctio n
.

o f th e p se ud o p o dium s timul a t e d until th e n or m a l cr e e ping p o s i

tio n h as b e e n a tta in e d C o nt a ct with foo d al s o r e sult s in p osi

.

tive re a cti o n s as de s cr ib e d o n p ag e s 4 7 a nd 4 8 Ame ba the re

, .
,
 AM EBA

fore r e acts n e ga tive ly to a strong m e chani c al stimul us and p osi

,

ti v e ly t o a w e a k o ne .

C h e m o tr opi s m Am e ba is se n sitiv e to ch a nge s in th e ch e mi

.

cal c o mp o siti o n o f th e w a t e r sur r o undi ng it If a ch e mica l is .

FI G . 20 . M e th od by whi ch a floa ting Ame ba pas se s to a so li d .

( From Je nni ngs ) .

bro ught into co nt a ct with o ne sid e by m e ans of a fi ne c apill ary

gl ass r od or tub e th e p ar t a ff e ct e d c o nt ra cts a nd th e a nim al
,

m ove s off in so m e o the r di r e cti o n ( Fi g It h as b e e n sh o wn .

t o r e a ct n e g ativ e ly wh e n th e f oll o wing

sub s ta nc e s co m e in c o nt a ct with one side
o f i t s b ody : m e thyl e n e blu e m e thyl g r e e n , ,

so dium chl o rid e so dium ca r b o n a t e o t a s

, p ,

sium nit ra te p o t assium hyd ro xid e a c e tic

“
, ,

a cid hyd r o chl o ric a cid c a n e s uga r d i s FIG N g tiv 21 . e a e {e a c

, , ; ,
nt h m l nm o o C e l ca s u
till e d w ate r tap w a te r a nd w a te r f ro m l ti n i n Am b A
, ,
a o e a .

o th e r cultu r e s th a n th a t in Wt h th e littl m thyl g n d if e e re e

”
Amoeba und e r e xp e r im e nt a ti o n liv e s f g i n t th d u se s a a s e a

(4 1,
p .
v n i ng n d i ng a c e , ca u s

Th e r m otr opi s m n w d di t b
Am e ba giv e s a n e g a
.
P P e se u O O a o e

th t t nd d ‘ ng rus ou a a a e
tive re sp o n se if l o cally a ffe ct e d by h e a t i n th d i ti n f m v
.

e re c
’

o o o e
Thi s m ay b e a cco mpl i she d by b ri ngi ng m nt ( F m J nni ng ) e . ro e s .

a he a te d n e e dle in f r o nt o f an Ame ba
w hi ch i s c r e e ping al o ng o n th e und e r su r f a c e o f a c o v e r gl a s s .

Whe n th is i s do n e th e a nim al m ov e s o ff in a n o the r di r e ctio n

,
.

T he d e g re e o f t e mp e r a tu r e a l so h a s a n influ e nc e up o n th e r a t e
o f th e n o r m al m o v e m e nt s C o ld a nd e x ce ssiv e h e a t r e t a r d its
.

a ctiviti e s whil e a m o d e r a t e a m o unt o f h e a t inc r e a se s t h e m o v e

,
5 6 AN I N T R ODU C T ON T O I Z OOLOGY

m e nt Activity ce ase s
.

v i i a lt o g e th e r b e tw e e n
30
°

°
a nd 35 C .
,
a nd th e ani
kill e d by coagul a ti o n of th e
m al i s
p r o topl a sm if th e t e mp e r a tu r e i s
ra i s e d t o 4 0
°
C .

P h ototr opi s m If a s trong light .

str ik e s Am e ba fr o m th e side it will ,

o r i e nt it se lf in th e di r e c ti o n o f th e

r a ys a nd m o v e a w a y f ro m th e so u r ce

o f illumin a ti o n ( Fi g Or din a ry whit e light .

c a u se s a n Ame ba to stop m o ving

C h r o m o tr opi s m — T h e e ff e ct o f blu e light up o n
.

Am e ba i s sim i l ar t o th a t o f o r din a r y whi t e light .

R e d light a pp a r e ntly h as no e ff e ct wh a t e v e r up o n
i ts n or m a l a ctiviti e s whi l e o th e r co l o r s c a u se a v a r i ,

a bl e a m o unt o f d e c r e ase in i ts m o v e m e nt .

E l e ctr o tr opi s m If a n e l e ct ri c cu r re nt i s p a s se d
.

thr o ugh a n Am e ba it c o nt ra ct s o n th e p o sitive o r ,

a nod e side o f th e b o dy a nd m o v e s t o wa r d the

c a th o d e or n e g a tiv e p o l e .

In Am e ba th e r e ar e no org a ns th a t ca n b e com
p a re d with wh a t w e c all se n se o rga n s in highe r org an
i sm s and w e mu s t a ttr ibut e its r e a cti o n s to stimuli
,

to th a t fund a m e nt a l p ro p e r ty Of p ro t o pl a s m c a ll e d
i rr ita bility T he sup e r fici al l aye r Of .

x cy toPl as m r e ce iv e s th e s timulu s a nd
tr an s fe r s th e e ffe cts t o so m e o the r p a r t
o f th e b o dy ; thu s m ay b e s h o wn th e

ph e no m e n o n o f int e r nal
F R t n f Am b t l gh t T h
Tr r l t a b m
IG 22 e ac i o o e a o i e
ty o r co nd u cti v
. . .

w i nd i t t h d i ti n f th ligh t
a r ro s ca e e re c o o e

ra y nd t h n m b
s a e th u iv p i
e rs
I t)”
e
P e
s u cce s s
r h a p s th e m os t
e os

ti no m d b y th
s a ss u e ni m l Th A m be a p ri miti v e m e tho d I n th e
a . e e a

lw y m v w y f m th f a nim a l kingd o m o f a r e

_
a a s o es a a ro e s o ur ce o

light ( F m J nni ng f t D v np t )
. ro e s a
a cti o n t o s timuli is
er a e
i11 ,s
or .

,
 AM E B A
57

t ra t e d by sp o n se o f a d e finite p oint o n th e b o dy o f
th e re

Ame ba to a st ro ng m e ch a nic a l s timulu s Thi s m ay be c o mp a r e d

.

with wh a t h app e n s wh e n a m a ss o f in o rg a nic m a t e ri al i s act e d

upo n by a for c e ; h e r e a r e a ctio n i s p ro duce d dir e ctly a t th e
p o int o f co nta ct In o ri e nting itse lf in th e di r e ctio n o f th e light
.

r a y s A me ba sh o w s a r e s p o n se t o a co ntinu o u s s timulu s w hi ch
,

a ff e c t s th e e nti r e b o dy ( F i g if t h e s o u r c e of illumin a ti o n
.

i s ch a nge d th e a nim a l ch a ng e s i ts o r i e nt a tio n

,
T he ca u se o f .

th e r e a cti o n a s in th e a b o v e in s t a n ce
,
s e e m s to b e in m os t
,

case s a ch a ng e in th e e nvi r o nm e nt T h e r e sp o n se i s a lw a y s p r o
.

d u ce d by a stimul a ti o n di r e ctly p r e c e di ng it but t h e a nim a l s oo n ,

b e co m e s a cclim a te d to th e ne w e nvi ro nm e nt a nd i ts a ctiviti e s ,

b e co me n orm al Thi s is sho wn wh e n it i s t r a n s fe rr e d f ro m o ne

.

cultu re to a n o th e r (Ch e m o t ropism) or wh e n whi te light i s thro wn

up o n it (ph o to t ropi s m) .

T h e re a cti o n s o f A m e ba t o stimuli a r e o f und o ubt e d v a lu e to

th e individu a l a nd to th e p re s e r v a ti o n o f th e r a c e f o r in e v e r y ,

in s ta nce th e n e g a tiv e r e a cti o n i s p r o du ce d by inj ur i o us ag e nts

such as s tr o ng ch e mi cal s h e a t a nd m e ch a nic a l imp a cts whi l e
, , ,

p ositiv e r e a cti o n s a r e p r o duce d by b e n e fi ci al a ge nt s T h e r e .

spo nse s th e r e f o r e
,
in th e fo r m e r ca se s ca rr y th e a nim a l o u t o f
,

d a nge r in th e l a tt e r to w a r d a sa fe r h a ve n
, ,
.

A me ba i s o f fund a m e nt a l inte r e s t t o a nim al p sy ch ol o gi s ts ,

sin ce it r e p r e se nt s the a nim al mind in i ts m os t p ri m itive

f o r m T he b e h a vior o f Ame ba in th e a bs e nce o f e xte r n a l s tim uli
.
,

f o r e x a mpl e wh e n it i s s u sp e nd e d fr e e ly in th e w a t e r sh o w s th a t ,

s o me o f i ts a ctiviti e s a r e initi a te d by inte r n a l c a u se s Whe the r .

o r no t th e a nim a l i s in a n d e g r e e c o n s ci o u s i s a qu e s ti o n s till
y
un a n swe r e d If Am e ba h a s r e co gniz abl e se n sa ti o n s th e y mu s t
.
,

b e infi nit e ly l e ss in b o th qu a lity a nd qu a ntity th a n in high e r

o r g a ni s m s F u r th e r m o r e it i s un abl e t o l e a r n f r o m th e f e w kinds
.
,

o f e xp e r i e nc e s it do e s p ass th r o ugh a nd i s th e r e f o r e l a cking in

,

m e m ory im age s One ca se in whi ch a n Am e ba a tt e mpte d t o

.

capture a n o the r i s re co r d e d by Je nning s a nd s e e m s t o p o int t o ,

s o m e thing hi ghe r th a n m e re phy sica l a nd ch e mic al a tt r acti o ns .

5 8 AN IN T R OD U CTION TO Z OOLOGY

In this in s t a nc e a l ar g e a nim al Sh o we d de cide d p e rsi s t e nc e in its

e nd e a v o r s t o e ng u lf a s m a ll e r A m e ba ; se v e ra l tim e s it r e v e r se d

i ts c o u r s e a nd co ntinu e d th e pu rs uit .

A r e vi e w o f th e f a cts thu s f a r Obt a in e d se e ms to s h o w th a t

f act o r s a r e p r e se nt in th e b e h a vi o r o f Am e ba co mp a r a bl e to th e
h a bit s re fl e x e s a nd a uto m a ti c a ctiviti e s of highe r o r g a ni s m s
, ,
”

(3 9 ,
p . a nd if Am oe b a w e r e a l a r g e a nim a l
,
so a s t o c o m e

within th e e v e r yd a y e xp e r i e n ce o f hum a n b e ing s i t s b e h avi o r

,

w o uld a t o nce c all f or th th e a ttr ibuti o n t o i t o f s ta te s o f pl e a s ur e

a nd p a in o f hung e r d e si r e
, , ,
a nd th e lik e ,
o n p r e ci s e ly th e s a m e
”
b a si s a s w e a ttr ibut e the se things t o th e d og (4 1 p ,
.
60 AN INTRODU CTION TO Z OOLOGY

o utside o f th e e ctosar c,
Lying in the e ct opl asm
are a g re a t numb e r O f
minute sacs th e tr i chocysts
whi ch di scha rg e l ong
th re a ds to th e e xte ri or
wh e n p rop e r ly s timul a t e d .

One l ar g e contr acti le va cu ole

i s s itu a t e d n e ar e ith e r e nd
o f t h e b o dy ,
cl ose to the
d or sa l s ur f a ce whi l e a
,

v a r i abl e numb e r of food

vacu ole s m ay usu a lly
be s e e n . T h e nu cle i a r e
a - b ‘ t w o in numb e r a l a rg e ,

macr onu cle u s ( m a .

) a nd a

sm a ll e r mi cr onu cle u s (m i ) ;
the se ar e su sp e nde d in the
e c t o pl as m n e a r th e m o uth

o p e ning . Th e a na l s pot
( )
a n c
. a n b e o b s e r v e d o nl y
wh e n solid p ar ticl e s a r e di s
ch a rge d . I t i s situ a t e d
ju s t b e hind the p o st e ri or
e nd o f th e o ra l g r o o v e .

FI G . 23 . vi e w e d f rom th e o ra
P a r a me ci u m l D e taile d Ana to my .

s u r f a ce L l e f t s i d e ; R r i ght s i d e
T h e e nd opla sm ( Fi g 2 3
. .
, ,

e ct o sa r c ;
,
.

an . a nu s ;
,
ec en e nd o
.
, .
,

sa r c ; f v f o o d v a cu o l e s ; g gu ll e t ;
. .
,
e n ) o f P a r a m e
.
ci
,
u m o ccu .

m m o u th ; m a m a cro nu cl e u s ; m i
.
,
pi e s th e ce ntral p ar t of
.
, .
,

m i cr o nu cl e u s ; o g o r a l gr o o ve ; p t h e b o dy
. .
, It i s supp o se d .
, .

pe lli cl e ; tr t r i cho cys t l aye r T h e t o b e a lv e ol ar in s t r uctu r e

.
, .

a rr o w s S h o w t h e d i r e cti o n o f m o ve
M os t o f th e l ar g e r g r a nul e s
m e nt o f th e f oo d v a cuo l e s ( Fr o m
c o nt a in e d ithin it
.

Je nni ngs ) .
w are

h own by mi croch e mical

s

re a cti o ns to b e r e s e r v e f oo d p a r ti cl e s ; th e y flo w f r o m pl a c e t o
 P AR AM EC IUM 6I

pl a c e indic a ting th a t th e p r o t opl asm i s o f a fluid n a tu re T he

,
.

e ctoplas m ( e c ) d o e s no t co nt a in a ny o f t h e l a r g e g r a nule s ch a r
.

a ct e r i s t i c o f t h e e nd opl a s m s inc e i t s d e n sity p re v e nts th e i r e n

,

t ra nce In thi s r e sp e ct th e two kinds o f cyt opl a sm r e s e mbl e

.

t h e e ct opl a sm a nd e ndopl a s m o f A me ba Outs id e of t h e e cto

.

pl a sm i s a de li ca t e e l a s tic m e mb r a n e th e pe lli cle (p ) o r cu ti cle

, . .

If a d rop o r tw o o f 3 5 pe r c e nt a l co hol i s a dd e d t o a d r op o f
w a te r co nta ini ng P a r a me ci a t h e p e lli cl e will b e r a ise d in so m e
,

sp e cim e n s in th e fo r m o f a bliste r Unde r th e hi ghe r p o we rs o f

.

th e mic r os c op e t h e p e llicl e i s th e n s e e n t o b e m a d e up o f a g r e a t
numb e r o f he x ag o n a l a r e a s p ro du ce d by s tri a ti ons o n th e s u rf ace
( Fi g 24 A) Th e se st r i a ti o n s a r e r e a lly v e r y fi ne g roo ve s ( Fi g
.
,
. .

24 B ) which c r o s s o ne a n o th e r obliqu e ly
, , .

T h e di s t r ibuti o n o f th e m o til e o r g a n s th e ci li a c or r e sp o nds

, ,

to t he a r r a ng e m e nt o f t h e s t r i a ti o n s o n th e cuticl e sinc e o ne ,

cilium p roj e cts fro m th e c e nt e r of e a ch h e x ag o n a l a re a ( Fi g 24 .

,

A) . The s e t hr e a d lik e str uctu r e s o ccu r o n all p a r ts o f th e b o dy

-
,

tho se a t the p os t e ri or e nd b e ing slightly l o ng e r th a n e l se wh e re

( Fi g . A cilium m a y b e c o mp a r e d to a v e r y fi ne ps e ud o
p o dium whi ch h as b e co m e a p e r m a ne nt str uctu r e It i s a n ou t .

g ro wth o f t h e c e ll p r o topl a sm c o ming f ro m a b asa l b o dy ca ll e d a

,

mi cro s o m e ( Fi g 24 B B ) which app e ar s t o a r i se f ro m th e nucl e us

.
, ,
.

T h e st r uctu r e a nd m ov e m e nt s o f cili a h a v e b e e n s tudi e d by i nv e s

ti ga t o r s f or m a ny y e a r s a nd th e f o ll o wing th e o r i e s h a v e b e e n
,

p rop o se d t o e xpl ain th e m : ( 1 ) C ili a ar e s o m e wh a t s ti ff life l e ss ,

p ro c e sse s a tt ache d to the c e ll a nd m o ve d by a ctive e l e m e nt s

withi n th e c e ll b o dy ( )
2 C ili a a r e h o ll o w e l a s tic s h e a th s

into which a fluid i s inj e cte d a nd withd r a wn 3) C ili

. a c o n s i s t o f
a co mpl e x fluid o wing i t s c o nt r a ctility t o th e p r e se n ce o f a
fil a m e nt o f kine tic g r a nul e s pl a ce d a l o ng o ne e dge o f th e cilium
, ,

th e co nt r a cti o n o f thi s th r e a d fu r ni shi ng th e p o w e r o f t h e cilium

(52 p , .
(4 ) C il i a a r e fib r illa r t h e m o v
,
e m e nts b e ing d u e t o

t h e c o nt r a cti o n of t h e fib r il s We a r e inclin e d to fa vo r th e l a st
.

the o ry since th e mo ve m e nts ca n o nly b e a d e qu a te ly e xpl aine d

,

by the p re s e nc e o f a c o nt r a ctil e fr a me wo rk Fib ril s have b e e n .

62 AN INTRODU CTION TO Z OOLOGY

d e m o n s tr a t e d in th e flage lla o f se v e r a l uni ce llul ar a nim al s a ,

fl age llum b e ing a l a r g e cilium F or e x a mpl e in a P r o to zoo n .

,

Eu gle na ( se e p th e fl a g e llum co n si s t s Of f o u r fib r il s w hi ch
.

e xt e nd i t s e nti re l e ngth Th e se a r e twi s t e d a b o ut o ne a no the r

.

in a spi ra l o f tw o a nd o ne h a lf tu r n s Th e y ca n b e t r a ce d int o .

th e a nim al wh e r e th e y b r a n ch o u t int o a s y s t e m o f r o o tl e ts
,

( g
F i . In t h e s m a ll e r cili a th e s e fib r il s h a v e no t b e e n s e en ,

FI G . 25 . St r u ct ur e of th e fl age ll um of E u gle na .

( Fr o m D e lli nge r i n J ou r n M or ph ) . .

but ar e p rob ably p r e s e nt ; a t l e as t w e ca n a cco unt f or all th e ,

m o v e m e nt s m a d e by cili a if s uch a co n di ti o n e xi s t s a s in th e flage l

lum o f Eu gle na A fu sio n o f cilia f r e qu e ntly t a k e s pl a ce f or ming
.
,

m e m br a ne lle s In P a r a m e ci u m thi s h a s o ccu rr e d withi n th e

.

m o uth cavity p r o duci ng th e u nd u la ti ng m e mbr a ne ( F i g 2 9 M b)

,
.
,
.

Thi s i s a tta ch e d to th e d o r sal w all o f th e m o uth a nd guide s th e ,

f oo d p a r ti cl e s th a t a r e s w e pt within i t s r e a ch .

Jus t b e n e a th th e cili a e mb e dd e d in th e c o r tic a l l a ye r o f th e

,

e ct opl as m i s a unif o r m l a y e r o f spindl e sh a p e d s t r u ctu r e s T 3 3 5

4 -
,

mm in l e ngth lying with th e i r l o ng a x e s p e rp e ndi cul ar to th e

.
,

s u r f ace ( F i g 2 4 A tr
. Fi g 2 4 B T)
, ,
.
,
Th e se a r e tri chocysts
. . .
, ,

The i r di s t ributi o n i s sho wn in s u r fa ce vi e w in Figu r e 24 A The y ,

.

a pp e a r t o b e c a viti e s in t h e e ct o pl a s m fill e d with a se mi liquid -

h o m oge n e o u s sub s t a n ce whi ch i s v e ry r e f r a ctiv e Th e y a rise .

in th e n e ighb o rh oo d o f th e nucl e u s a nd p r ob ably f ro m it ,

A sm a ll a m o unt o f o s mi c o r a ce tic a cid wh e n a dd e d t o a d rop o f ,

w a te r co n ta ining P a r a m e ci a ca u se s in so m e c a se s th e discha rge

,

o f th e t ri ch ocy sts to th e e xt e r i or th ro ugh v e r y sm all ca n al s .

Thi s e xp l osi o n i s d u e t o th e p r e ss ur e d e r ive d f ro m th e co nt ra e

tio n Of the co r tica l l a ye r o f th e e ctopl asm Afte r th e e xpl osi o n .
,
FI G . 24 . Par ts of h o wing cili a a nd t r i ch o cys t s A Su r fa ce vi e w
P a r a me ci a s .
,

S h o wi ng s t r i a ti o ns cili a a nd t r i ch o cys t s ( tr )
, ,
. B p a r t o f a cro ss s e cti o n
,

s h o wi ng r i d ge s ( L) o n t h e s u r f a ce ,
cili a ( C ) co m i ng f r o m m i cro so m e s

( )
B t r i ch t
o cys s c t
u l n
o g it u d i na ll y ( ) y g
T l i n i n t h e e c t o s a r c ( C o),
a n d
,

t r i cho cys t s ( T) a nd a f o od v a cu o l e ( N ) l yi ng i n th e e nd o p l a sm (E n) .

,
t i h t v
C r c o cys s i n a r o u s s gi t a e s o f e x t r u s i o n T k,
b o d y o f t h e s t r u c t ure ;

Tf th e,
h i li j ti
a r k e p ro e c o n ( A f
. r o m S c h u b e rg; B ,
C f ro m M a i e r i n
A r ch i v f . P r o ti s t .
)
 P ARAM E C IUM 63

th e tr ichocysts app e a r a s l o ng thre a ds whi ch ha v e b e e n e xt e nd e d

t o ab o ut e ight tim e s th e i r f or m e r l e ngth ( F i g 2 4 C ) Tr icho
.
,
.

cysts a re supp ose d to functi o n as we ap o ns of off e ns e a nd d e f e ns e .

It is s a id th a t the i r co nt e nts a r e disch arg e d with c o n side rabl e

for ce and tha t the y c o nt ain a p o i so n s t ro ng e n o ugh to p ara lyz e
a ny s i ngl e c e ll e d a n
- i m al T h e o nly e vid e n ce w e h a v e th a t th e
.

t richo cy s ts a re w e ap o n s o f de f e n se i s fu r ni sh e d whe n P ar a me ci u m
e nc o unt e r s i t s e n e my Didi ni u m If th e s e izing o r g a n o f thi s
.

P r o t o z o o n b e co m e s f a s t e n e d in th e P a r a me ci u m a g r e a t numb e r ,

o f t r ich o cysts n e a r th e pl ace o f th e inju r y a r e disch a r g e d ( F i g .

The s e p ro duce a subst a nce whi ch b e

co m e s j e lly lik e o n e nt e r ing th e w a t e r ;
-

thi s t e nds to fo r c e th e tw o a nim al s ap a r t ,

a nd if th e P a r a m e ci u m
,

is a l arg e o ne f r e qu e ntly ,

it s ucce e ds in m a king
its e sc ap e

FI G . 26 . P a r a me ci u m d e f e nd i ng it se l f f ro m a n a tt a ck by a P ro t ozoo n
Did i ni u m T h e t r i ch o cys t s a r e d i s ch a rge d a nd m e ch a ni ca lly f o rce th e
.

e ne m y a w a
( Fro m M a s t i n B i ol B u t )
’

y . . .

T wo contr acti le p r e s e nt o ccupying de finite p o si

va cu ole s a r e ,

ti on s one n e a r e ithe r e nd o f th e b o dy The y li e b e twe e n th e

, .

e ct opl a s m a nd th e e nd o pl a s m cl ose t o th e d o r s a l s u r f a c e a nd co m ,
,

m u ni ca te with a l ar ge p o r ti o n o f th e b o dy by m e a n s o f a sy s t e m
o f r ad i a ti n ca nals s i x t o t e n in numb e r T h e v a cu o l e s gr o w in
g ,
.

siz e by th e a dditi o n o f liquid w hi ch i s e x cr e t e d by t h e p r o t o pl a s m

into th e c a n als a nd i s the n p o u re d into th e m Whe n th e full .

64 AN INT R ODU C TION TO Z OOLOGY

siz e is ch e d th e w all s c o nt r a ct a nd th e c o nt e nt s are disch ar g e d

re a ,

t o th e e xt e r i o r p r o b a bly th r o ugh a p o r e
,
T h e t w o v a cu o l e s d o .

no t co nt r a ct a t t h e sa m e tim e but a lt e r n a t e ly t h e int e r v a l b e , ,

tw e e n s ucc e ssiv e c o nt r a cti o n s b e ing te n t o twe nty se co nd s T h e .

e xpul s i o n o f th e flui d c o nt e nts o f th e co nt r a ctil e v a cu o l e s m a b e

y
s e e n in th e f ol
l0wing way
P a r a m e ci a Sh o uld
be m o unte d in
w a t e r int o whi ch
h as b e e n r ubb e d
up a s ti ck of In di a
or C hi n e se ink .

The y th e n app e ar
whit e ag ain s t a
b l a ck b a ck
g r o und P a r t o f .

th e w a t e r s h ou ld
be withdrawn
F G
I .
7 P
2 . m i wi m m i ng i n
ar a ec a l ti n f I nd i
s
f r o m b e n e a
a so u
th t h
o
e o a

i nk h wi ng th d i h g f t h
,
s o e nt til c ov e r gl a ss thu s
sc ar e o e co ra c e ,

v l t th
a cuo e s t id o ( F m D hl g n nd slightly c o m p r e ss
e ou s e . ro a re a

K pn f t J nni ng )
e er a er e s .

ing th e m If no w .

a sp e cim e n in p r o fi l e i s f o und a nd w a t ch e d th e di s ch a r g e pr o ,

duce s a b r ight sp o t o ut sid e in th e o p a qu e liquid ; thi s l a s t s f ro m

o ne t o t w o se co nd s a nd i s th e n d r iv e n Off by th e cili a ( 6 0 F i g
, ,
.

Wh a t h a s b e e n sa id o f th e fun cti o n o f th e c o ntra ctil e .

v a cu o l e in Am e ba (p 3 9) appli e s a s we ll t o th ose o f P a r a me
.

ci u m i e it a ct s a s a n o r g a n o f e x cr e ti o n a nd r e spi r a ti o n a nd i s
,
. . ,

p r ob ably hyd ros t a ti c

Lo com o t oni — T h e o nly m o ve m e nt s o f P a r a m e ci u m th a t in
.

n
a yw y a r e s e mbl e th o s e o f A m e b a a r e se e n wh e n t h e a nim a l p a s s e s

th ro ugh a sp a ce s m all e r th an i ts sh or t e r di a m e te r ; it wi ll the n

e xhibit an e as cl ti ity w h i ch a ll o w s it t o s qui r m th r o ugh In a .

f r e e fi e ld P a ra me ci u m s wim s by m e a n s o f i ts ci li a The se a r e .
66 AN INTRODU CTION TO Z OOLOGY

o ra l side i s to th e l e ft i s to the r ight ; wh e n th e o r al side i s ab ove

, ,

t h e b o dy s w e r v e s d o wnw a r d ; wh e n th e o r a l sid e i s to th e r ight ,

th e b o dy sw e r v e s to th e l e ft e tc ,
He nc e th e s we r ving in a ny
.

give n di r e cti o n is co m p e n sa te d by an e qu al sw e r ving i n th e

o pp osit e di r e cti o n ; t h e r e s ult a nt i s a spi r a l p a th h a ving a s t r a ight

a xi s ( 6,
2 p 44
.
; F i g .

R o t a ti o n i s thu s e ffe ctive in e n abling a n un s ymm e t r ic a l a nim al

t o swim in a st r a ight c o u rs e th r o ugh a m e di um which all o ws
de vi a tio n s to r ight o r l e ft a nd up or do wn It i s we ll kn o wn th a t
,
.

a hum a n b e ing ca nn o t k e e p a st ra ight c o u rse wh en l os t in th e

w oo ds alth ough h e h as a ch anc e to e rr o nly t o th e r ight o r l e ft

, .

N u tr i ti on. T h e f ood o f P a r a me ci u m c on s ist s p r in cip a lly o f

b acte r i a a nd minut e P r o toz oa T h e a ni m al d oe s not wait f o r
.

th e f oo d to c o m e withi n i ts r e a ch but by c o ntinu a lly s wim m ing

,

f r o m pl ace to pl a ce i s abl e to e nt e r r e gi o n s whe r e fa vo r abl e fo o d

c o n di ti o ns p r e v ail T h e cili a al so ai d in b ringing in foo d p a r ticl e s
.
,

S inc e a s o r t o f v o r t e x i s f or m e d by th e i r a rr a ng e m e nt a b o ut t h e

o r a l g r oo v e whi ch di r e ct s a s t e a dy st r e a m o f w a t e r t o w ar d th e

m o uth .

F igu r e 2 9 illu s t ra t e s th e fo r m a t io n of a food va cu ole ( N e ) .

F oo d p a r ticl e s th a t a r e s w e pt int o th e m o uth ( M u ) a r e ca r r i e d

d o wn into th e cyt oph a r ynx (s) by th e und ul a ting m e mb ra ne
( M b) ; th e y a r e th e n m o ve d o nwa r d by th e cili a lini ng th e cyt o
ph arynx a nd a re fin ally g a th e r e d t og e the r a t th e e nd o f th e
p as sag e w a y int o a v acu ol e whi ch g r a du ally fo r m s in the e ndo
pl a sm ( N e ) Wh e n thi s v a cuo l e h a s r e a ch e d a ce r ta in size it i s
.
,

pin ch e d off fr o m th e e xt r e mity o f th e cytopha rynx by a co nt ra e

tio n o f th e su r ro unding p r o topl asm a nd th e fo r m a tio n o f a n o the r
,

v a cu ol e i s b e gun A f o o d v a cu ol e ( N ) i s a d ropl e t o f w a te r with

.

f oo d p ar ticl e s su sp e nd e d within it As s o o n a s o ne i s se p a r a te d
.

f ro m the cytoph a rynx it i s swe pt a wa y by th e ro t ar y str e a ming

,

m ove m e nt o f th e e ndopl a s m kn o wn as cyclosi s Thi s c arri e s .

th e f oo d v a cu ol e ar o und a d e finit e co u rs e whi ch b e gin s just ab ov e

a nd b e hind t h e cyt o ph a rynx p a s se s b ackw a r d t o th e p o s t e r i or
,

e nd th e n f o r w a r d n e a r th e d o r s a l su r f a c e t o th e a nt e r i or e nd
, ,
FI G . 29 . A se c tio n
f ro m t h e r e gi o n o f t h e m o u th o f P ara me ci u m sh o wing
th e f or m a ti o n o f a f o o d v a cu o l e . E n , e nd o sa r c ; M a , m a cr o nu cl e u s ;
. .

Mb , u nd u l a ti ng m e m b r a ne M u , m o u th N f oo d va cu o l e N e , f ood
. . .

va cuol e f orm ing ; S gulle t ( Fr o m M a i e r i n Ar chi v f P r oti st )

. . .

FI G 3 0 . B i nary fi s s i o n o f P a r a me ci u m a u r e li a
. . 1 , 3 , 4 , 5 , co nt r a ctil e v a e n

o le s ; 2 , 6 d ivi d i ng m a cr o nu cl e u s ;
, 7, 9, g u ll e t ; 8 ,
1 0 , d ivi d i ng m i cr o

nuc e li
( Fr o m La ng s Le h r b uch )
.
’
.
 P ARAM EC I UM 67

a nd fin ally downw ar d a nd al o ng th e ve ntr al su r fa ce t o war d th e

m o uth (indi c a t e d by a r r o ws in F i g 2 D u r ing thi s j o u r n e y
.

dig e s ti o n t a ke s pl a ce .

Unlik e Ame ba a sp e ci a l a na l spot ( Fi g 2 3 a n ) is p re s e nt in .

,
.

P a r a me ci u m th r o ugh which indig e s tibl e s o lid s a r e di sch a r g e d t o

th e o ut sid e Thi s Op e n s o n th e v e ntral sur fa ce ju s t b e hind th e
.

m o uth It ca n b e s e e n o nly wh e n m a t e r i al is c a s t o u t It i s no t
. .

y e t kn o wn wh e th e r th e a n a l sp o t is a p e r m a n e nt o r ific e wh ose

lip s a r e so tightly cl o s e d as t o b e invi s ibl e to u s or wh e the r a fre sh

o p e ning i s m a d e a t e a ch disch ar g e T h e p r o c e sse s o f dig e s ti o n
.
,

a b so rpti o n di ssimi l a ti o n e xc re ti o n r e spi ra ti o n a nd g r o wth a r e

, , , ,

s o simi l ar t o th o s e d e sc r ib e d f or A me ba th a t th e y n e e d no t b e

c o nside re d fu r the r a t thi s pl a ce ( se e pp 4 9 .

R e pr o d u cti o n P a r a m e ci u m r e p r o du ce s o nly by s impl e

.

bi na r y di vi si on Thi s p ro ce ss i s inte r r upte d o cc a sio nally by a

.

te mp o r a ry uni o n (conj u ga ti on) o f two individu al s a nd a su b se

quent mutu a l fe r ti li za ti on .

B inar y fi s s i o n — I n bin a r y fi ss i o n th e a nim a l div id e s t r a n s

.

v e rs e ly ( Fi g . T h e fi r s t indic a ti o n o f a fo r thc o ming divi s i o n

i s s e e n in th e mic r o nucl e u s whi ch und e rg oe s a s o r t o f m it os i s
,

( Fi g 3 0 8 a nd
.
,
i ts s ub s t a nc e b e ing e qu a lly d ivid e d b e tw e e n
th e t w o d a ught e r nu cl e i ; th e se se p a r a t e a nd fin a lly c o m e t o lie
o ne n e a r e ith e r e nd Of t h e b o dy F igu r e 3 0 sh o w s tw o dividi ng
.

mi cro nucl e i since th e re a r e t wo Of th e s e in P a r a me ci u m a u r e li a

,
.

T h e m acr o nucl e u s e l o ng a t e s a nd th e n divid e s t r a n sv e r se ly

( Fi g 3 0
.
,
T h e gull e t p r o duc e s a bud whi ch d e ve l o p s int o
a n o th e r gul l e t ; th e s e t w o s t r uctu r e s m o v e a p ar t t h e o ld gull e t ,

a dv a n cing t o th e v e nt r a l middl e lin e o f th e f o re p a r t Of th e b o dy ,

a nd th e ne w o ne t o a simil a r p o siti o n i n t h e p o s t e ri o r h alf

( g 3 7
F i . 0 ,
a n d T h e undul a ting m e mb ra n e r e m a in s with t h e

Old gull e t whil e a ne w o ne a r i se s in c o nn e cti o n with t h e ne w gull e t .

A ne w co nt r a ctil e v a cu ol e ( Fi g 3 0 I ) a ri s e s n e a r th e a nt e r i o r e nd
.
,

o f th e b o dy a n o th e r j u s t b a ck o f th e middl e lin e ( Fi g 3 0 .
,
,

Whi l e the se e v e nt s a r e t aking pl a ce a c o n s t r ictio n app e a r s n e a r

th e middl e o f th e l o n gitudin a l di a m e t e r o f th e b o dy ; thi s cl e a v ag e
68 AN INT RODU CT ION TO ZOOLOGY

fu rr ow b e c om e s d e e p e r and d e e p e r until only a Sl e nde r th re a d of

p r o topl a sm holds th e two h alv e s o f th e b o dy toge th e r Thi s .

co nn e cti o n i s fin a lly s e v e r e d a nd th e t w o d a ught e r P a r a m e ci a

a r e f r e e d f r o m e a ch o th e r E a ch co nt a in s b o th m a c r o a nd mi cr o
.

nucl e i tw o co ntra ctil e v a cu ol e s a nd a m o uth with gull e t T h e

, ,
.

e nti r e p ro c e ss o ccupi e s a b o ut t w o h o u rs T h e tim e h o w e v e r

.
, ,

v a r i e s co nsid e r ably d e p e nding up on th e t e mp e r a tu r e o f th e w a t e r

, ,

th e qu ality a nd qu a ntity o f f o o d a nd p r ob a bly o th e r f a ct o r s

,
.

T h e d a ught e r P a r a m e ci a inc r e a s e r apidly in siz e a nd a t th e e nd o f ,

tw e nty fo u r ho ur s di vid e a gain if th e t e m p e r a tu r e r e m ain s a t

-

f ro m 1 5 to 1 7 C ; if th e t e m p e r a tu r e i s r ai se d t o 1 7 2 0 C
° °
.
— ° °
.
,

t wo divi si o n s m ay t a k e pl a ce in o ne d a y
C onj u ga ti on —.At a ce r t a in tim e in th e lif e cy cl e o f P a r a me ci u m
c o njug a ti o n occu r s T h e conditi o ns th a t initi a t e thi s p roce ss ar e
.

no t ye t kn o wn but th e c o m plic a t e d st a g e s h a v e b e e n quit e fully

,

w o r k e d ou t Whe n two P ar a me ci a w hi ch a r e r e ady to conjug a t e

.
, ,

c o m e to g e th e r th e y r e m ain a tt a ch e d t o e a ch o th e r b e ca u se o f th e
,

a dh e siv e s t a t e o f th e e xt e r n a l p r o t o pl asm T h e v e nt r a l su r f a ce s
.

o f th e tw o a nim a l s a r e Opp ose d a nd a p r o t o pl a smic b r idg e i s

,

co n s t r u ct e d b e tw e e n th e m AS so o n as thi s u ni o n i s e ff e ct e d the
.
,

nucl e i p a ss th r ough a se ri e s o f st age s whi ch h av e b e e n lik e ne d to

th e m a tu ra ti o n p r o ce sse s o f m e t a z o a n e gg s ( C h a p VI I p .
,
.

R e f e r e nce to Figu r e 3 1 will h e lp to m a k e cl e a r th e fo ll o wing d e

s c r ipti o n . T h e mic ro nu cl e u s m ov e s f r o m its n o r m a l p o s iti o n in
a c o n cavity o f th e m acr o nu cl e u s ( Fi
g 3 m i ) a nd gro ws l a rg e r
2 .
, , ,

its ch ro m a tin b r e aking up into g r a nul e s whi ch r a di a t e fro m a

divi sio n ce nte r a t one e nd ( Fi g 3 1 a ) T h e nu cl e u s th e n l e ngth
.
,
.

e ns , fo r ming a spindl e a nd s ub se qu e ntly di vide s into two

,

The se imm e di a te ly divid e again witho ut th e int e rv e nti o n o f a

r e s ting s t a g e .T h e r e su lt a nt fo u r nucl e i (c) h a v e b e e n co mp a r e d
to th e f o u r sp e r m s p r o duc e d by a p r im a r y sp e r m a t o cyt e o r t o a n
e gg with i t s p o l ar b o d i e s a nd th e divi s i o ns a re c o n sid e r e d a s th e
,

fi r s t a nd s e cond m a tu r a ti o n mit o se s (s e e pp 1 0 3 Thr e e

.

o f th e fo u r nucl e i d e ge n e r a t e (d) th e fou r th divid e s a g a in

,
.

Du ri ng thi s d ivi si o n th e re a r e no d e fin it e spindl e fib e r s a nd no

 P ARAM E C IUM 69

l ongitudin al splitting of the ch ro m oso m e s but th e gr a n ul e s of,

ch r o m a tin c o nt a in e d in th e nucl e i se p ar a t e int o tw o g ro up s o ne ,

sm all e r ( Fi g 3 2 A m n ) th a n th e o th e r ( Fi g 3 2 A f
.
, ,
. . .
, , .

i t

3 1 . Di a gr a m sh o wi ng t h e s t a ge s i n t h e co n j u g a ti o n o f t w o P ar a m e ci a

a nd th e su b se q u e nt d ivi s i o ns o f t h e co nj u a nt s d ur i ng t h e p e r i o d o f
g
nu cl e a r r e cons t r u cti on. F o r d e s cr i p tio n se e t e x t .
( Af t e r M au pas ) .
70 AN INTR ODU CTION TO ZOOLOGY

m . 73 .

FI G 3
. 2 . T w o vi e w s o f th e m i cr o nu cl e i d u r i ng th e j
co n u gati on o f P a ra m e ci um .

A ,
th e sp i nd l e f orm e d d ur i ng t h e d ivi s i o n o f t h e m i cr o nu cl e u s whi ch

r e s u lt s i n t h e p r o d u cti o n o f a l a r ge f e m a l e nu cl e u s ( f n ) a nd a sm a ll e r
. .

m a l e nu cl e u s (m n ) B t h e f u s i o n o f th e m a l e nu cl e u s (m n ) o f o ne
. .
,
. .

a ni m a l with t h e f e m a l e nu cl e u s ( f n ) o f t h e o th e r co nj u ga nt
. . .
(From
C alk ins a nd C u ll i n Archi v f P r oti st )
. . .
72 AN INTR ODU CTION To Z OOLOGY

of vitali ty r apid c e ll multiplic a tio n a nd g rowth and a ctiv e

, ,

functio n s ; a t m atu ri ty th e ce ll divi si o n b e c o m e s l e ss rapid

, ,

gr owth e nds a nd th e o rga m sm b e co me s s e xually m a tu r e ; in old

,

a ge c e ll s b r e a k d o wn functi o n s b e c o m e imp e r f e ct a nd d e g e n e r a
, , ,

ti o n s e ts in e nding in n a tu ra l de a th The se thr e e st ag e s a re fo und

,
.

in the life cycl e o f P a r a me ci u m p r o vide d w e a cce pt th e a b o ve

,

d e fi niti o n of th e p ro to z oa n individu al ; y o uth i s cha r a cte rize d

by rapidly dividi ng c e ll s ; m a tu r i ty by the a ttainm e nt of full
size a nd co njuga ti o n ; a nd old age by de g e n e r a tio n and n a tura l
d e a th if co nj ug a ti o n is p re v e nt e d .

In the s t age o f you th P a r a me ci a d o no t c o njug a te e ve n if m any

a re c o nfin e d in a limit e d sp a ce T h e a nim a l s divid e v e r y ra pidly
.

a t thi s tim e Th e y a r e u su ally alm os t t r a n sp a r e nt a nd fr e e

.

fro m re s e r ve m a t e ri a l o f a ll kinds a nd a r e abl e to re si s t a dve r se

,

co nditi o ns sho wing a hi gh g r a de of vit ality ; this is d u e to the

,

e xc e s s Of c o n s t r uctiv e o v e r d e s t r uctiv e m e t a b o li s m The re is .

no d e finit e limit to y o uth M a tu r ity co m e s o n imp e r ce ptibly

.
,

a nd m a tu r e a nim a ls ca n b e r e c o gni z e d o nl y wh e n in so m e ph a s e

o f se xu a l a ctivity .

In a cultu r e und e r c o ntinu al ob se rva ti o n a de clin e in th e ra te

o f r e p r o ducti o n indic a t e s t h e a pp r o a ch o f m a tu r i ty T h e p r o to .

pl a sm a t thi s st ag e unde r go e s a ch ang e b o th physic ally a nd

che mic ally ; th e s ur fa ce l a ye r b e co m e s s ti cky so th a t whe n two ,

c e ll s m e e t the y fu se and c o njug a ti o n r e s ults This fr e que ntly

, .

o ccu r s in a l a r g e numb e r o f a ni m a l s in a s ingl e cultu re a t th e s a m e

”
tim e and a so c a ll e d e pide mic o f co njug a ti o n m ay th e n b e
-

Ob se r v e d .T h e co njug a nt s a r e s m a ll e r th a n th e o th e r sp e cim e n s ,

b e ing o nly 2 1 mm l o ng whi l e th e u su al l e ngth i s a b o ut 3 mm

. .
,
.

T h e imm e di a t e r e s ult o f c o njug a ti o n i s a pp are ntly th e r e j u v e na

ti o n o f th e co njug a nt s .

If th e P ar a m e ci a a re k e pt in a c o n s t a nt m e dium e g h ay i h ,
. .

”
fu si o n the y unde r go a p e ri o d o f physiol o gi ca l d e p re ssio n ab o ut
,

e v e r y th r e e m o nth s a s sh o wn by t h e d e c r e ase in th e i r r a t e o f
,

divi sio n The se p e ri o ds o f d e p r e ssi on ar e d u e to unkn ow n m e t a

.

b oli e conditi on s whi ch l e s se n th e r api di ty of divi si o n but d o not

, ,
 P AR AM E C IUM

c a us e th e de a th of the a nim al Se m i annu al p e rio ds al so o ccur

.
,

but r e cov e r y f ro m the se doe s not t a k e pl a ce if th e a nim als a re

k e pt unde r c o nst a nt c o n di ti o n s o r co nj ug a tio n i s p r e v e nt e d but ,

t h e p r o t opl as m d e g e n e ra t e s a nd b e c o m e s v a cu o l a t e d a nd th e a ni
m al s l ose the i r e n e r gi e s a nd fin ally di e .

E xp e r im e nt s h a v e b e e n p e r f or m e d which s e e m to Sh ow th a t
in a var i e d e nvi r o nm e nt n e ithe r co njug a ti o n nor de a th fro m old
a ge n e c e ss a r ily o c cu r s .Thu s Wo o dr uff ( 1 90 9 1 9 1 0) h as ca rrie d
,

a cultu r e o f P a r a m e ci a th r o ugh a p e r i o d o f thi r ty s e ve n m o nths

-

by ch a nging th e ch ar a cte r o f th e m e di um d aily Du r ing thi s .

tim e the re we re s e v e nt e e n hund r e d a nd nin e ty fi v e g e n e r a ti o ns

- .

At th e e nd o f th e thi r ty se v e nth m o nth th e a ni m a l s w e r e n or m al

-
, .

T h e cycl e m a y thu s b e p r ol o ng e d fr o m si x m o nths t o o v e r thi r ty

s e v e n m o nths by e mpl o ying a v a r i e d cultu re m e di u m Since in .

n a tur e th e s timuli d e r iv e d f r o m ch a ng e s in th e e nvi r o nm e nt

p rob ably a re p r e se nt th e l e ngth o f th e cycl e m ay p e rh ap s be
,

p r ol o ng e d ind e finit e ly .

B e h avi or. P a r a m e ci u m is a m or e a ctiv e a ni m al th a n Ame ba ,

s wimming a cr o s s th e fi e ld o f the mic ro s co p e so rapidly tha t care

ful ob se rva ti o n s a r e n e c e ss ar y to di s cove r th e de ta il s o f i ts m ov e
m e nt s As in Am e ba its a ctiviti e s a r e e ith e r spontane ou s tha t i s
.
, , ,

initi ate d b e c a u se o f so m e int e r n al influ e nce o r re sult f ro m s o me

,

e xte r nal s ti m u lu s
. Thi s s timulus is in a ll case s a cha nge i n the
e nvi r onm e nt . F o r e x a mpl e if a d r o p o f di s till e d w a t e r is a dd e d to
,

a d ro p o f o r di n a r y cultu r e w a t e r co nt a ining a numb e r o f P a r a

me ci a all o f th e a nim al s will e nt e r a nd r e m ain in th e di still e d

,

w a te r ; the y ar e sti m u la te d to a c e r t a in kind o f a ctivity by the

'

change in th e c o mp ositi o n o f th e w a te r The y wi ll soo n b e co me

.

a c clim a t e d t o th e i r ne w s u r r o un di ng s a nd will b e h a v e th e m se lve s

,

within th e di still e d w at e r in a n or m al m anne r until a n o the r ch ange

in the i r e nvi r onm e nt stimul a te s th e m t o fu r the r r e a cti o ns .

P ar a me ci u m r e sp o nd s t o s timuli e ith e r ne ga ti ve ly o r posi ti ve ly

Th e n e g a tiv e r e sp o ns e i s kn o wn as th e a void i ng r e acti on

it ta k e s pl ace in th e fo ll o wing m a nn e r Wh e n a P a r a me ci u m
.

r e c e iv e s an inju r i o u s stimulu s a t i t s a nt e r i or e nd it r e v e rs e s i ts
,
74 AN INT RODU CTION TO ZOOLOGY

cili a a nd s wim s b a ckw a r d f o r a shor t di st ance ou t of th e r e gi on of

stimul a ti o n ; the n its r ot a ti o n de c r e a se s in r apidity a nd it sw e rv e s
t ow ar d th e a b or a l side m ore s t ro ngly th an unde r n o r m al c ondi
ti on s I ts p ost e r i or e nd the n b e co m e s a s or t o f piv o t up o n which
.

th e a nim a l s wing s a b o ut in a ci r cl e ( Fi g D u r ing thi s r e v o

.

lu ti o n sa mpl e s o f th e s u r r o unding m e dium a r e b r o ught int o th e

o r a l gr oo v e Wh e n a s ampl e no l o nge r c o nt ain s the stimulu s
.
,

t h e cili a r e s um e th e i r n o r m al b e a ting a nd th e a nim al m ov e s

7 .
1 .

FI G 3 3
. D i agr a m
. of vo i d i ng r e a cti o n o f P ara me ci u m A i s a so li d
th e a .

o b j e ct o r o th e r so u r ce o f s ti m u l a ti o n 1 6 su cce s s iv e po s iti o ns o ccu p i e d

.
—
,

b y th e a ni m a l.
(T h e ro t a tio n o n th e l o ng a xi s i s no t sh o w n ) ( Fro m
.

Je nni ngs ) .

f or w a r d a g ain If thi s o nce m o r e b ring s it int o th e r e gi o n o f th e

.

stimulus t h e a v o iding r e a cti o n i s r e p e a te d ; thi s g o e s o n a s l o ng

,

a s t h e a nim a l r e c e iv e s t h e s timulu s T h e r e p e titi o n o f th e a v oid

.

ing r e a ctio n i s v e r y w e ll sho wn whe n P a r a me ci u m e nt e r s a d r op

o f 5 5 p e r c e nt a c e ti c a cid In a tt e mpti ng t o ge t o u t o f th e d rop
1
.

th e s u r ro un di ng w a t e r i s e n co unt e r e d ; to thi s t h e a v o iding r e a c

ti o n i s giv e n a nd a ne w dir e cti o n i s t a k e n within th e a cid whi ch ,

o f c o u r s e l e a d s t o t h e w a t e r a nd a n o th e r n e g a tiv e r e a cti o n T he .

a c co mp anying F igu r e 3 4 s h o w s p a r t o f t h e p a thw a y m a d e by a

s ingl e P a r a m e ci u m und e r th e s e co nditi o n s .

P osi ti ve Re a cti on If a littl e a cid i s pl ace d in th e ce nte r of a

.

l arge dr op of wa te r c ont aining a numb e r o f P a r a me ci a a ll of ,

 P AR AM E C IU M 75

th e nim als in the d r o p will s o on e r o r l a te r e nco unte r the a cid

a
,

a nd h a ving o n ce e nt e r e d are un a bl e t o e s c a p e ju s t a s in th e c a se ,

de sc rib e d ab o v e A g r o up i s th e r e fo r e f o r m e d in t h e a cid illus

.
,

t ra ti ng wh a t i s ca ll e d a p os itiv e
r e a cti o n a nd in thi s c a se i ti
pos ve

che motr opi s m Thi s e xp e rim e nt .

F I G 3 4 P a th f o ll o w e d by a s i n
. .

gle P a r a me ci u m i n a dr op o f
ac id .
( Fro m Je nni ngs ) .

p e a t e d u sing a lg m ay be re

p e r c e nt s o luti o n o f c o mm o n
s alt in which a re pl ace d a
numb e r o f sp e cime n s If a .

d rop o f 1 5 pe r c e nt s oluti o n 1

Of t h e s a m e ch e mi ca l i s no w

a dd e d th e P a r a me ci a wi ll ,

s wim int o a nd di r e ctly a cr o s s

it but o n r e a ching th e b o und ,

FI G 3 5 D i g m
. . h wi ng P m i a r b e twe e n th e tw o s oluti o ns
a ra s s o a ra ec a
y
ll t d b t b bbl f C O i n
co ec e a ou a u e o 2
o n th e O l w r Sl d e o f th e d r o p
t h Op ti m m
e n nt ti n ; i
u

t h
co
f d
ce
n
ra
t n W
o
11 b e
a s a
o
,

b b bl u f i b f C0 e oA h w a r,e a v o I l
o g r e a c i o
2 .
1
s o s

th p p e ti n t w m i n t
re ar a ft give n Soo n th e w e ak e r so
o o u es a er .

th i ntm d e ti n f th C 0 ; B t W uc
lu ti o n will co nt ain all o f th e
o O e 2 ,
0

m n t il t ; C ght n m n t a ni m a l s whi ch h a ving o nce

u es a er ,
ei ee i u es
,
l t ( F m J nn ng )
a er ro e i s
e nt e r e d ca nn o t e sca p e In
. .

.
,

m any ca se s a s a b o v e th e p assa ge fr o m a s t r ong soluti o n to a

, ,

we ak soluti o n ca u se s no r e a c ti o n F or ce r tain s ub s t a nce s h ow . ,

76 AN INTROD U CTION TO Z OOLOGY

e ve r the r e is a de finit e str e ngth which s e e m s t o s uit th e P a r a

,

me ci u m b e tte r th a n a ny o th e r a nd no re a cti o n t a k e s pl a c e o n
,

e nt e r ing it P as s ag e fr o m s uch a soluti o n t o e ith e r a w e a k e r o r

.

a st r o ng e r c a lls f o r th th e a v o i di ng r e a cti o n T h e co nce nt r a ti o n

“
.

”
i s th e re fo re c all e d th e opti mu m F o r e a ch ch e mic a l th e r e i s
.

a c e r t a in Op t imum c o nce nt r a ti o n in whi ch t h e P a ra m e ci a a r e no t

c a use d to re a ct P assag e fro m .

thi s optimum to re gi o n s o f
e ith e r g re a t e r or l e ss co nc e nt ra

ti o n ca use s th e a vo i di ng r ca c
ti on so th a t th e a nim al s t e nd ,

“
a t o r e m a in in th e r e gi o n o f th e
Optimum a nd if thi s r e gi o n i s ,

s m a ll t o f o r m h e r e a d e n se Col ,

l e cti o n ( 6 2 p 66 Fi g ,
.
,
.

P a r a me ci a m a y giv e a ny o ne
Of thr e e re ac ti o ns t o contact
s ti mu li ; th e fi rs t t w o are n e g a

FI G 3 6 P
. . m i m t
ara ec u t with n tiv e th e thi r d p o sitive ( I ) If
a re s a ,
.

t i nd g i n t
e r or e a m a f bs P a r a m e ci u m s wim s a g a in s t a n
a a ss o ac

t i l 81 h OW i ng h
er a 2 00 09 3
t O b s t a cl e
S
or if th e a nt e r i o r e nd
e
, ,
nts
curr e by th h ° Cl
whi ch i s m or e s e n sitive th a n
a '

Fm m J nn ng )
,

e i s
th e o th e r p a r t s o f th e b o dy i s ,

t o u che d with a gl a s s ro d th e a vo idi ng r e a cti o n i s give n ( 2) Wh e n

,
.

a ny o th e r p a r t Of th e b o dy i s s timul a t e d in a lik e m a nn e r
-
,

th e a ni m al m a y s imply s wim f or w a r d (3 ) F r e qu e ntly a P

. a r a

me ci u m up o n st r iking a n o bj e ct wh e n s wimming s l o wly co m e s to

, ,

r e st with its cili a in c o nt a ct with th e o bj e ct ( Fi g Thi s .

p ositiv e r e a cti o n oft e n b ring s th e a nim al s into a n e nvi ro nm e nt

r ich in foo d .

P a r a m e ci a d o no t r e sp o nd in a ny w a y t o or di n a r y vi sibl e li ght ,

but give th e av o iding r e a cti o n wh e n ultr a -vi ol e t r a y s a re thr o wn

up on th e m ; if unabl e to e scap e de a th e n su e s in fr o m te n to fifty ,

s e conds
Th e opti mum te mpe r a tu r e f o r P a r a me ci u m li e s und e r o r din a ry ,
 P AR AM E C I U M

c o nditi o n s b e tw e e n 24 and 2 8 C A numb e r o f ani m als pl a ce d

,
° °
.

o n a slid e which i s h e a t e d a t o ne e nd wi ll s wim a b o ut in all d i r e c

, ,

ti o n s giving th e a v o iding r e a cti o n wh e r e s timul a te d until the y

, ,

b e co m e o ri e nte d so a s to m ov e t ow a r d th e c ool e r e nd Thi s is th e .

m e tho d Of tr i al and e rr or th a t i s th e ani m al t rie s all di re cti o ns

, ,

until th e o ne i s di sco v e r e d whi ch all o ws it to

e s c ap e f ro m th e r e gio n Of injur io us stimul a
ti o n .

Gr a vi ty in s o m e unkn o wn w ay c a us e s P a r a me
ci a to o r i e nt th e ms e lv e s with th e i r a nt e r i or e nds

p oint e d upw a r d Thi s b r ing s th e m n e ar th e

.

sur fa ce o f th e w a t e r If a numb e r are e qu ally

.

di st ribut e d in a t e s t tub e o f w a t e r th e y will ,

g ra du ally find th e i r w ay to th e t op ( Fi g .

T h e m o st p r ob a bl e th e or y t o a ccou nt f or this
‘

i s th a t th e sub s t a nc e s with in th e b o dy b e ing ,

Of di ff e re nt s p e cific g r a viti e s m o v e a b o ut wh e n
,

th e a nim al ch a ng e s i ts p o siti o n a nd a ct as s tim

uli r e li e f b e ing obt ain e d o nly wh e n th e a nim al
,

h as pl a c e d i t s b o dy with i ts lo ng di a m e t e r
p e rp e ndicul a r to th e e a r th s su r fa ce a nd i ts
’

FIG 3 7 P m ia
a nt e r i o r e nd up Und e r ce r t a in c o ndi . . ara ec

ll t d t co ec e a
ti o ns P a r a me ci u m e xhibit s p ositiv e g e o t r opi sm th
.

t p f e o o a

If P ar a m e ci a a r e pl a c e d i n r u nni ng wa te r v ti l t b
, er ca u e .

the y o ri e nt the m se lv e s with the a nt e r i o r e nd s (F m J n ro e

upst r e a m a nd swim a g a in s t th e cu r r e nt T hi s .
i n g f t h s a er

i s p r ob ably ca u se d by th e int e r fe r e n ce o f th e
J n n) e se .

cu rr e nt with t h e b e a ting o f t h e c i li a f o r a s so o n a s a n a nim a l

,

r e a ch e s a p o s iti o n with a nt e r i o r e nd up s t r e a m th e wa t e r no ,

l ong e r t e nds t o r e v e rse th e cili a .

P a r a me ci a m a y b e s ubj e ct e d t o a n e le ctr i c cu r r e nt in th e l a b o r a
to ry but the y a r e n e v e r influ e nc e d by it unde r n o rm a l co nditio ns
,
.

A we a k cu rr e nt c a u se s a m o v e m e nt t o w a r d th e c a th o d e ; a st r o ng
curre nt re v e r se s t he di r e cti o n o f th e b e a ting o f th e cili a a nd c a use s
th e a nim a l s t o swi m b a ckw a r d t o w a r d th e a n o d e M a ny o the r.
7 8 AN INT RODU C TION TO Z OOLOGY

int e r e s ting phe n om e n a might b e cit e d but the e nti r e subj e ct is

,

t o o c o mpl e x f o r b ri e f di s cus si o n .

F re qu e ntly P a ra me ci u m m a y b e stimul a t e d in m or e th a n o ne
w ay a t t h e sa m e tim e F o r e x a mpl e a sp e cim e n which is in
.
,

c onta ct with a solid i s a ct e d up o n by g ra vity and m ay b e a ct e d

,
-

up o n by ch e mical s h e a t cu rr e nt s of wa t e r and o the r s timuli

, , ,

( g
F i . It h as b e e n f o und th a t g r a vity alwa ys give s way to
o th e r s timul i a nd th a t if m o r e
,

th a n o ne o the r f a ct o r i s a t wo r k
t h e o ne fi r s t in th e fi e ld e x e r t s
t h e g r e a t e r influ e nc e .

B o th th e sp o nt a n e o us a cti vi
ti e s a nd r e a cti o n s d u e t o e x
t e r n a l s timuli a r e d u e t o cha ng e s
F G 38 P
I . . m i ara ti m l t d by in th e int e r n a l co ndi tio n of th e
ec a s u a e a

c h e m i ca l a nd by co nt a ct a t t h e
a nl m a l T h e phys w logi ca l con
s a m e ti m e T he y h a ve f o rm e d
.

di ti n O f P r a m e ci u m th e r e f o r e
a r i ng a bo u t a b u b b l e o f C 02
o a , ,

a nd h a v e th e n co m e to r e s t d e t e r min e s th e ch ar a ct e r Of i ts
a ga i ns t th e gl a ss s u ppo r ti ng r e sp o n se Thi s physi ol o gica l
.

r o d s f o r m i ng t wo d e ns e gro u s
, p st at e is a dyn a mic co nditi o n
.

,
( Fro m J e nm ngs )
ch a nging c o ntinu ally with the
'

p r o ce s s e s Of m e t ab oli s m g oing o n within th e living s ubsta nc e o f

t h e a nim al Thu s one phy si ol o gic a l s t a t e r e solv e s its e lf into
.

a n o th e r ; thi s b e co m e s e a si e r a nd m or e r apid a ft e r it h a s t ak e n
pl a ce a numb e r o f tim e s ( 6 2 p ,
gi ving u s g r o unds f or th e
.

b e li e f th a t s timuli a nd r e a cti o n s h av e a di s tinct e ffe ct up o n s u c

ce e d i ng r e sp o n s e s .

We m ay s u m up th e e xt e r n al f a ct o r s th a t p r o duc e o r d e t e r
min e r e a ctio n s a s foll ows : ( I ) T h e o rg a ni s m m ay r e a ct t o a
cha nge ,
e v e n th o ugh n e ith e r b e n e fici a l no r i nju r i o u s ( 2
.
) A n y
thing th a t t e nds t o int e r fe r e with th e n o r m al cu rr e nt o f life
a ctiviti e s p ro duc e s r e a cti o ns Of a ce r t a in s o r t

(3 ) A n y c h a ng e th a t t e nd s t o r e s t o r e o r f a v o r t h e n o r m a l lif e

p ro ce sse s m ay p r o duce r e a ctio n s o f a d iffe r e nt sor t p o sitive

(4) C h a ng e s th a t in th e m s e lv e s n e ith e r int e r f e r e with n or a ssi s t
 C HAP TE R VI

OTHER P R OT OZ OA

r . C LAS SI F I C ATI ON OF TH E P R OTOZ OA

""
A P R OTOZ OON is a p rimi tiv e a nim al orga nism usu ally con
Si s ting o f a singl e c e ll who s e p ro topl asm b e co m e s di s tribute d
'

am ong m a ny f r e e living c e lls The se r e p ro duc e the ir kind by

.

divi sio n by buddi ng or by sp or e fo r m a tio n th e r a c e thu s for m e d

, ,
“
p assi ng th r o ugh di ff e r e nt fo r m cha nge s and th e p r o topl asm
through v a r i o u s st a ge s of vit ality c oll e ctiv e ly known a s th e life
cycl e ( 78 ; p .

P r o t o z o o ns m ay b e s e p ar a t e d int o g r o ups a cc or ding to the

p r e se n ce o r ab se nc e of l o co m o t or o r gan s a nd the chara cte r o f
-th e s e whe n p r e se nt Fo ur divisi o n s a r e u su ally r e co gniz e d
.

( I ) R hiz o p o d a with pse ud op od i a ( 2) M a stigoph or a with fl a ge ll a

, ,

( 3 ) I nfus or i a with cili a a nd (4) Sp orozoa which a re p ar a sitic in

‘

, ,
‘

ce ll s and witho ut m o til e o r gan s .

T h e siz e Of P ro to z oa r a ng e s within wid e limits So m e a r e .

v e ry l a r ge ,
f o r e x a m pl e a p a r a sit e P or ospor a gi gante a whi ch
, , ,

liv e s in th e alim e nta rycan al o f th e l ob s t e r r e a che s two thi r ds of ,

an in ch in l e ngth ; so m e a r e ju s t a t t h e limit o f vi s i o n with t h e

m os t p o w e r ful mic r osco p e s w h e r e a s o th e r s lik e th e y e ll o w fe v e r

, ,

p ar a site do ubtl e ss e xi s t tho ugh the y h av e n e v e r b e e n se e n

, ,
.

Individu als b e l onging t o o ne Sp e ci e s v ar y in Siz e a cco r di ng to

th e i r food co nditi o n s and age T h e v ar i a ti on s in s tr uctur e and
.

lif e hi s t ori e s o f th e P r o to zo a a r e so num e r o u s a s to p r e clud e d e

scripti o n in a b oo k Of thi s ch ar a ct e r We Shall the r e fore give a n
.

abb r e vi a t e d cl assifi ca ti o n Of t h e phylum a nd c o nsid e r typic al

e xa mpl e s und e r e ach cl a s s .

 OT HER P R OTOZ OA

P hylu m P r otozoa

C l a ss I . Rhi zop o d a Typ e : Ame ba pr ote u s P ro to zo a us uall y

. .

Simply in s t r u ctu r e m o ving by m e a ns Of p s e ud op o di a ;

,

p ro to pl a sm n a k e d .

Or d e r Gym na m e b a Witho ut She lls

. .

F a mily Am e b i dee P s e udo p odi a l o b o s e

. .

Ge nus Am e b a E ctopl a s m a nd e nd o pl a s m di s ti nct

. .

Sp e ci e s p r o t e u s ( N a m e d a ft e r a se
. a go d w h o h a d th e

p o we r o f a ssuming di ff e r e nt Shap e s ) .

C l ass 2 . M a stigophora Ty pe s : Eu gle na vi r i di s Volvox globa tor

.
, .

P r o to zo a with o ne o r m ore fl a ge ll a ; Simpl e st f or m s

re s e mbl e b a ct e r i a

Ord e r E ugl e nid a La r g e f or ms with o ne o r t wo fl a g e ll a ;

.

m o uth a t b ase of fl a ge llum thro ugh which co nt ra ctil e va cuol e

Op e n s to o uts id e .

F a mily E ugl e nid ae E l o ng a t e ; o ne fl a ge llum ; r e d e ye Sp o t ;

.

g re e n chr o m a topho r e s .

Ge nu s E ugl e n a B o th e nds co nt r a ct e d
. .

Sp e ci e s vi ridi s (g r e e n ) .

Or d e r Volvocina .C o l o ni a l f or ms with two fl a g e ll a .

F a mily Vo lv o c a c e ae .

Ge nu s Vo lvo x C ol o ni e s Of m a ny c e ll s in a sph e re
. .

Sp e ci e s glob a t or ( a b a ll ) .

C la ss 3 In fu so r i a Type f P a r a me ci u m cau da tu m Lo como t or

. . .

o r g a n s cili a p e r m a n e nt o r limit e d to y o ung st a g e s ;

, ,

t w o kinds o f nucl e i m a cr o nucl e i a nd mic ro nucl ei ,

.

Or d e r H ol o t r ichi d a C i li a Simil a r all o v e r b o dy ; t rich o cys t

.

p re se nt .

F a mily P a r a m e ci dm Or a l g r oo v e p r e s e nt
. .

Ge nu s P a r a m e cium M o uth n e ar middl e o f b o dy ; ph ar

.

y n x S h o r t ; o r a l g r o o v e Obliqu e .

Sp e ci e s ca ud a t um ( t a il e d) .

c
82 AN IN T RODU C TI ON To Z OOLOGY

C l a ss 4 . Sp oro z oa Typ e : P las modi u m vi vax P ar asitic

.
1
. P ro
t ozo a ; no m o til e or g a ns .

Or d e r H e m osp o ridi a B l oo d dw e lling Sp or o z o a

.
-
.

Ge nus P l as m o dium .

Sp e ci e s viva x ( liv e ly) .

2 . E U G LE N A

( Eugle na vi r idi s E h rb g ) .

Eugle na is th e a nim a l us u ally s e l e cte d in p re p ara to ry

vi r id i s

co u rs e s t o re p re se nt the C l a ss M as tigopho r a It i s f o und in .

f re s h-wa te r p o nds a nd m a y app e ar in cultu r e s p r e p are d a s d e

sc rib e d o n p ag e 3 8 It i s g r e e n in co l or a nd tho ugh a singl e
.
, ,

a nim a l ca nn o t b e se e n with t h e n a k e d e ye wh e n a g r e a t m a ny a r e ,

m a ss e d t og e the r the y imp ar t a g r e e n tint to th e wa t e r .

Ana tomy . Eu gle na ( Fi g 3 9) i s a singl e e l o ng a t e d c e ll p o int e d

.

a t th e p o s t e r i or a nd blunt a t th e a nt e r i o r e nd
, T w o kind s o f .

cyt o pl a sm m ay be di s tingui sh e d in Eu gle na a s in Am e ba a nd

P a r a m e ci u m a d e n se o ut e r l a y e r t h e e ctosa r c a nd a c e nt ra l m a s s
, , , ,

t h e e nd os a r c w hi ch i s m o r e fluid
,
A thi n cu ti cle is p r e s e nt a s in.
,

P ara me ci u m co ve r ing t h e e nti r e sur f a c e o f t h e b o dy

,
P ar all e l .

thicke ni ngs of this cuticl e ru n oblique ly a r o und th e a nim al ,

m a king it app e ar s tr i a te d ( B ) A littl e to o ne Side of th e c e nte r

.

o f th e a nt e r i o r blunt e nd o f t h e b o dy i s a funn e l s h a p e d d e pr e s -

sio n kn o wn a s th e m ou th (A At th e b o tt o m o f thi s d e pre s

,

si o n is a n Op e ni ng whic h l e a ds int o a shor t duct c all e d th e gu lle t .

1
Only o ne o f t oz o a n t ype s (P las m odi u m vi va x) d i scu ss e d i n thi s
th e p r o
boo k i s p a r a s iti c T h e r e a r e h o w e v e r p a ra s iti c sp e ci e s i n e ach o f th e cl as s e s
.
, ,

o f P ro tozo a . F o r e xam pl e t h e R hi z o p o d E nta m e ba hi s tolyti ca i s h e l d r e

, , ,

s po ns i b l e f o r a m e b i c d ys e nt e ry th e F l a ge ll a t e Tr ypa nos oma ga m bi e ns e i s a

, ,

bl oo d p a r a s it e whi ch cau se s sl e e p i ng s i ck ne ss i n Af r i ca ; a nd th e C ili a t e ,

B a la nti di u m coli whi ch i s f o u nd i n t h e ali m e nt a ry ca na l Of m a n i s s u ppo s e d

, ,

t o pl ay a p a r t i n ca t ar r h a l i nfla m a t i o n o f t h e i nt e s ti ne E a ch cl a ss o f P ro .

t o z oa co nt a i ns m a ny o th e r p a r as it e s o f bo th m a n a nd t h e l o w e r a ni m a l s so m e ,

be i ng app ar e ntly h a rml e s s wh e r e a s o th e r s a r e d ange r o u s a n

,
d f r e q u e ntly f a t al .
84 AN INT RODU CTION TO Z OOLOGY

Thi s in tur n e nt e rs a l ar ge sphe r ical ve sicl e th e r e se rvoi r (r ) , ,

into whi ch se ve r a l minut e contr acti le va cu ole s (A c v ) dis charge ,

. .

the ir c o nt e nt s .

A c o nspicu o u s s t r uctu r e in Eu gle na i s th e r e d e ye spot or s ti gma

( g 39
F i . A ,
This
,
I S pl a c e d n e a r t h e inn e r e n d o f t h e g u ll e t
cl o se t o th e r e s e r voir It co n si sts o f p ro t opl a sm in which a re
.

e mb e dd e d a numb e r o f gr a nul e s o f h ee m a tochr om e T h e a nt e r i o r .

e nd o f th e b o dy Of E u gle na i s s a id t o b e m ore s e n sitiv e t o light

th a n any othe r p a r t a nd it i s supp ose d by so m e tha t th e stigm a

,

functi o n s as a r a th e r p rimitiv e vis u a l Orga n Thi s vi e w i s m a d e

-
.

p r ob a bl e by th e p r e se nc e Of l e n s like para mylu m gr a ins ju st -

a nt e r i or t o it Th e h mm a to ch r o m e a l so h a s m a ny o f th e ch ar a c
.

"
,

t e r i sti cs o f t h e pigm e nt s in t h e e y e s o f high e r o r g a ni s ms If .

ke pt in t he d a r k E u gle na so o n l o se s i ts r e d pigm e nt A r e ce nt
,
.

vie w i s th a t th e h ae m at och rom e Sh a de s a se nsitive p ar ticl e o f

p ro t o pl asm .

E u gle na c o nt a in s a s ingl e o va l nu cle u s ( Fi g 3 9 n ) lying in a .

,
.

de finit e p o sitio n a littl e p os t e ri or t o th e ce nt e r o f th e b o dy It .

h a s a di s tin ct m e mb r a n e a nd c o nt a in s a ce nt r a l b o dy which h a s
,

b e e n c a ll e d a nu cle olu s ( E) but p r o b ably i s no t sinc e i t functi o n s

, ,

a s a divi s i o n c e nt e r du r ing mit o si s .

E u gle na d e rive s i t s g r e e n c o l o r fr o m a numb e r Of o v a l di sks

susp e nde d in t h e p ro t o pl a sm The se a r e kn o wn a s chr omato .

p h o r e s ( Fi g 3 9 A . Th e y a r
,
e a rr a ng e d a b o ut a c o ll e cti o n o f
g r a nul e s s i t iI a te d in th e ce nt e r Of th e b o dy a nd c o nt ain chl oro ,

phyll which i s diff u se d th ro ugho ut th e i r p r o t o pl asmic co nt e nt s

, .

The y m a nufa ctu r e fo o d by a p ro ce ss co mm o n in g r e e n pl a nts but

.

r a re in a nim a l s call e d photos ynthe s i s ( se e p 1 8 a nd Fi g

,
. .

T h e chl o r o phyll i s a bl e in th e p re se n ce Of light t o b r e a k d o wn t h e

, ,

c a rb o nic a cid ( C 0 ) thu s s e tting fr e e t h e o xyge n a nd t o unit e th e

2 , ,

c arb o n with w a t e r fo r ming a substa nce a llie d to s t a r ch call e d

,

p a r a m y lu m ( A a nd B If s p e cim e n,
s a r e k e pt in g oo d light
c o ntinu ally a l a r ge a m o unt o f p a r a mylum will b e s to r e d up f o r
,

fut u r e u se b e ing l aid do wn a r o und so m e g ra nul e s o f p r o t e id sub

,

st a nce n e ar th e c e nt e r Of th e b o dy The se g ra nul e s are c a ll e d .

 OT HER PR OTOZ OA 85

pyr e noi d s . B o th th e
py re n o id s a nd chr o m a t o ph or e s a r e p e r m a

n e nt c e ll st r uctu r e s a nd in cr e a se in numb e r by divi si o n a nd no t by ,

th e o r igin Of ne w o n e s f ro m th e o th e r p a r t s Of th e b o dy .

Lo com o ti on E u gle na ch a ng e s i t s s h a p e fr e qu e ntly b e co m

.
,

ing Shor t e r a nd thi cke r a nd Sh o ws c e r ta in s qui r ming m o v e m e nt s

,
.

Th e s e p ro ve th a t it p o s se ss e s co n side rabl e e lasti ci ty since th e nor ,

m al sh ap e i s r e g a in e d if e n o ugh w a t e r i s p re se nt Oft e n in a .

f avorabl e Sp e cim e n a thr e a d —like str uctur e m ay b e s e e n p roj e ct

,

ing fro m th e a nt e ri o r e nd o f th e b o dy a nd b e nding to a nd f ro

‘

d ra wing th e a nim a l a fte r it Thi s is th e f lage llu m It ar is e s f ro m

. .

a numb e r o f b ra n ching r oo t -lik e fib r ils within th e b o dy p a sse s ,

thr o ugh th e w all of th e m o uth de p re s s io n a nd e xt e nds for wa r d to ,

a di s t a nc e Oft e n e qu a l t o th e l e ngth o f t h e a nim a l T he p a r t o f th e

.

fl age llum o ut side o f th e b o dy is c o mp ose d Of fo ur co nt ra ctil e

fib ril s which a re wo und t o ge th e r spi r ally ( Fi g 2 T h e co n
.

t r a cti on of the s e fib r il s i s s upp ose d t o p ro duc e all o f th e m o ve

m e nt s ch a r a cte r i s tic o f this s t r ucture If th e fl ag e llum
c ann o t b e s e e n in th e living a nim al a littl e i o din e pl a c e d unde r
,

th e c o ve r gl a s s will h e lp t o b r ing it o u t .

N utr i ti on — .Alth o ugh E u gle na h a s a m o uth a nd gull e t it is ,

ve r y do ubtful if a ny fo o d i s t a k e n in F ood is m a nufa ctur e d as

.

i n g re e n pl a nts by th e a i d o f th e chl orophyll in t h e ch ro m a t o

,

ph ore s . Thi s m o de of nut r iti o n i s kno wn a s holophyti c Th a t all .

th e f o o d n e c e s sa r y f o r th e life o f th e a nim a l i s no t p ro cu r e d in this

w ay is sh o wn by th e fa ct th a t th e a nim a l is a bl e t o liv e in th e d ar k
f or o v e r a m onth wh e r e a s chlor ophyll d e m a nds light b e fo re th e
,

p ro ducti on Of p ara mylum is p ossibl e Thi s s e e ms to indica t e .

tha t o rg a nic s ub s t a nc e s in soluti o n ar e abs orb e d th ro ugh th e

s u r f a c e o f th e b o dy th a t i s s a r o h ti c nut r iti o n suppl e m e nt s t h e
p p y
, ,

ho l ophytic Th e nut riti o n Of E u gle na di ff e rs fro m th a t o f th e

.

m aj ority of anim al s since th e l a tt e r live by ing e sting so lid par

,

ti cle s Of f oo d a nd are s aid to b e holozoi c .

E ncys tm e nt Occa sro nally Eu gle nw ar e fo und which h a ve

.

b e com e a lmos t sphe r ic a l a nd a r e s u rr o und e d by a r athe r thi ck

g e la tinous co ve ring which th e y h ave s e cr e t e d Such an ani m al .
86 AN I NTR ODU CTION TO ZOOLOGY

is s aid to be e ncy s t e d (Fi g 3 9 F a nd G) In thi s co nditio n p e ri ods

. .

o f d r o ught a re s ucc e ss fully p ass e d t h e a nim a l s b e c o m ing a ctiv e

,

wh e n wa te r i s a gain e nco unte r e d Usually in cultur e s b ro ught

.

into the l ab or a t or y m a ny cys t s ar e fo und o n the side s of th e di sh .

F I G 40 Di agr a m o f t h e r e a cti o n o f E u gle na wh e n th e li ght i s d e cre ase d

. . .

T he o rga ni s m i s s wi m m i ng f o r w a r d a t I whe n it r e a che s 2 i t i s s h a d e d .

It th e re upo n s w e r ve s to w a r d th e d or sa l s i de a t t h e sa m e ti m e co nti nu
,

i ng to r e vo lve o n t h e l o ng a xi s so th a t i ts a nt e r i o r e nd d e s cr i be s a ci r cl e
, ,

t h e E u gle na o ccu pyi ng s u cce s s iv e l y t h e po s iti o ns 2 —6 . F ro m a ny Of

the se it m a y s t ar t f or wa rd i n th e d i re ctions i nd i ca te d by th e a rro w s .

(From Je nni ngs ) .

E ncys tm e nt fr e qu e ntly t a ke s pl ac e witho ut a ny app ar e nt c a u se ,

t h e a ni m a l r e s ting in thi s co nditi o n f or a tim e a nd th e n e m e r g

ing a g a in to i ts fr e e s wimming h abit B e for e e ncys tm e nt th e.

fl ag e llum is thr o wn off a ne w o ne b e ing p ro duc e d whe n a cti vity

,

i s a g ai n r e sum e d .
88 AN INTRODU C TION To ZOOLOGY

by m e a ns of its chlorophyll Eu gle na wi ll swim a wa y fro m the

.

di re ct ra ys o f th e sun Di r e ct s unlight will kill the org ani sm if

.

a ll o w e d to act f or a l o ng tim e If a drop o f wa t e r co ntaini ng

.

E u gle nce is pl a c e d in th e di r e ct s unlight a nd th e n o ne h a lf o f it is

sh a d e d th e a nim al s will a vo id th e sha dy p ar t a nd als o t h e di re ct
,

su nl ight b o th o f whi ch a r e inju ri o u s to the m a nd will re m a in in a

, ,

sm all b a nd b e tw e e n th e t wo in the light b e s t s uite d f or th e m ,

th a t i s the i r opti mu m ( Fi g
,
B y Sh a ding v ari o u s p or ti o n s Of
.

th e b o dy o f a E u gle na it h a s b e e n f o und th a t th e r e gi o n in f ro nt
o f th e e ye sp o t is m o r e s e n sitiv e th a n a ny o th e r p a r t It Sho ul d .

b e n o t e d th a t wh e n E u gle na i s s wimmi ng th r o ugh th e w a te r it is

thi s a nt e ri or e nd whi ch firs t r e a che s an injuri o u s e nvi ro nm e nt ;
t h e a nim a l s giv e th e a vo iding r e a cti o n a t o nc e a nd are thus c a rr i e d ,

o u t o f d a ng e r .

3 . P L ASMODI U M

( P la smodi u m vi va x Gr a s si)

Th e Sp o r o zo a p a ra sitic P r o to z oa a nd a re r e sp onsibl e f o r
a re

m a ny o f th e mo s t m align a nt a nima l di se a s e s One o f th e b e s t .

kno wn Of all Sp o r o zo o n s i s P la sm odi u m vi vax which ca u se s M al a ,

ri a l F e v e r . Thi s minut e a nim al w a s di s c o ve r e d in th e bl o o d of

m al ari a p a ti e nt s by a Fr e n ch milit ar y d o ctor Lave ra n It w as ,
.

sugge s t e d by this inve s tig a t or in 1 89 1 th a t th e p ara site is p rob

, ,

a bly t ra n s mitt e d f ro m m a n t o m a n by s o m e bl o o d -sucking in se cts ,

a nd this hyp o th e si s w a s p ro v e d t o b e corr e ct by th e w o r k o f M a j or

R o s s in 1 8 99 . N o t o nl y w a s it d e m o n s tr a t e d th a t m al a r i a is
Sp re a d by in s e ct s but it w a s p r o v e d th a t hum a n b e ing s ca n o nly
,

b e com e infe ct e d by th e bit e Of a m os quito b e l o nging t o th e g e nus

Anophe le s . T h e two m os t c o mm o n g e n e r a o f m os quit o e s a r e
C u le x a nd Anophe le s One Of th e e a si e s t m e th o ds Of di s tinguish
.

ing o ne fro m th e o the r i s by Ob se rving the i r p ositi o n whe n a t

re st. It will b e fo und th a t th e har ml e ss C u le x ho ld s i ts abdome n
a pp r o xim a t e ly p ar alle l with t h e s ur f a ce o n which it a light s ,

whe re as th e abdo m e n Of Anophe le s i s h e ld a t a n a ngl e .

Th e re are thr e e w e ll kn o wn typ e s o f m al a ri a ; th e se m ay be

-
 S C HI Z O GO NY

FI G 4 2
. . D i a gr a m ill u s t ra ti ng th e li f e hi s to ry Of th e M a l a r i a l F e ve r paras it e .

s h o w n a bo v e t h e li ne O f d a s h e s a re p a sse d th r o u gh i n th e bl o o d O f a h u m a n b e ing

b e lo w t h e li ne i n t h e b o d y o f a n Anophe le s m o sq u it o
, ,
. F o r e xp la na ti o ns o f t h e
—
( F ro m M i nchi n i n La nk e s t e r s T r e a ti se a f t e r v a r i o us a u th o r s )
’
s e e p a ge s 8 9 9 1 . .
90 AN INTRODU CTION TO ZOOLOGY

c all e d m e lani n (9 a nd 1 0 ,
the s e a re wa ste m a tte rs tha t ha ve
a ccumul a t e d du r ing th e g r o wth p e r i o d b e ing m o difi e d p r o ducts
,

of th e di g e s ti o n o f ha e m ogl o bin .

R e pr oducti on no w t a k e s pl a c e by a p oce ss t e r m e d schi zogony

—
( I V a nd 6 th e p a r a s it e a ft e r it e nt e r s t h e bl oo d c orpuscl e ;
b e ing kno wn a s a schi zont D ur ing this p r o ce ss th e nucl e us
.

divid e s a numb e r o f tim e s witho ut th e inte r ve nti o n o f ce ll wall s

until fro m twe lve to sixte e n da ughte r nu cl e i a r e p re se nt
Ea ch nucl e us with p ar t Of th e s u r r o undi ng cyt o pl asm i s th e n cut
, ,

Off f r o m i ts f e ll o w s r e s ulting in t h e p r o du cti o n o f a numb e r Of

,

sma ll ce ll s call e d m e r ozoi te s incl o se d in th e m e mb ra ne of th e

, ,

c o rpu s cl e s T h e m e l a nin g r a nul e s a r e no t includ e d in th e

d a ught e r c e ll s but re m a in in th e ce nte r o f th e m o the r p a r a
,

s it e ( 9
,
T h e m e r o z oit e s fin a lly b r e a k th ro ugh th e w a ll s o f
t h e co rpu scl e a nd e s ca p e int o t h e bl oo d pl a s m a T h e m e l a nin
.

g ra n ule s a re al so lib e r a t e d a t thi s tim e the y a re ca r

r i e d by th e bl oo d t o t h e liv e r kidn e y spl e e n lung s o r b r a in f r e
, , , , ,

qu e ntly re s ulting in the pigm e nt a ti o n a nd hyp e r trophy o f the se

o rg a n s. An a tt a ck Of f e ve r c o incide s with th e lib e ra ti o n o f the
me r o zoite s a nd th e inte rv al b e twe e n the p e n e t ra tio n o f a bl oo d
,

c orpuscl e a nd th e p r o du cti o n o f m e ro z oit e s i s t e r m e d the p e r i o d

o f in cub a ti o n . Thi s p e ri o d in P lasm odi u m vi vax as s ta te d ab o ve
, ,

i s fo r ty e ight h o u r s
- . All th e m e ro z o it e s Simult a n e o u sly a tt a ck
o th e r c o rpu s cl e s ( 1 0 t o I ) p r o ducing th e chill whi ch is a sympt o m
,

of m al a ri a .

Se ve ra l a s e xu al cy cl e s like th a t ju s t d e s cr ib e d a re p a ss e d
thro ugh ; but fin ally f or s o m e unkno wn r e a so n th e m e ro zoite s
, ,

de v e l op into s e xu a l pha s e s (VI XII) P a r t Of the m n a m e d

— .
,

m acr oga me tocyte s (VI I b ) b e co m e c a p abl e Of p r o ducing a s ingl e

,
.

f e m a le ga m e te or m a cr oga m e te ( X ,
wh i l e t h e re s t b e c o m e

mi cr oga m e tocyte s (VI I ,

e ach c a p a bl e o f p r o ducing as m a ny as

e ight m a le e le m e nts o r m i croga m e te s ( X T h e g am e t o cyt e s

,

c ann o t p r oduce g a m e t e s within th e hum a n b o dy but mus t fi r st

b e s uck e d up int o th e alim e nt a r y ca n al Of a n A nophe le s m o s qui to .

He r e all sta g e s of th e p a r asit e e xc e pt th e g a m e t o cyt e s a r e di

 OTHER P R OT OZ OA 9 1

ge ste d ; the l a tt e r s e e m stimul a te d by the dige stiv e j uice s to f ur

the r de ve l opm e nt .

T h e macr oga m e tocyte no w g oe s th r o ugh th e p r o ce ss o f m a tu r a

ti on ( s e e p . o ne sm all p o l a r b o dy b e ing e xt r ud e d (X ,

wha t re m ains i s a l a rg e m a cr og a m e te T h e nucl e us o f th e mi cr o

.

g a m e tocy t e ( IX a n ),
d .i vid e s int
. o e ight d a ught e r nucl e i which

mig ra te to th e p e riphe r y a nd e nt e r the l o ng p ro to pl asmi c p ro ce sse s

tha t h ave in t h e m e a ntim e b e e n thr u s t ou t fro m the ce ll (X
, , ,

The se flage lli f or m nucl e a t e d p ro ce ss e s the n b r e a k a wa y fro m th e

micro ga m e t o cyt e a s m al e c e ll s mic ro g a m e t e s o r sp e r m a to z o a
, ,

a nd m o v e a b o ut until th e y e nc o unt e r a m a c r o g a m e t e with whi ch

the y fuse (XI) Wh a t r e m al ns o f the mi c r o ga m e t o cyt e sl owly

.

di sinte gr a te s Only o ne mi c r og a m e t e fu se s with a m a crog a m e te

.

in the p r o ce ss o f f e r ti li za ti on .

T h e r e sul t o f thi s fu s i o n i s a zygote which h a s b e e n giv e n th e

n a m e oole i ne t (XII) Thi s zyg o t e b e co m e s spin dle sh ap e d (XIII)
'

.
-

a nd m a k e s its w a by v e r mif o m m v e m e nt s t o t h e e pith e li a l c e ll s

y r o

lining the alim e nt a ry c a n al o f th e m os quit o ; it p e ne t r a t e s the se

a nd r e a ch e s t h e und e r lying ti s s u e s H e r e it g r o w s r a pidly ;
.

th e nucl e u s divid e s f o r ming m a ny d a ught e r nu cl e i a nd by t h e

, ,

sixth d ay as m a ny c e ll s a r e p r o duce d a s th e re a r e nucl e i (X I V

X VI ) . Ea ch c e ll i s a s por obla st (X VI S p T h e sp o ro bl a s t
,
.

for m s by divi si o n a numb e r o f g e r m s the spor ozoi te s (X VII ,

XVIII) ; the se m a tu re in a b o ut f o u r t e e n d a y s a nd a re the n se t

f e int o th e b o dy c avity Of the m os quito (X VIII XIX ) F r o m
r e — .

he r e the y a r e ca rr i e d by th e ci r cul a ting pl a sm a to all p ar ts Of th e

b ody fin ally c oll e cting in th e a nt e ri o r r e gi o n The y m a ke th e i r
, .

wa y int o th e s a liv a r y gl a nd s a nd a r e th e n r e a dy to p a ss int o th e

,

bl oo d o f th e hum a n b e ing a t th e time th e m os quito m a k e s i ts

ne xt m e al .

M al a r i a l fe v e r is no t p r e s e nt wh e r e v e r Anophe le s is fo und ; f o r
e x a mpl e Ano he le s i s c o mm o n in E ngl a nd but no m a l a r i a o c cu r s
, p ,

the re . It i s s upp o se d th a t m os quito e s m ay b e c o me immun e to

.
a ll kind s o f bl o o d p a r a sit e s A m o s quit o th a t i s a bl e t o di g e s t
.

th e p ar as i te b e c o m e s h a r m l e s s t o m a n T h e m osquit oe s o f t h e
.
9 2 AN INTRODU CTION TO ZOOLOGY

ge nu s Cu le x are abl e to dig e s t all s t age s Of P la smodi u m vi vax and ,

a r e th e r e f o r e immun e On th e o th e r h a nd thi s m os quit o doe s

.
,

no t di g e s t a ll o f t h e o r g a ni s m s o f th e m a l a r i a which a tt a cks bi r d s
a nd thu s b e c o m e s inf e ct e d a nd is th e m e a n s o f t r a n s mitti ng th e

fe v e r fr o m one bir d to
a n o th e r .

Q uinin e is th e re m
e d y c o mm o nly use d
a g a in s t t h e m al a r i a l
p ar a sit e It a ct s di .

r e c t ly u p o n the
y o unge r st age s of the
organism ca using ,

d e a th .

4.VOLV OX G LOB ATOR

AN D I T S AL LI E S

C e r t a in or g a ni sm s
w h i c h li e o n t h e
b o r d e rl and b e twe e n
pl a nt s a nd a ni m al s are
p e culi a rly f avo r abl e
f o r illu s t r a ting h ow a
FI G 4 3 Chla mydomonas A, i ngl e v e ge a
s t tiv e
.

ce
.

ll : a, s ti gm a ; chr
.

ch r o m a t o p h o r ; g fl
m a
u lti ce llul a r a ni m al
,
.
,
.

llu m ; k , nu cl e u s ; py , pyre no i d ; v va c
hke ( 8
F 1 °
4 6)
ge . . .
,

u o le s B , F o ur d a u ght e r ce ll s s till
. h av e e v olv e d withi n m a y
th ll w ll f th p nt ; C th
e ce a o e a singl e -c e ll e d
a re ,
e u ni o n f r o m

Of t w g m t ; D
o yg t a ( F m Olt a nc e s t o r
e es ,
z
The se o r
o e . ro
.

m nn ) a s
ni m s b e l o ng t o t h e
'

ga s

F a m i ly Vo lvo ca c e ae The y a re imp or t a nt no t o nly f o r the i r phy

.

loge ne ti c sig nific a n ce but a l so b e c a u se th e y illu s t ra t e th e e volu

ti on of s e x St a r ting with th e uni ce llul ar C hla mydom ona s the
. ,

line Of e v o luti o n p a sse s th ro ugh S pondylom oru m P and or i na , ,

Eud or i na a nd fin a lly e nd s with Volvox

,
94 AN INT RODU CTION TO Z OOLOGY

of all th e o the r s . R e p r o ducti o n like wis e t ak e s pl ace in ea ch

c e ll .

SO f ar kn o wn r e pr odu cti on of onl y o ne kind o ccu rs ; th i s i s

as

th e Sim u lt a n e o u s l o ngitudin a l divi si o n o f e a ch c e ll int o t w o f o u r , ,

e ight a nd fin a lly s ixt e e n d a ught e r c e ll s

,
Th e s e d a ughte r c e ll s
.

d o no t s e p a r a t e a s d o t h e da ught e r ce ll s wh e n C hla mydomonas

di vide s but r e m ain f as te ne d t og e the r by a ge l a tin o us m a trix
, .

E a ch c e ll o f th e m o th e r col o ny thu s p ro duce s a d a ught e r c ol o ny

o f sixt e e n c e ll s If a n o the r kind o f r e p ro du cti o n d oe s t a ke pl a c e
.
,

it is p rob ably by th e u ni o n Of Simil ar ga me te s f ormi ng a zygo te

simil ar to tha t of C hlamydom onas .

P and ori na m oru m (Fi g 4 5) i s lik e wi se a c o l o ni al for m f o und in

.

fre s h-w a te r p onds It co nsi s t s Of Sixte e n c e ll s which are h e ld

.

toge the r by a ge l a tin o u s m a trix p a r t of whi ch is s e cre te d by e ach

,

ce ll ( I) E a ch c e ll p o ss e ss e s o ne p ai r o f fl ag e ll a a nd a n e ye sp o t
.
,

a nd c o nt a in s chlo r ophyll It is thus e n a bl e d t o c arr y o n all Of th e

.

p ro c e sse s n e c e ss a r y t o sust a in life to g ro w a nd to r e p roduce , ,

.

R e pr odu cti on in thi s sp e ci e s t a k e s pl a ce in t w o w a ys F i r st .

,

a s in S pond ylom or u m e a ch ce ll m a y divid e t o f or m t w o f o u r

, , ,

e ight a nd th e n sixt e e n d a ught e r ce ll s whi ch b e c o m e a ne w c o l o ny

, ,

( g 5
F i 4 . II ) T h e
,
n e w co
. l o ni e s e s c a p e f ro m t h e m o th e r c o l o ny
by the di ssoluti o n o f th e g e l a tin o us e nve l op e a nd swim a w ay to ,

l e a d a s e p ara te e xi s te nce Thi s m e tho d o f r e p ro ductio n o ccu rs

.

r e p e a t e dl y but fin ally con j g ti n i in ugu t d E ch f t h

‘

, u a o s a r a e a o e .

sixt e e n c e lls o f th e m o the r c ol o ny p r o duce s by di vi si o n Sixte e n or

thi r ty tw o d a ught e r ce ll s E a ch d a ughte r ce ll de ve l op s fl ag e ll a
- .

a nd se p a r a t e s no t o nly f ro m th e m o th e r c ol o ny but f r o m i t s si s t e r ,

c e ll s in th e d aught e r col o ny ( III ) Thi s se p a r a ti o n r e s ults fro m

.

a soluti o n o f no t o nly th e g e l a tin o u s e nv e l op e o f t h e m o th e r c ol o ny

but al so tha t whi ch h old s th e da ught e r c e ll s t o ge the r The se .

i sol a te d ce lls are kno wn a s ga me te s T he g am e t e s a re not all o f .

t h e s a me size s o m e b e ing l ar g e r t ha n o th e r s Wh e n tw o g a m e t e s
,
.

m e e t the y b e com e fa ste n e d t oge th e r by the i r a nt e ri o r e nds a nd

,

gr adually fus e ( I V—V) T h e fuse d ce ll s c o n stitute a zygote co n

.
,

t ai ni ng t wo e ye sp o ts a nd b e a ring fo u r fl ag e ll a (V) T h e fl age ll a .

 OT HER P R OT O OA Z 95

a re s oo n ca st Off a nd the zyg o te sink s to the b o tto m Of the p o nd

whe re it s e c re te s a thi ck re d w a ll a b o ut its e lf (VI VI I ) It re
— .

m a ins in t hi s co nditi o n thr o ugho ut th e summ e r be co nu ng d r y ,

whe n the p ool o f wa te r in which it live s e vap ora te s In th e .

a utumn a ctivity is a g ain r e s um e d T h e zyg o t e p ro du ce s f ro m

.

FI G 4 5 I, ve ge ta tive l
co o ny ; I I ,
f o r m a ti o n o f d a u ght er

l
co o ni e s ; III , e s ca pe of ga m e t e s ; I V, V, VI , VI I , f u s i o n o f t wo
ga m e te s to f o r m a z ygo t e ; VIII I X p ro d u cti o n Of s w a rm spo r e s by a
, ,

z ygo t e
; X v e ge ta tiv e co l o ny f o r m e d by a s w a r m spore
, ( F r o m Olt .

manns ) .
9 6 I
AN I N T R OD U C T ON T O Z OOLOGY

o ne th re e swar m spor e s which e sc a p e f r om the cy s t a nd swim

to
a w a y ( VIII IX ) E a ch s w a r m sp o re d e ve l ops at o n ce by bin ar y
— .

di vi si o n a ne w c ol o ny o f sixte e n c e ll s (X) .

T h e g a m e t e s Of P a nd or i na a re no t e qu a l in siz e as in C hla my
dom ona s but a pp e a r to b e o f t w o kind s s o m e l a r g e r th a n o the rs
, , .

Wh e re s uch a co ndi ti o n e xi s ts the s m all e r ga m e te s a r e c all e d

m ale ce lls o r s pe r ms t he l a r ge r f e m ale ce lls o r e ggs
, , S e xu a l .

r e pr odu cti on c o n si s ts in t h e uni o n o f a m a l e c e ll with a f e m a l e

ce ll ,
a nd ce r t ai n inv e s tig a t o r s b e li e v e th e l a rg e r g a me te s o f
P a nd or i na t e nd to fu s e with t h e s m a ll e r If this is t r u e w e h a ve
.
,

in thi s o rga m sm th e b e ginning o f th e e voluti o n Of s e xual r e p ro

du cti on .

In Eu d or ina e l e gans a n o the r m e mb e r Of the fa mily Vo lvo

,

ca ce ae thi s di s tincti o n b e tw e e n m a l e a nd f e m a l e g a m e t e s i s m o r e
,

cl e a rly se e n E u d or i na is a sphe r ic al c ol o ny c o nt aini ng thi r ty

.

t w o r a r e ly Sixt e e n ce ll s w hi ch r e s e mbl e th e c e ll s Of P a nd ori na

,
.

R e pr odu cti on by s impl e divi si o n t a k e s pl a ce a s in P a nd or i na .

Thi s ca nn o t go o n inde finit e ly f o r fin ally s o m e col oni e s a re

,

f o und whose c e ll s h a ve g r o wn l a r g e r th a n u su al The se a re .

f e m al e col o ni e s a nd th e c e ll s a re m acr oga me te s Oth e r c ol o ni e s

,
.

p ro duce micro g a m e t e s ; e a ch o f th e thi r ty two c e ll s o f th e se -

m a l e co l o ni e s divid e s p r o du cing a fla t pl a t e o f sixt e e n o r thi r ty

,

t w o spindl e Sh a p e d m i cr oga m e te s e a ch with a p a i r o f fl a g e ll a

-

e xt e nding f r o m i t s a nt e r i o r e nd Wh e n a pl a t e Of mic rog a m e t e s

.

e nc o un t e r s a c o l o ny o f m a c r o g a m e t e s it b e co m e s a tt a ch e d t o

it th e mic ro g a m e t e s s e p a r a t e f r o m th e pl a t e a nd m a k e th e i r
,

wa y th r o ugh th e g e l a tin o u s e nv e l o p e s o f th e m a cr og a m e t e s ,

thus f e rti li zi ng the m a nd fo r ming zygote s .

It sho uld b e n o t e d th a t the o rg a ni z a ti o n of E ud or i na is m ore

co mpl e x tha n th a t of th e o the r thr e e sp e ci e s co n side r e d a nd tha t ,

thi s a dv a nce in co mpl e xity i s g r a du a l In Eu d or i na al so the

.
, ,

s m a ll mic ro g a m e t e s a re p e r f e ctly di s tin ct f ro m t h e lar ge m a c ro

'

g a m e t e s a nd fe r tili z a ti on a lwa ys con si s t s in th e uni o n o f a m a cro

,

with a mi cro ga m e te This Size di ff e re nc e i s no t SO e vid e nt in

.

P a nd or i na .
FI G .
46 . Volvox globa tor . B, a co lo ny r e pr o d uc i ng by m e a ns o f e ggs a nd

sp e r m s : o .
,
e ggs ; s ,
. spe r m s .
( F r o m La ng )
. C, a s i n g l e ce ll s h o wi n g
t l
p ro o p a s m ci ti o ns pl
co nne c ,
.
,
ce ll p r o t o pl a s m ; g . j
,
e ll y; m . li,
ne s

s h o wi ng li m it s Of ce ll e nv e l o p e .
( F r o m O lt m a n ns a f t e r v a r i o u s a u th o r s ) .
9 8 AN INT R ODU C TION TO Z OOLOGY

of th e l a rg e r ce ll s with o ut fl ag e ll a So m e Of the se gr o w l arge r and

.

m ay b e r e co gni z e d a s f e ma le ce lls o r m acr oga me te s o th e r s

p ro duce by simpl e divi sio n a fla t pl a t co nt a ining ab o ut o ne hun ,

dr e d a nd twe nty e ight spindl e sh ap e d m a le ce lls or mi cr oga m e te s

- -

(s) The se e sca p e into th e ce nt r a l c a vity a nd fu se with the

.

m a cr oga m e te s T h e zygote thu s fo r m e d se c r e t e s a s ur ro unding

.

wall co n si s ting o f t w o l a ye rs th e o ute r Of whi ch i s re ddi sh in col or

,

a nd co v e r e d with s h o r t spin e s In thi s co nditio n th e winte r is

.

p asse d T h e foll o wing sp r ing th e zygo t e b r e a ks o u t Of th e w all

.

a nd by divi s i o n p r o duc e s a ne w co l o ny T h e sm a ll e r so m a tic

.

c e ll s co nt a in e d in th e m o th e r co l o ny f a ll t o th e b o tt o m a nd di si m
t e gra te a s soo n as th e ne w co l o ni e s p ro du ce d by th e f e r tiliz e d
g e r m c e ll s h a ve e s c ap e d .

In Volvox tr u e s om a ti c ce lls a re e nco unt e re d f or th e fi rs t tim e

, ,

tha t i s c e ll s which functi o n o nl y ve g e ta tive ly a nd a re un abl e to

,

r e p r o duc e t h e c o l o ny In the o the r fo r m s de s c rib e d e ve ry ce ll

.

h a s th e c a p a city o f r e p r o du cing t h e wh o l e Volvox a l so co nt a in s .

tr u e ge r m ce lls th a t i s c e ll s th a t h a v e giv e n up nut ritiv e functi o n s

, ,

t o ca rr y o n r e p r o du cti o n F u r th e r m ore a C l e a r c a se o f na tu r a l
.
,
'

de a th o ccu r s in t h e so m a t i c ce lls wh e n th e y f a ll to th e b o tt o m o f
th e p o nd a nd di sint e g r a t e T h e b o di e s o f high e r a nim als co n sist
.

o f m a ny ce ll s whi ch m a y b e se p a r a t e d int o so m a tic a nd g e r m c e lls .

T h e l a tt e r a r e e ith e r m a l e o r f e m a l e In m o s t c a se s a fusi o n o f a
.

m al e ce ll with a fe m al e ce ll i s n e c e s sa ry b e fo r e a ne w a nim al ca n
b e r e p ro du ce d . At a ny r a t e s o m e o f th e se g e r m ce ll s m a int a in
,

th e co ntinu a ti o n o f th e s p e ci e s by p r o du cing ne w indi vidu als

whi l e th e so m a tic ce ll s p e ri sh wh e n t h e a nim al di e s .

Th e G e r m P l a s m Th e or y Figu r e 4 7 illu s t r a t e s t h e th e o ry
.

o f t h e co ntinu ity o f t h e g e r m -pl a s m whi ch i s h e ld by m os t z oolo

gi s t s a t t h e p r e se nt tim e B y g e r m —pl asm i s m e a nt tha t p a r t Of

.

t h e p ro t o pl a s m which i s s e t a sid e f o r r e p r o ductiv e pu rp os e s a nd

de t e r min e s th a t th e Offsp r ing sh a ll r e se mbl e the i r p ar e nt s ;
thi s sp e ci al m a t e ri a l i s s tor e d in th e ge r m c e ll s In th e sp ring .

o f th e y e a r th e Volvox r a ce i s r e p r e s e nt e d by fe r tiliz e d e ggs

‘

( zyg o t e s) o n lv : e a ch o f th e s e divid e s p r o du cing a n a nim a

,
l con
 OT HER P R OT OZ OA

t aining as m a ny a s tw e lve tho usa nd c e lls a f e w of whi ch a re ,

p
re ro ductiv e c e lls a nd c o nt a in t h e g e r m -pl as m In th e a utumn .

t h e g e r m -pl a sm is se g re g a t e d in th e e gg s a nd sp e r m a to zoa which ,

fuse two by two an e gg with a sp e r m p ro ducing zygo t e s T h e

, ,
.

s o m a tic o r b o dy c e ll s f a ll to th e b o tt o m a nd d i e but t he zygo t e s ,

live thro ugh t h e wint e r a nd ge r min a t e in th e sp r ing thus a ssu ring ,

t he c o ntinu e d e xist e nce o f th e r a c e .

0 no .

F I G 4 7 D i a gra m to ill us t r a t e th e th e o ry o f th e co nti nu it y o f th e ge rm -p l as m

. .

by m e a ns o f Volvox T h e f e r tili z e d e gg o r z ygo t e

.
p r o d uce s by d ivi
s i o n i n t h e s pr i ng a co l o ny (c) co nt a i ni ng a gre a t m a ny ce ll s , so m e O f

whi ch a re se t as i de fo r re tiv
pro d u c e p ur po s e s T . h e se pro d uce ne w co ol
ni e s d ur i ng th e s u m m e r by t h e a se xu a m e l tho d
ll s ( spcr . M al e ce

m a to z o a s p) a nd f e m a l e ce ll s ( e ggs e ) a r e
, ,
fo r m e d a l so
I n th e a u t u m n .

th e b o d y d i e s b u t t h e f e r tili z e d ge r m ce ll s (22) pro d uce d b y th e u ni o n o f

,

e ggs a nd spe r m a t o z oa ( ) s ur viv e t h e wi nt e r d ivi d i ng to f o r m ne w co lO

f ,
~

ni e s the ne xt spr i ng .

C e r t a inp ro ce ss e s in the de ve l opme nt Of the fe r tilize d e gg s o f

s ome o f the m ore co mpl e x a nim a l s s t r e ngthe n t he b e lie f in the
p ass age o f th e ge r m -pl a sm fro m o ne ge n e ra tio n t o a n o the r a nd ,

se e m to in di c a t e th a t t h e b o dy i s no t th e p r o duc e r o f t h e g e r m

ce lls but i s simply a ve hicl e which c arr i e s th e m a b o ut until the y

,

r e a ch m a tur ity T he b o dy the n die s a nd di sint e gr a t e s the

.
,

imp or t a nt functi o n o f r e p r o ducti o n a nd c on se qu e ntly th e s u r,

vi v al o f th e r a c e b e ing l e ft in ch a r ge o f t h e m a tu r e ge r m c e lls
,
.
 C HAP T ER VI I

AN INTR ODU CTI ON T O THE M ETAZ OA

1 . C E LLU LAR DI F F E R E N TI ATI ON TI SSUE S

THE R E i s no sh a r p lin e b e twe e n the M e t a zo a a nd P r o to zoa .

We h a ve s e e n ( C h ap VI ) th a t th e Vo lvo c a c e ae which a r e Ofte n

.
,

include d in t h e P hylum P r o t o zoa a re m a ny ce ll e d co l o nie s the

,
-
,

ce ll s Of which are e ithe r e nti r e ly ind e p e nd e nt a s in S pondy ,

lomor u m ( Fi g . o r a r e c o nn e ct e d by p r o t o pl as mi c s t r a nd s a s ,

in Volvox ( Fi g . He re th e c e ll s o f t h e c o l o ny h a ve ce a se d t o
b e ind e p e nd e nt but h a ve unit e d t o f o r m a di s tin ct b o dy whi ch
,

c a rri e s o n nut r itiv e p ro ce sse s p ro duc e s ge r m ce ll s a nd th e n di e s

, ,
.

T h e c e ll s Of thi s b o dy a re c a ll e d so m a tic c e ll s The ir functio ns

.

a re l o co m o t or a nd nut ritive but not re p r o ductive Such a n

,
.

a gg r e g a ti o n o f ce ll s i s kn o wn a s a ti s s u e A ti ssu e i s a n a sso ci a
.

tio n o f Simil ar ce ll s o r igina ting f r o m a p a r ticul a r p a r t o f th e

e mb r y o a nd w ith sp e ci a l functi o n s t o p e r for m In Volvox the r e
.

is o nly o ne kind o f ti ss ue ; in so m e Of the simpl e M e t az oa the re

a r e t w o kind s o f ti s s u e r e s ulting f r o m t h e di ff e r e nti a ti o n o f t h e

so m a tic ce ll s ; in t h e m a j or ity h o w e v e r th e r e a r e a t l e as t thr e e

, ,

a nd u s u a lly m or e T he m a ny di ff e re nt kinds o f ti s s ue in t he
.

M e t a zo a m ay b e cl assifi e d a cc or ding t o th e i r st r uctur e a nd

functi o n s int o fo ur g ro up s .

( 1 ) Epith e li al ti s s u e ( Fi g 4 8 A B ) c o n sis ts o f c e ll s which c o ve r

i
.
,

a ll th e s u r f a ce s o f th e b o dy b o th with o ut a nd wit hi n In the .

simpl e r a nim a l s thi s i s the o nly kind o f ti ss u e p r e se nt Th e .

so m a tic ce ll s o f Volvox m ay b e co n side r e d e pithe lium In the .

m o r e co mpl e x a nim al s e pithe li a l ce ll s b e co m e va r i o u sly m o difie d

b e c a u se the y are the m e an s o f communic a ti o n b e twe e n t he or
g a n i s m a nd its e nvi ro nm e nt ; nut r itiv e m a t e r i a l p a ss e s t hr o ugh
1 00

o r 0 0 0 0
 AN INT RODU C TION T O T H E M ET A OA Z I Ol

the m into the b ody a nd e xc re tor y p r o duct s p ass th ro ugh the m on

,

the i r w ay ou t o f the b o dy ; the y a ls o co nta in th e e nd o rga ns o f

th e se ns o r y a pp a r a tus a nd p r o t e ct th e b o dy f ro m phy s ic a l c o nt a ct
with the o uts ide world In m a n the cuticl e a nd lining o f th e
.

a lim e nt a r y c a n a l a r e e x a mpl e s Of e pith e lium .

( 2) S u ppor ti ng and C onn e ctive Ti s s u e s ( Fi g 4 8 C ) m ay be e n .

c o unte r e d i n alm os t a ny p a r t o f t he b o dy The i r chie f functio n s

.

a r e t w o in numb e r ; ( 1 ) th e y bind t o g e th e r v ar io u s p a r t s o f the

b o dy a nd ( 2) the y for m r igid st r ucture s c ap abl e Of r e si s ting

,

sho cks a nd p re ss ure s o f all kinds Th e se tissue s c on sist l a rge ly

.

o f no n living s ubst a nc e s fib e r s pl a t e s a nd m a sse s p r o duc e d by

-
, , ,

t h e c e lls e ithe r wit hi n t h e c e ll w a ll o r o ut s ide Of it

-
T h e t e nd o n s .

which unite muscl e s t o b o n e s a nd the b o n e s the ms e lve s illus

, ,

t ra t e the two ki nds Of ti ss ue s i n thi s g ro up .

(3 ) M u s cular ti s s u e s ( Fi g 4 8 D E) are the a ge nts o f a ctive

.
,

m o ve m e nt We fo und ( C h a p I V) tha t the l o co m o tio n o f Ame ba

. .

c o uld be e xpl a in e d by the p r e se nc e o f co nt r a ctil e fibe rs In o the r .

P r o t o zo o n s b e l o nging t o t h e C l a sse s I nfusor i a a nd Sp o ro z oa

the r e are muscul ar fib r ils c a ll e d m yo ne m e s in th e m e mb ra n o us
c o v e rings In m o st o f th e hi ghe r org a nis ms sp e cia l muscl e c e lls
.

a r e di ff e r e nti a t e d f or p e r for ming t h e v a r i o u s m o v e m e nt s o f t h e

b o dy The se c e lls p o sse s s muscl e fib rils whi ch are a bl e to co n

.

t ract with gre a t for c e a nd in quick s ucce ssio n The fib rils a re .

us ually of two kinds ; ( a ) c r oss s t ri a t e d ( D) a nd ( b) s mooth no n

st ri a te d ( E) T h e l a tt e r fo r m a l e ss highly de ve l ope d ti ss ue tha n
.

th e f o r m e r a nd a re fo und in th e Simpl e r in a ctive a nim a ls a nd ,

in th o se inte r n al org a n s of highe r o rg ani s ms no t subj e ct to the

will of the a ni m al .

(4) N e rvo u s ti s s u e i s c omp o se d of ce ll s whi ch a r e SO a ct e d

upo n by e xt e r n al phy sic al a nd che mic al a ge nts th a t the y a re
a bl e t o p e r c e iv e a s timulus t o c o nduct it t o s o m e o th e r c e ll o r
,

ce ll s of the b ody a nd to stimul a te still o the r c e lls to a ctivity

,
.

All p r o t opl a sm i s i r r i table ; a ni m a l s with o ut n e r v o us sy s t e ms ,

e g A me bw a r e ca p a bl e o f r e a cting t o a s timul us but in m or e

. .
, ,

c omple x org a ni sm s ce r t ain ce ll s a re spe ci alize d f or the sol e pur

1 02 AN INTR ODU C TION TO Z OOLOGY

p o s e of p e r fo r ming th e functi o ns de scrib e d ab ove as chara cte ris tic

o f n e r v o u s tissue .

T HE GE R M C E LL S
’

2 .

We ha ve se e n th a t th e sub sta nce o f whi ch all a nim a ls a re com

pose d m ay b e s e p a ra te d int o re p r o ductive a d no n re p ro ductive
n - .

T h e l a tt e r i s call e d s o m i p
a t c las- m t h e f m g
or,
e r e r m - l
p m NO
as .

o rg a ni s m a r i se s fully de v e l o p e d but u s u a lly o r igin a t e s f r o m a

,

union o f two ge r m c e ll s which h ave b e e n p ro duc e d by a p r e e xi s t

'

ing orga ni sm The r e is a g r a dual de ve l o pme nt of an individua l

.

fro m the fe r tilize d g e r m c e ll th e org a ni sm p a ssing thr o ugh a

,

numb e r o f we ll d e fi ne d s t ag e s b e f or e th e a dult c o nditio n is at

-

t a i ne d . T o b r ing th e v ar i o u s s t e p s o f t hi s p ro c e ss cl e ar ly b e fo r e
u s w e s h a ll no w di s cu ss t h e o r igin a nd r ip e n i ng o f th e g e r m c e lls ,

the i r unio n in fe r tiliza ti o n a nd th e s ub se que nt de v e l opm e nt al

,

his t ory of the zyg o te thu s f or m e d Th e se s ubj e ct s will b e co n

.

s i d e r e d in so m e d e t a il f or e a ch typ e s tudi e d in t h e f o ll o win ch a p

g
t e rs SO w e sh all he re giv e o nly a g e n e ral o utlin e which will s e r ve
,

a s a f o und a ti o n f o r futu r e discu ss i o n s .

E a r ly in t h e hi s t o r y o f th e individu a l c e r t a in c e ll s a re s e t a sid e
f o r th e s ol e pu rp ose o f r e p r o ducti o n ; th e se ar e t h e ge r m ce lls .

All th e o th e r c e ll s o f t h e d e v e l oping or g a ni s m b e c o m e m or e or
l e ss sp e ci aliz e d f or the v a r i o u s v e g e t a tive functi o n s a nd m ay ,

e v e ntu a lly b e r e c o gniz e d a s n e r v e c e ll s mus cl e ce ll s e t c

,
Only
,
.

th e g e r m ce lls r e m a in in th e i r p r imitiv e co nditi o n a nd sinc e , ,

the y t a k e no p ar t in th e da ily life Of th e a nim al w e m ay think o f ,

the m a s b e ing p a ssive ly ca rr i e d ab o ut a nd p ro t e cte d by the so m a

tic ce ll s Ju s t b e f o r e th e individu al b e co m e s se xu ally m a ture the
.
,

g e r m ce ll s a wa ke t o a ctivity a nd a numb e r Of co mpl e x p ro c e sse s

a r e in a ugu r a t e d w hi ch r e s ult in t h e c a s ting o u t o f c e r t a in Of th e m

by th e b od y t o p r o duce ne w individu als Sim il ar t o th e p are nt

o rg ani s m .

Spe r m a toge n e s i s . T h e d e v e l o pm e nt o f th e m al e g e r m c e ll or
sp e r m a t ozo o n i s t e r m e d sp e r m a t og e n e si s As sho wn in Figure 4 9
.
,

thi s p ro c e s s m ay b e di vide d int o thr e e p e r io ds : ( 1 ) the multiplica

1 04 AN INTRODU CTION TO ZOOLOGY

divide e a ch for ming tw o spe r matids E a ch Sp e r ma tid r e ce iv es

, .

o ne h a lf o f e a ch chr o m o s o m e fr o m t h e s e c o nd ar y sp e r m a t o cyt e

th a t giv e s r i s e t o it T h e sp e r m a tids a re the n m e t a m orph ose d

.

int o spe r m atozoa .

T h e s pe r m a tozoa Of v a r i o u s a ni m al s a r e u su ally e a sily di s

t i ngui sh e d o ne fr o m a n o th e r but a r e m o s tly c o nst r uct e d o n th e
,

R M R AL
P I O DI
G RM
E '
OE LL

M U LT L A T
IP IC ION
R D
P E IO

OOGONIA

MAT URKTTJON
£EB IOD

FI G 5 0 D i a gr a m ill u s t r a ti ng t h e s t a ge s o f oo ge ne s i s
. . T he pr i mo rd i a l
.

g e r m ce ll i s re p r e se nt e d a s posse ss i ng f o u r ch ro m oso m e s .

s am e pl a n The y r e se mbl e a n e l o ng a t e d t a dp ol e h a ving a he a d

.
,

fill e d alm o st e nti re ly with nucl e ar m a te r i al a nd a l o ng flage llu m

like t ail which i s th e o rg a n Of l o co mo ti o n ; th e middl e pi e c e
,

j o ining the s e two i s th e c e nt ro so m e Th e sp e rm a to zo a a re .

th e a ctiv e g e r m c e lls ; it i s th e i r duty t o s e e k o u t a nd fe r tilize

th e l ar g e r st a ti o n ary e gg c e ll s F r e qu e ntly th e y are o nly
.

“ 1
17 76 50
7 50 t h
“
e siz e o f t h e e gg a n d in th e ,
s e a u r chin T oxopne u s te s , ,

the i r bulk i s ab o ut 3 6 3 35 0 th e v olum e Of th e o vum

-
5
-

Oo ge n e s i s T h e o r igin o f th e e gg i s c all e d o oge n e si s ( Fi g

. .

St a ge s ar e p a s se d th r o ugh by th e ge r m c e ll s co r r e sp o nding al
m os t e x a ctly t o t h ose d e scr ib e d und e r sp e r m a t o ge n e si s ( Fi g .

B e fo r e th e g r o wth p e r i o d th e g e r m c e ll s whi ch wi ll p r o duc e e gg s

a re kn o wn a s oogoni a ( Fi g 5 0 ; Fi g 5 1 . a) At t h e co mpl e ti on
.
,
.
 AN INT R ODU CTION T O T HE M ET AZ OA 1 05

of th egr owth p e ri o d the y a re t e r m e d pr i mary oocyte s ( Fi g 5 1 b) . .

T h e p r im a r y oo cyt e s c o nta in o nly o ne h a lf t h e numb e r of ch r o m o

s o m e s chara cte ri s tic of th e s o m a tic ce lls a nd oogoni a As in .

th e p r im ar y sp e r m a t o cyt e s th e s e ch ro m o s o m e s a r e bi vale nt
,

FI G 5 1 D i a gr a m s ill u s t r a ti ng th e m a t ur a ti o n f e r tiliz a tio n a nd cl e a v a e

.

g
.

, ,

Of a n e gg T h e pr i m o r d i a l ge r m ce ll i s r e r e se nt e d as o sse ss i n f ou r
p p g
.

chr o m o so m e s.
1 06 AN INTRODU CTION TO Z OOLOGY

re sulting f ro m th e unio n two by two o f th e unival e nt chr o m o s ome s

Of th e o og o ni a T h e p r im a r y oo cyt e di vid e s in th e f oll o wing
.

m anne r Its nucl e us c all e d th e ge r mi nal ve si cle ( Fi g 5 1 a)

.
,
.
, ,

m ov e s to th e p e r iphe r y ( b) whe re a mito tic figur e is for me d pe r

p e ndi c u lar t o t h e su r f a c e o f t h e e gg ( c) A sm a ll
. bud lik e p r o t ru -

si o n i s now f or m e d into whi ch p ass one unival e nt chro m os om e

fro m e a ch of th e bival e nt chro m os om e s p r e se nt in the p ri m ary
o o cyt e (d) T h e bud i s th e n pinche d Off
. T wo s e cond a ry oocyte s
.

a r e p r o du ce d by this divi s i o n e a ch c o nt a ining a n e qu a l a m o unt o f

,

ch ro m a tin but o ne with a gr e a t d e al m ore cyt opl asm and yolk

,

tha n t he o the r (e ) T h e sm all one is kn o wn as the fir st polar

.

body ( e p b I ) a nd is no t functi o n al ; t h e l a r g e r is th e e gg E a ch
,
. . .

s e co ndar y o o cyt e now p r e p ar e s f or di visi o n (f ) T h e fi rst p ol ar .

b o dy in s ome c as e s doe s no t divide ; whe n it do e s the di vi si o n is

e qu a l (g p b ,
T h e e gg th ro ws Off a s e cond pola r body (g
. .
,

p . b .2 ) which co nt a in s o n e h a lf o f e a ch c h r o m o s o m e Th i s .

se c o nd p ol ar b o dy di s int e g ra t e s a s d oe s th e fi r st ,
.

F e r tiliz a ti on T h e m a tu r e o vum now b e c o m e s th e c e nt e r o f

.

th e inte r e s ting p ro ce s s o f fe r tiliza ti o n T h e spe r m a to zo o n .

s o m e tim e s e nt e r s th e e gg b e f or e t h e p o l a r b o di e s a r e f or m e d a nd ,

so m e tim e s a ft e r w a r d In t h e illu s t r a ti o ns ( Fi g 5 1 f ) we have

. .
,

sho wn the sp e r m e nte ring the e gg a t th e e nd o f th e fi r s t oo cyte

divi si o n T h e sp e r m b rings int o th e e gg a nucl e us a c e nt ro
.
,

s o m e a nd a v e r y s m all a m o unt o f cytopl as m T h e sp e r m nucl e us

, .

s oo n g ro ws l ar g e r by th e a bs orpti o n Of m a t e ri al f r o m the cyto

pl a sm of th e e gg a nd th e ce nt roso m e b e gin s i ts a ctivity A mi
, .

t o ti c figu r e soo n g ro ws up (g) a nd m o v e s t o w a r d th e c e nt e r Of t h e

e gg . T h e e gg nucl e u s a l so m ov e s in thi s di r e cti o n ( h) a nd fin a lly ,

b o th th e m al e and fe m al e nu cl e i ar e b r ought t oge the r in th e mids t

o f t h e Spindl e p r o duc e d a b o ut t h e s p e r m nucl e u s ( i ) Thi s co m .

pl e te s th e p ro ce ss usu ally kn o wn a s fe rtiliz a ti o n In this p ro ce ss .

t h e chi e f a i m S O f a r s e e m s t o b e the u ni on of two nu cle i one of ,

ma te r na l or i gi n the othe r of pa te r nal or i gi n We s h all se e l a t e r

,
.

th a t fe r tiliza tio n i s r e ally no t co n s umma te d until t h e a nim al

which de ve l o ps fro m th e e gg h a s b e co m e s e xu ally m a ture .
1 08 AN INT RODU CT ION TO Z OOLOGY

mate r nal ori gi n . F u r th e r m it o tic di vi sio ns insu re a like di s tri bu

ti o n to e ve r y ce ll in the b o dy . Aft e r nu cl e a r divisi o n c om e s th e
di vi sio n o f the e nti r e ce lls into tw o (13 a nd l) .

Typically th e f e r tiliz e d e gg divide s into two ce lls the s e tw o ,

into fo ur the s e fo ur into e ight e t c e ach cl e a va g e pl a n e b e ing

, ,
.
,

p e rp e ndi cul ar to the l a st p r e ce ding pl a ne ( Fi g This i s .

kno wn a s total cle avage a nd i s Chara ct e ristic Of holoblasti c e ggs

,
.

Oth e r e ggs a r e said t o b e m e r oblas ti c b e c a u se th e y e xhi bit pa r ti a l

cle a va ge th a t is o nly a s m a ll p a r t o f th e e gg e nt e r s int o c e ll divi
, ,

s i o n th e r e m a inde r se r ving a s nut r itiv e m a t e r i a l f o r t h e cl e a v a g e

,

ce lls In all w e can r e co gnize fo ur di s tinct typ e s of cl e av ag e :

.

( ) q
1 e u a l cle a va ge wh e r e
, t h e e gg di vid e s int o t w o e qu a l h a lv e s

( Fi g 5 2 A) ; ( 2) u ne qu al cle ava ge whe r e th e fi r st divi sio n Of th e

.
, ,

e gg r e s ult s in o ne l a r g e a nd o ne s m a ll c e ll ( Fi g 5 2 B ) ; ( 3 ) di s .
,

ooid al cle a vage wh e r e th e e ntir e e gg d o e s no t di vid e

,
but sm all ,

c e lls a re cu t Off a t the surf a ce a nd for m a di sk Shap e d r e gio n -

( Fi g 5 2 C ) ; a nd (4) su pe rfi ci al cle avage wh e r e th e nucl e us o f th e

.
, ,

e gg di vid e s ra pidly ; t h e d a ught e r nu cl e i mig ra t e t o th e p e r iph e r y

a nd f o r m a singl e l a y e r Of c e ll s a t th e s u r f a c e ( F i g 5 2 D) .
,
.

Tha t p ar t of o ntog e ny which co n ce r n s th e de ve l opm e nt Of a n

a nim a l f r o m th e e gg t o m a tu r ity i s kn o wn a s e m bryoge ny Ce r .

t ain s ta ge s in thi s de ve l opm e nt h a ve b e e n r e co gnize d as co mmo n

t o a ll high e r a ni m a l s a nd h a v e b e e n giv e n n a m e s
, Th e s t a ge s .

o ccu r in a ce r t a in r e gul a r o r d e r a nd a s a n int r o ducti o n to th e m or e

,

de ta il e d sp e ci al a cco unts giv e n f or e a ch typ e s tudi e d in sub se

que nt ch apte r s w e sh all p r e s e nt a b ri e f e mb ry ol o gi ca l hi s t ory of a
,

typica l h ol obl as tic e gg T h e st age s to b e co n side re d are : ( I )

.

cl e av ag e ( 2) th e m o r ul a (3 ) th e bl a s t u l a (4) th e g a s tr ul a ( 5) the
, , , ,

for m a tio n of ge r m l a y e r s a nd ( 6) o r g a no ge ny
,
.

C le avage in a h ol obl a sti c e gg ( Fi g 53 ) r e sul t s in th e p r o ducti o n

.

o f tw o ( B ) f o u r ( C D ) e ight ( E ) sixt e e n F ) e t c c ll
, , , ( ,
e s a pp r o x i ,
.
,

m a t e ly e qu a l t o o ne a n o the r and g ro wing sm all e r a s the i r numb e r

incr e a s e s E ach o f th e se ce ll s i s kno wn as a blas tome r e T h e
. .

bl as to m e r e s d o no t se p a r a t e a s d o th e d a ught e r ce lls p ro du ce d
by the bin a ry divi s io n Of P a r a me ci u m ( Fi g 3 1 o q) but r e ma i n
— .
FI G 5 2
. . F i gure s ill u s t r a ti ng f o ur d i ff e re nt k i nd s
l v age A e q ua l
of c ea .
,

cl e a va ge o f t h e s e a u rchi n e gg ; B u ne q u a l cl e a v a ge o f t h e e gg o f
“
,

m a r i ne w o r m ; C d i s co i d a l cl e a va ge o f th e e gg o f a s q u i d ; D su pe r fi ci al
, ,

cl e a v a ge o f a n i ns e c t s e gg —B f ro m Wil so n C f ro m Wil so n a f t e r
( A
’
.
, ; ,

Wa t a se ; D f ro m Ko r sch e lt a nd He i d e r )
, .
I IO AN IN TR ODU C TION TO Z OOLOGY

a tt a che d to one a n o th e r a s was n o te d i n th e c ase of P and or i na

p
( 94 Fi g 4 5 II ) T h e r e s e mbl a nce o f th e gro up o f bl a s
.
,
.
,
.

t o m e r e s to a mulb e rr y sugg e s t e d th e t e r m mor u la w hi ch i s o ft e n .

us e d in de sc ribing th e e gg during th e e a rly cl e a va g e s tage s

, .

B l a s tu la As cl e a va g e a dv a nc e s a c a vity b e c o m e s n o tic e abl e

.
,

in the c e nte r o f th e e gg ( Fi g 53 H) b e co ming l a rg e r a s d e ve l op

.
, ,

m e nt p ro c e e ds until th e whol e re se mbl e s a holl o w r ubb e r b all ,

th e r ubb e r b e ing r e p re se nt e d by a s ingl e l a y e r o f c e lls At thi s .

st age th e e gg i s call e d a blas tu la th e c a vity th e cle avage o r s e g ,

me ntati on cavi ty a nd th e c e llul a r l a y e r th e blas tode rm

, The .

bl a stula re se mbl e s s o m e wha t a Singl e col o ny o f Volvox ( Fi g .

46 , A) .

G as tru la . c e ll s o n o ne side o f th e bl a s tul a a re s e e n to be

The
thick e r tha n e l se wh e r e ( Fi g 53 K) a nd b e gin to inva gin a te .
,

( Fi g 53 L) Thi s p ro ce ss r e s ult s in a cup sh ap e d s t r uctu re with

.
,
.
-

a w a ll o f two l a y e rs a n o ut e r l a y e r o f sm a ll c e lls a nd a n inne r

,

l ay e r o f l a rg e r c e ll s T h e e mb ry o m ay now b e call e d a gas tru la

.

( M ) a nd th e p r oce ss by whi ch it d e ve l o p e d f r o m th e bl a stul a

,

i s t e r m e d ga s tr u la ti on T h e cl e a v a g e c a vity i s a lm o st ob lit e r a t e d
.

du ring th e invagin a tio n whil e a ne w ca vity th e p rimitive di g e s

, ,

tiv e tr a ct o r a r che nte r on i s e s ta bli sh e d , .

G e rm lay e r s T h e c e ll s o f o ne l a ye r o f th e g a s t r ul a r e s e mbl e
.

o ne a n o th e r but di ff e r in a pp e a r a n ce f r o m th e c e lls o f th e o th e r
,

l a ye r E a ch l a y e r give s r i se to ce r ta in d e finite p ar ts o f th e b ody

.
,

a nd i s th e r e fo r e t e r m e d a ge r m laye r ; th e o ut e r is th e e ctode r m
‘

( g 53
F i . N , ,
t h e inn e r t h e e ntod e r m ( N
,
A nim a l s wit h ,

o nly th e s e t wo l a y e r s a r e s a id t o b e d i ploblas ti c; but th e m aj o rity

o f th e high e r a nim a l s h a v e a thi r d l a y e r w h i ch u s u a lly a pp e ar s ,

b e tw e e n th e fi rs t two a ft e r th e ga s t rul a h as b e e n fo r m e d Thi s is .

th e middle l a y e r o r m e s od e r m,
It o rigin a te s e ithe r fro m th e pr o
.

li f e r a ti o n of a f e w sp e ci a l ce ll s whi ch m ay b e re c ogniz e d in th e
e a r ly cl e a v a g e s t a g e s o r f r o m ce ll s budd e d Off f r o m th e inn e r
,

sur face o f b o th th e e cto d e r m a nd e nto de rm or fro m p o uche s ,

a r i s ing f ro m t h e w a ll s o f th e e nt o d e r m ( Fi g 5 3 N) Anim al s .
,
.

with th re e g e rm l a ye rs a re s aid to be tri ploblasti c .

 AN IN T R ODU C TION T O T HE M ET AZ OA

Organoge ny . E a ch
g e r m l ay e r giv e s ris e to ce r t ain org ans
o f th e a dult a nim a l An organ m ay b e d e fin e d a s a n a sso ci a ti o n
.

o f tissu e s h a ving a d e fi ni t e f or m a nd sp e ci a l functi o n In a ny .

o f th e m o r e c o mpl e x a ni m a ls e i ght s yst e ms Of o r g a ns ca n b e r e c o g

ni z e d : ( 1 ) di g e s tiv e ( 2 ) ci r cul a t or y (3 ) r e spi ra t o ry (4) e xcr e

, , ,

tory ( 5) s ke l e t al and inte gum e nt ary ( 6) re p r o ductive ( 7) m u s

, , ,

cula r ( 8 ) n e r v o u s
,
T h e s tudy o f th e o r igin o f th e s e o r g a ns f ro m
.

th e g e r m l ay e r s is kn o wn a s or ga noge ny T h e o r g a n s d e r iv e d
.

from the di ffe r e nt ge r m l a y e rs m ay b e b ri e fly li st e d a s foll ow s .

Fr o m th e e ctode r m a ri se th e e pid e r mi s e pith e lium o f v a r i o us ,

or g a n s a nd th e n e r v o us s yst e m ; fro m th e me sod e r m c o m e th e

,

muscl e s co nn e ctiv e a nd supp or ting ti ssu e s a nd bl oo d and bl o o d

,

v e sse l s ; th e e ntode r m b e c o m e s th e e pith e lium of the dige s tiv e

t ra ct pharynx a nd re spi ra t o ry t ra ct
, ,
.

C oe lom F igu re 5 3 N 0 Sh o ws o ne m e th o d o f or igin o f th e

.
, ,

c oe l o m a c a vity of g re a t imp or t a nce in the b o di e s o f m a ny of th e

,

compl e x M e t azoa T wo p o uche s ( N) f or m e d fr o m th e inn e r

.

l aye r of th e g ast r ul a a r e pinche d Off a nd co m e to li e o ne on e ithe r

Sid e o f t h e a lim e nt a r y c a n a l ( O) T h e c a viti e s o f th e se s a cs
.

constitut e th e coe l o m By a l a te r a l gr o wth a nd a b re ak ing

.

th ro ugh a t th e e nds of the sa cs the ca viti e s uni t e to for m a Singl e

,

sp a ce b e twe e n th e al im e ntary c a n al a nd th e b o dy w all All .

M e t a zo o ns a r e fr e qu e ntly s e p ar a t e d int o tw o g ro up s th e coelo ,

ma ta a nd the acce loma ta a cc or di ng a s th e coe l o m i s p re se nt or

,

a b s e nt Of th e typ e s co n sid e r e d in th e fo ll o wing ch apt e r s

.

Hydr a m ay b e m e nti o n e d a s a n a c oe l o m a t e th e e a r thw or m as ,

a c oe l o m a t e .

T abl e III c o nt ains a se r i e s o f di a gra m s a nd d e scr ipti ons which

a re int e nde d t o r e p re s e nt th e m e th o d s o f r e p ro ducti o n e xhi bit e d

by the typ e s al re a dy s tudi e d a nd to sh o w th os e of ce r t ain M e ta

,

zo a a s w e ll In o r de r t o m a k e th e di a gr a m s a s simpl e a s p ossibl e
.

ce r ta in de t ails h a v e b e e e n o mitt e d ; n e v e r th e l e ss it i s b e li e v e d ,

th a t a c orr e ct id e a o f th e r e p ro ductive p ro ce ss e s wi ll b e g ain e d

in e v e ry cas e .
1 12 AN IN TR ODU C TION TO ZOOLOGY

TABLE II I

DI AGRAM S I LLU STR ATI N G THE M ETHOD S OF R EP R ODU CTI ON IN P ROTOZ OA

AN D M ET AZ OA

A . A simpl e P ro t o z o o n (Ame ba) .

CE LL V
DI I SI ON T M NT A D S
EN CY S E N UB SE CE LL V DI I SI ON
( Bi na ry F iss i o n) Q U T C LL D V
EN E I I SI ON ( B i nar y F i ss io n)
An i nd e fi ni te nu m be r of ( Sp l ti n)
o ru a o An i nd e fi ni te nu m be r
ge ne ra t io ns . M a ny pse u d opod i o s po re s psp) o f ge ne rat i on s .

( young Ame bas) prod uce d .

B . A simpl e P ro t o z o o n (P a r a me ci u m) .

CE LL DI V I SI ON TE MP O RAR Y C ON J U CE LL
DI I SI ON V
CE L DI I SI ON L V
( B ina ry F is s i o n) G ATI ON ( P e ri o d o f R e co n ( Bina ry Fiss ion)
An i nd e fi ni t e nu m ( Fe rt ili za t io n) s t ru ct i o n) An i nd e fi ni te num
be r of ge ne rati ons . Each ce ll f t ili
er ze s l l
E a ch f e rt i ize d ce l be r o f ge ne ra i ons t .

th e o th e r. v
gi e s r ise t o f o ur
norm a ani mals l .
I I 4 AN IN T R ODU C TION TO ZOOLOGY

E . A mo r e c o mpl e x co l o ny Of c e lls ( Volvox) .

CE LL
DI I S I ON C E V DI I SI ON C E LL V LL
DI I SI ON P E R M N E N V CE DI I SI ON A T LL V
l
( Co o ny f o r ( As e xua R e l
( Ga m e te f or C ON J I ON ( Co lo ny f or U G AT
m a ti o n) pr od u ct i o n) m a t i o n) ( F e r t i i za t i o n) m a t i o n) l
Z ygo t e ( z ) Ge r m ce s ll
C e r t a i n ge r m One s p e r m Z ygo te d e v e o ps l
vl
d e e o ps i nt o a gi e r ise t o v
ce s pr o d u ce ll
f use s w i t h o ne i n o a co ony t . l .

co o ny l ne w co o ni e s
. l
e ggs ( e ) ; o th e rs e gg f or m i ng a
.
, .

Spe r m a t o zo a zygo t e

F . A S impl e M e t a zo o n (Hydr a) .

-
n

—
Q

—
z
f
— J u — f a h w _ Z
J

CE LL DI I SI ON V B DDI N G U CE LL V
DI I SI ON P E M NEN R A
CE DI I S I ON T LL V
( Em bryo o g ica l l
( As e xu a l
( Ga m e t e f o r C O N JU GAI I ON ( Em b ryo o gi cal' ‘
l
vl
d e e o p m e nt ) R e p r o d u ct i o n) m a t i o n) l
( F e r ti i za t i o n) d e e o pm e nt ) vl
Z ygo te ( z ) p r o P a r t o f a ni m a l
C e rt a i n ge r m One s p e rm Z ygo t e ( 2 ) p r o
d uce s ani m a l
se p a ra t e s f r o m l
ce l s pr o d u ce u ni t e s w i t h o ne d uce s a ni m a , l
co nt a i ni ng p a r e nt a nd o ne e gg ( e ) a nd e gg f o r m i ng a , e tc .

ge r m ce s ll l
e a d s s e pa r a t e t w o po a r l
z ygo e ( z ) t .

a nd t w o a ye rs l e xi s t e nce bod i e s ( pb )
. .

o f s pe ci a li z e d o t h e rs p r o d uce

s o m a t i c ce llS, t h e m a ny s pe r ms
e ct o d e r m ( c c ) .

a nd e nt od e rm
 AN IN TR ODU C TION T O T HE M ET AZ OA 1 1
5

G . A co mpl e x M e t a zo o n (Lu mbr i cu s) .

pb . e

ms en
. . ec .
 C HAP T ER VIII

HYDRA AND C G LENTERATE S I N GENERAL

I . H Y DR A

( y f ca Linnae us)
H d ra u s

Hydr a fu sca i s impl e M e t a zoo n a bund a nt in f re sh w a te r

a s

p o nd s a nd s t re a m s If a qu an tity o f a qu a tic Ve ge ta ti o n i s g a th
.

e re d a nd pl a ce d in gl a ss di s h e s full o f w a t e r th e se lit tl e f r e s h -w a t e r ,

p oly ps will be fo und clinging to t he pl a nt s a nd the sid e s a nd bo t

to m Oi the di s h Th e y a re e as ily se e n with the n ak e d e ye b e ing
.
,

f ro m 2 to 20 mm in l e ngth a nd m a y be like ne d to a sho rt thick

.
,

th re a d f ra zzl e d a t the un a tt a ch e d di s t a l e nd T he g re a t v ari a , .

ti o n in l e ngth i s d ue to t h e f a ct th a t b o th b o dy a nd t e nt a cl e s a re
c ap abl e o f re m a rk a bl e e xp a nsi o n a nd c o nt ra cti o n b e ca u se o f the
pr e se nce o f sp e ci aliz e d mus cl e fib rils in m any o f th e c e lls .

T ABLE I V

THE N AMES AN D C HARACT E RI STI C S o r T HE P RI N C I P AL S P EC I E S o r H DRA Y ( 1 05 )

A UAL WHE N Sax

L R U A LL
l

i N EM S EX
G EN U S S PEC I E S CO O
EN '
.

Y
M AT U
TO C Y S TS C O N Dm ON
RE

Hyd ra f us ca bro w n 1 0-6 l a rge He r m a phro 2 cm . Se p t —Oct . .

or fe we r d i ti c
vi r id i s gre e n 5
-1 2 s ma ll
gr is ca gra y 5
-I S v e ry
l a rge
to De c .

cm . Oct -De c .
1 1 8 AN INTR ODU CTION TO ZOOLOGY

u nfa vorabl e condi tio n s a nd e xt re m e a ge re sult in a de c r e as e

F r e qu e ntly sp e cime n s of Hydr a a re fo und which p osse ss buds

in va ri o u s s t a ge s of d e ve l opme nt Se ve r al bud s a r e o ft e n f o und
.

o n a Singl e a nim a l a nd th e s e in mi n m a y h e a r buds b e f or e d e t a ch

,

FI G 5 4 D i agr am o f a l o ngit u d i na l s e cti o n o f Hydr a ; b , b u d ; b d ,

. . . . .

d i sk ; bl , b l a s t u l a ; e c , e ct o d e r m ; e n , e nt o d e r m ; g , ga s t ru l a ; gv c ,
. . . . . .

g a s t r o v a scu l a r ca vit y; b h
y , yp o s t o m e m ,
m o u th ; m e ,
m a t ure e gg ;
. . .

m e s , m e so gl e a ; m t , m a t u r e t e s ti s ; n , ne m a t o cys t s ; p b , po l ar bod
. . . . . .

i e s ; t , t e nt a cl e ; y e , yo u ng e gg ; y t , yo u ng t e s ti s
. . . .All o f t h e s tr u c
. .
~

tu r
es s ho wn d o no t o ccu r o n a Si ngl e an i m al a t o ne ti m e .
 H Y D R A AN D C CELENT ER AT E S I N G EN E R AL I I g

m e nt fr o m th e p ar e nt In thi s w ay a s or t o f p rimitiv e Hydr a

.

c ol o ny i s f or me d re se mbling s o m e wha t th e ase xu a l c ol o ni e s o f

,

s o m e of t h e m o r e co mpl e x C oe l e nt e r a t e s to b e d e sc rib e d l a t e r
( g 5, )
F i 6
. A .

R E P R ODU C TI V E OR GAN S m ay b e Obs e r v e d on s p e cim e ns Of

Hydr a f u sca in Se pte mb e r a nd Oct ob e r B o th . a n ova r y a nd te s te s

are p r o duce d o na singl e individu a l ; t h e for m e r is kn ob like o ccupy -
,

ing a p ositi o n a b o ut o ne thi r d th e l e ngth o f th e a ni m al a b ov e the

b a sa l di sk ; the te s t e s u su ally t wo o r m o r e in numb e r a re c o nic al
, ,

e l e v ati o n s p ro j e cting f ro m th e di s t a l thi r d o f th e b o dy .

S tru ctu r e ( Fi g . Hydr a i s a di pl obl a stic a ni m a l co n

s i s ti ng o f t w o c e llul a r l a y e r s a n o ut e r thin c o l or l e s s l a y e r t h e
, ,

e ctod e r m a nd a n inn e r l a y e r t h e e ntod e r m , twic e a s

thick as th e o ute r a nd co nt a ining th e b r o wn b odi e s w hi ch giv e
Hydr a f u sca i t s ch ara ct e r i s tic c ol o r B o th l a y e r s a r e c o mp o s e d
.

o f e pi the li a l ce lls A thin sp a c e c o nt a ining a j e lly lik e s ubst a nc e

.
-
,

t h e m e s ogle a ( m e s ) s e p a r a t e s e ct o d e r m f r o m e nt o d e r m Not .

o nly t h e b o dy w a ll but a ls o t h e t e nt a cl e s p o s s e s s th e s e th r e e
, ,

d e finit e r e gi o n s T h e b o dy with th e e xc e pti o n o f th e b a sal di s k

.
, ,

i s c o v e r e d by a thin t r a n sp a r e nt cu ti cle B o th b o dy a nd te n .

t a e l e s a r e holl o w t h e singl e c e nt r al sp a ce b e ing kno wn a s t h e

,

ga str ovas cu la r ca vi ty

E C T ODE R M . t d e r m i s p r im a rily pr ote cti ve a nd s e ns ory

T he ec o ,

c ont aini ng s t r uctu re s ch a r a ct e ri s tic o f the se functi o ns Slight .

di ff e r e n ce s in s tr uctu r e a r e o b se r v a bl e b e twe e n th e e ct o de r m o f
t h e t e nt a cl e s a nd th a t o f t h e b o dy w a ll whi l e t h e l a tt e r di ff e r s
,

fr o m tha t o f th e b asal di s k In t h e e ct o d e r m o f the b o dy wall

.

a r e t wo p rin cip a l kind s o f ce ll s l a r g e e pithe li o mu s cul a r c e ll s

, ,

a nd s m a ll int e r s titi a l ce ll s T h e l a tt e r giv e r i s e t o c e ll s c a ll e d

.

cnid o bl a s t s w hich f o r m n e m a t o cy s t s a nd t o b o th m a l e ,
a nd

fe m al e g e r m c e ll s T he e pi the li omu scu la r ce lls a r e shap e d lik e

.

inve r te d co n e s At th e i r inn e r e nd s a re o ne o r m o r e co mp a ra
.

t i v e ly l o ng ( s o m e tim e s 3 8 mm ) un s t r ip e d.con tr acti le f i b e rs

whi ch f o r m a thi n l o ngitudin a l mu scul a r l a ye r The se mus cl e .

fib e r s e xpl ain th e r e m a r k abl e p o we rs o f co nt ra cti o n e xhibite d

1 20 AN INT R ODU CTION TO Z OOLOGY

by Hydr a wh e n stimula t e d N e a r th e middle o f e a ch c e ll e m .

,

b e dde d i h th e alve ol ar cytopl a s m i s a nucl e us co nt aining one o r ,

tw o nucl e oli a nd a n e tw o r k o f ch r o m a tin .

N E MATOC Y S TS o r stinging c ap s ul e s ( F ig s 54 n ; 5 6) a r e p r e s e nt .
,

o n a ll p a r ts o f th e b o dy o f Hydr a e xc e pt th e b asa l di s k b e ing ,

m os t num e r o us o n
th e t e nt a cl e s ( Fi g .

55 A ) E a ch is co n ,
.

t a i ne d i n a c e l l
kn own a s a cnid o
bla s t Th e se in tur n .

are e mb e dd e d in lit

tle tub e r cl e s o n th e
s u r fa c e which give
t h e a nim a l a r ou gh
a pp e ar ing o utlin e .

T he tub e rcl e s a r e
e ct o d e r m c e ll s e a ch ,

o f whi ch u s u a lly po s

s e ss e s o ne o r m o r e
l arg e n e m a t ocysts
surrounde d by a
FI G 5 5 N m t y t
. . e nd th i
a oc ti n A p
s s a
numb e r Of a s m a ll e r
e r ac o .
,
or

ti n f t nt l h wi ng th b tt i
o o a e ac e f v a r i e ty Th re e k l nd s
S o e a er es o .

n m t y t ; l
e a oc ni d
s s il ; B i n t l v o f n e m a t o cys t s a re
c .
,
c oc s ,
se c ar a

v d with n m t y t
co e re e lt f p fo und in H dr a T h e
a oc s s as a re su o ca
y
t u re by Hyd ( F m J nni ng )
ra ro e s
l arg e s t is 0 1 3 mm
.

l o ng a nd 00 7 mm thick ; b e fo r e b e ing di sch a rge d it i s p e a r shape d

. .
-

a nd o ccupi e s a lm o s t t h e e nti r e c e ll in whi ch it li e s ( Fi g 5 6 .

,

Within it i s a coi le d tu be (t) a t wh ose b a s e a r e th r e e l arg e a nd a num

b e r o f s ma ll s pi ne s P r o j e cting f r o m th e ce ll n e ar th e o ut e r e nd o f
.

th e n e m a t o cy s t i s a t r igg e r lik e spin e th e cni d oci l - Ne ma ,

é

t o cys t s m ay b e e xpl o d e d by a dding a littl e a ce tic a cid o r b e tt e r , ,

m e th yl g r e e n to th e w a t e r T h e tub e whi ch i s co il e d within

,
.

th e m i s th e n e v e r te d Fi r st th e b ase o f th e tub e with th e spin e s

.
,
I 22 AN I
a NT I
R OD U C T ON T O Z OOLO GY

c e ll s e a ch c e ll p ro ducing o ne n e m a to cyst
,
F i r s t a cl e ar sp a ce ap
.

p e a rs in an int e r s titi a l ce ll ; thi s sp a ce e nl arg e s it a cqui r e s a d e fi

,
~

nite w a ll a nd i ts co nt e nt s s t a in d e e ply P r e se ntly it e l o ng a t e s a nd

, .
,

o ne e nd i s p r o duc e d t o f or m t h e thr e a d w hi ch a t i t s fi r s t a pp e ar
,

a n ce i s e v e r t e d a nd co il e d r o und t h e o ut sid e o f t h e sa c Aft e r a

.

tim e th e th re a d is int rov e r te d it i s no t qui te cl e a r h ow a nd

FI G 5 7
. . Th e ac ti o n o f ne m a to cys t s A a ne m a to cys t p i e r ci ng the chiti no u s
.
,

co v e r i ng o f a n i ns e c t ,
B ne m a t o cys t s h o l d i ng a small anhna l by co ili ng
a bo u t i t s sp i ne s
.
(Af t e r T o ppe i n Z ool Anz )
. .
 HY DRA AND C ( ELEN TERAT ES I N GEN ERAL

th e n e m a t o cy s t a ssum e s i t s fin a l f o r m Wh e n n e arly rip e a n e m a

.

t o cys t s till c o nt a in e d in i t s m o th e r c e ll o r cnid obl a s t mig r a t e s int o

, ,

th e in s id e o f a n e pith e li o mu scul a r ce ll a nd a pp r oa ch e s t h e su r f a c e .

Th e e xt e rn a l e nd o f th e cnid obl a s t i s p r o duc e d t o fo r m a cni d o cil

whi ch p e r fo r a t e s th e cuticl e (1 01 p
. Sin ce th e
,
.

tub e o f th e n e m a t ocys t c a nn o t b e r e tu rn e d t o th e c ap s ul e no r ,

a n o th e r o ne b e d e v e l o p e d by t h e cnid o bl a s t ne w cap s ul e s must ,

b e fo r m e d f r o m int e r s titi a l ce lls to r e pl a c e th ose a l r e a dy e xp l o d e d .

T h e s e c o nd kind o f n e m a t ocys t i s cylind ric a l ( Fi g 5 5 A) a nd .

,

co nt ains a thr e a d whi ch l a ck s th e b a rb s so ch a r a ct e ri s tic o f its

l arg e r n e ighb or T h e thi r d va r i e ty is a lm os t sph e r ic al a nd sm all e r
.

tha n th e o the r s m e as uring o nly 00 5 mm in di a m e t e r T he

,
. . .

thr e a d conta ine d in thi s n e m a to cys t like wi se b e a rs no b arb s but ,

whe n di sch a r ge d r e se mbl e s a cor kscr e w ( Fi g 5 7 B) It a ids in . .

th e c aptu r e o f p r e y by c o iling a r o und th e Spin e s o r o th e r s t r u c

tu re s th a t m ay b e p r e se nt
T h e int e rs titi al ce ll s a l so d e v e l o p a t a c e r t a in p e r i o d o f th e y e ar
( Se pt e mb e r a nd Oct ob e r) int o ge r m c e ll s T h e Origin a nd h i s .

t o r y o f the se ce ll s will b e fo u nd fully d e scrib e d o n p a ge s 1 3 4 1 3 6 —

.

Th e B AS AL DI S K ( 5 4 b d ) di f fe r s so m e wh a t in fun cti o n f ro m t h e
,
.

r e s t o f t he b o dy It i s th e p o int by which Hydr a a tt ache s itse lf

.

t o s olid Obj e ct s a nd f o r thi s pu r p o se se c r e t e s a s ti cky s ub s t a n ce

, .

It i s al so said to e ffe ct th e mo ve m e nt o f th e a ni mal fr o m pl ac e to

pl a c e by a so r t o f gliding m o ti on no t ye t fully e xpl ain e d but, ,

p o ssibly b r o ught ab o ut by p se udop o di a like p ro ce sse s thr u s t o u t -

fro m s o m e o f th e ce ll s Epith e lio mu scul a r ce ll s a nd a f e w int e r

.

s ti ti al ce ll s a r e p re se nt but no n e m a t o cy s ts a r e t o b e f o und h e r e
, .

T he c olumn a r e pith e li o mu s cul a r ce ll s a r e no t o nly p r o vid e d with

co nt r a ctil e fib e r s a t th e i r b as e s but b e ing s e c r e t o ry a l so co nt a in
, , ,

a l a r ge numb e r o f s m a ll r e f r ing e nt g r a nul e s a s s h o wn in F igu r e 8

, 5 ,

ad . se z .

Th e T E N T AC LE S ( Fi g p r o vid e d with an e cto d e r m

.
5 4 , t) are

co n si s ting o f l a r g e fla t ce ll s thi n a t t h e e dg e s a nd thi ck in th e

,

ce nt e r .T h e thi ck e r p o r ti o n s give th e s u r f a c e o f t h e t e nt a cl e
a l u mpy a pp e a r a nc e In th e ce nte r o f e a ch thi cke ni ng i s a
.
1 24 AN INTRODU CTION TO ZOOLOGY

nucl e us around whi ch a r e e mb e dd e d s om e tim e s a s m a ny as twe lve

n e m a to cyst s e a ch in i t s o wn cni dobl as t ( Fi g 5 5 A) Th e cni .
,
.

d o cils p r oj e cting fr o m th e cnid obl a s t s r e se mbl e gr oup s o f cili a .

Ea ch cni d o bl a s t i s d r a wn ou t a t its b a s e int o a c o nt r a ctil e fib r il

whi ch e nte r s th e l o ngitudin a l mus cular she e t a t th e b a se of the
e ct o d e r m c e lls .

E N TODE R M ( Fi g 5 4 T h e inn e r l a ye r o f c e ll s th e e nt o
.
, ,

d e r m o ccupi e s ab o ut tw o thi rd s o f th e b o dy w all Its fun ctio n s

,
.

a r e d i ge s ti ve a nd s e cr e tory Th e di .

g e s t i v e ce ll s a r e l o ng a nd Club
sh ap e d with t r a n sve r s e muscul ar ,

fib ril s a t the i r b a se for ming a ci r ,

cu la r sh e e t o f c o nt r a ctil e s ubst a nc e .

At th e l a r ge r e nd whi ch e xt e nd s ,

b m
e 0
into the c e ntr a l ga st rova scul ar
FI G 5 8 T h
. .
gl n d l ll ca
re e vity a rae t w o flg
a
u ar e lla P
ces e udso ,
.

f m th b
ro l d i k f Hy podi a m ay al so b e th r u st ou t f ro m
e a sa s o

4 ‘ ad Of
thi s fre e e nd Th e int e r n a l st r u c
31 w
.

.
.

a 1e s n mm
e 56 c lo
tu r e of th e s e c e ll s di f fe r s b e fo r e a nd
'

D hlg n nd K pn )
a re a e er .

a ft e r th e a mm a l I s f e d In a s ta rv ‘

ing Hydr a l a r ge va cu ol e s a pp e a r a lm o s t c o mpl e te ly filli ng the ,

c e ll the p ro topl as m b e ing re duc e d to a thin l aye r n e a r the ce ll

,

w a ll ; af te r a m e al ho we ve r th e ce ll s a r e go r ge d with nut ritive

, ,

sphe r e s m a ny o f which e sp e ci ally th e oi l gl obul e s mig ra te i nto

, , ,

th e e ct od e r m a nd a r e s t o r e d n e a r th e p e r iph e r y giving th e ,

a nim a l i ts b r o wn co l o r

T h e gla nd u la r ce lls a r e sm a ll e r th a n th e dige stive c e lls a nd l a ck ,

th e co nt r a ctil e fib r il s a t th e i r b a se The y a r e b roa d a t t he f r e e .

e nd a nd thin o u t t o a fi ne fil a m e nt which e nds in a kn o blik e

,

e nl a r ge m e nt wh e n t h e m e sogl e a i s r e a ch e d T h e gl and c e ll s .

a l so di ff e r in a pp e a r a n ce a c co r ding t o th e i r m e t a b olic a ctivity :

.

s o me a re fill e d with l a r ge v a cuol e s c o nt a ining se cre t ory m a tte r ,

whil e o th e r s h a ving di sch a r ge d the i r se cr e tum app e a r cro wde d

, ,

with fi ne gr a nul e s Int e r stiti a l c e ll s a re fo und lying a t th e b ase

.

o f th e o th e r e nt o d e r m c e lls .
1 26 AN I NTRODU C TION TO Z OOLOGY

t e nta cl e which h as captur e d th e pre y b e nd s to wa r d th e m o uth

, ,

with its l o a d of f oo d T h e o th e r t e nta cl e s no t o nly a ssi s t in thi s

.
,

but m a y u se the i r n e m a to cys t s in qui e ting th e victim T h e .

m o uth Ofte n b e gin s to Op e n b e fo re t h e fo o d h a s r e a ch e d it T h e .

e dg e s o f t h e m o uth g r a du a lly in cl o s e t h e o r g a ni s m a nd f o r c e it

int o th e ga st ro va scul a r c a vity T h e b o dy wa ll co nt ra cts b e hind

.

t h e f o o d a nd fo r ce s it d o wn until it r e a ch e s th e b a sa l e nd o f th e
b o dy He re it r e m a in s dur ing the p ro ce ss o f dige s tio n F r e
. .

qu e ntly or g a nism s m a ny tim e s th e siz e o f th e Hydr a a re s u cce ss

fully inge s te d .

R E AC TI ON S T O F OOD —. I t i s no t unco mm o n t o find Hydr a s th a t

will no t re a ct to foo d whe n it is p r e s e nte d to th e m Thi s i s d ue .

t o the f a ct th a t th e s e a nim a l s will e a t o nly wh e n a c e r t a in int e r v a l

o f tim e h a s e l a p se d s in ce th e i r l a s t m e a l T h e phy si ol o gica l
.

co nditi o n o f H yd ra th e re f o re d e t e r min e s i t s r e sp o n s e t o t h e f oo d
, ,

s timulu s . T h e colli s i o n o f a n a qu a ti c o r ga ni s m with th e t e nt a cl e

o f Hyd r a i s no t s u ffi ci e nt t o ca u se t h e f oo d t a king r e a cti o n s in ce
-
,

it ha s b e e n fo und tha t no t o nly a m e cha ni ca l s timulus but a l so a ,

che mi ca l s timulu s mus t b e pr e se nt A v e r y hungry Hydr a will

.

e v e n go th r o ugh t h e ch a r a ct e r i s ti c m o v e m e nt s wh e n it i s e x cit e d

by th e ch e mica l s timulus a l o n e Thi s h a s b e e n s ho wn by th e

.

fo ll o wing e xp e r im e nt Whe n the t e nt a cl e s a nd hyp o s to me o f a

.

m o d e ra t e ly hung ry Hyd r a a r e b ro ught into c o nt a ct with a pie ce

o f fil t e r p a p e r w hi ch h a s b e e n so a k e d f o r a tim e in th e s a m e cu l
,

tu re m e dium the re is no r e sp o n se If th e filte r p a p e r i s the n

, .

so a ke d in b e e f j uic e a nd o ff e r e d to th e Hydr a th e u su a l foo d ,

r e a cti o n s a r e giv e n .

B e e f jui ce a l o n e c a ll s fo r th no r e sp o n se in a m o d e r a t e ly hung r y
a nim a l ; but d o e s in a ugu r a t e t h e n o r m a l r e fl e x if a v e r y hung r y
,

sp e cim e n i s se l e ct e d f o r th e e xp e r im e nt T h e c o nclusio n r e a che d

.

i s th a t w e ll f e d Hydr as will no t r e sp o nd t o e ith e r m e ch a ni ca l or

-

che mica l s tim ul i whe n a cting a l o n e o r in co mbin a ti o n ; th a t

m o de r a t e ly hung r y a nim a l s wi ll r e a ct to a co mbin a tio n o f th e
t w o a nd th a t hung r y a nim a l s will e xhibit f o o d t a king m o v e m e nt s
,
-

e ve n if a ch e mi cal s timulu s al o n e i s e mpl o y e d

 HY DR A AN D C ( ELE N T E RAT E S I N GEN ER AL 1 27

DI GE STI ON Imm e di a t e ly a fte r th e ing e sti o n o f foo d the gl a nd

.
—

c e ll s in th e e nto de r m Sho w sign s of gr e a t a ctivity ; th e i r nucl e i

e nl a r g e a nd b e co m e g r a nul a r This is d ue p ro b ably to th e fo r m a
.

tio n o f e nzym e s which a r e di sch a r g e d into th e ga s t ro v a scul a r

c a vity a nd b e gin a t o n ce th e di sso luti o n of the foo d T h e a cti o n .

o f t h e dig e s tiv e j ui ce s i s m a d e m o r e e f f e ctiv e by t h e chu r ning o f

th e f o o d a s th e a ni m a l e xp a nd s a nd c o nt r a ct s T h e cili a e xt e nd
.

ing o u t into th e c e nt r a l c a vity a l so a i d in the diss olutio n o f th e

fo o d by Cr e a ting cu r re nts This m e tho d o f dig e s tio n diff e r s f ro m
.

tha t of Am e ba a nd P a r a me ci u m in b e ing c a r ri e d o n o utsid e o f th e

c e ll ; i e e xt ra c e llul a r The r e is e vid e nce th a t int r a c e llul a r
. . .

dige s ti o n a l so t a kes pl a ce in Hydr a ; th e p se udop o di a th r ust ou t

by th e e nto d e r m c e lls se iz e a nd e ngulf p a r ticl e s Of fo o d which a re
di sso lv e d within th e c e ll s Ho w e v e r m os t o f the fo o d i s di
.
,

g e s t e d in t h e ga s t ro v a s cul a r ca vity T h e dig e st e d f o o d i s .

a b s o r b e d by th e e nt o d e r m ce ll s ; p a r t of it e sp e ci a lly th e o il ,

gl obul e s i s p a s se d o v e r to th e e cto d e r m wh e r e it i s sto r e d

, , .

E GE S TI ON — All in so lubl e m a t e r i a l i s e g e s t e d f ro m th e m o uth

. .

Thi s i s a cc ompli s h e d by a v e r y sudd e n squi r t whi ch thro ws

th e d éb r i s t o s o m e di s t a n ce
B e h avi or . Hyd r a vi r idi s giv e s a m o r e p r o mpt a nd d e ci s iv e
r e s p o n s e wh e n s timul a t e d th a n a n th p e ci e s o f H d r a a nd
y o e r s y ,

f o r t hi s r e a s o n i t s b e h a vi or h a s b e n s tudi e d m or e th o r o ughly
“

th a n th a t o f th e o the r s T h e fo ll o wing p ara gr a phs h a ve b e e n

.

co mpil e d l a rg e ly f ro m e xp e rim e nt s up o n gr e e n Hydr as a ltho ugh ,

e n o ugh w o r k h a s b e e n d o n e with o th e r f o r m s to p ro v e th a t th e i r

r e a cti o n s a re p r a cti ca lly t h e s a m e o nly m o r e s lug i s h

, g .

N O R MAL P OSI TI ON OF H YDR A Hyd r a s m a y b e f o und a tta ch e d

.
-

t o th e s id e s o r b o tt o m o f a n a qu a r ium t o p a r t s o f w a t e r pl a nt s
, ,

o r h a nging f ro m t h e s u r f a c e film U s ua lly the y a r e n e a r th e to p

.

wh e r e m ore o xyg e n ca n b e o bt a in e d fr o m th e w a t e r tha n a t gr e a t e r

de pth s If a tt a ch e d t o th e b o tt o m th e b o dy i s u su a lly h e ld
.
,

up r ight ; if t o th e s id e s t h e b o dy i s in m os t ca se s hor izo nt al

, ,

t h e hyp o s t o m e g e n e r a lly b e ing l o w e r th a n t h e f oo t ; a nd if t o

th e s u r f a ce film t h e b o dy i s a ll o w e d to h a ng di r e ctly d o wnw a r d
,
.
1 28 AN I NT R ODU C T ION TO ZOOLOGY

Susp e nsi on fro m the sur fa c e fi lm m a y be comp a r e d with tha t o f a

ne e dle pl a c e d o n th e su r fa c e o f t he wa t e r Thr e a d s o f a
g e l a tin o us subst a nc e e xte nding o u t fro m the b a sal di sk he lp
, ,

su st ain th e b o dy whi l e in so m e c a s e s a l a rge a i r bubbl e a tt a che d

,

t o th e f o o t k e e ps th e a nim a l a flo a t T h e p ositi o n o f r e st
in e ve r y c ase g i ve s th e Hydr a th e g r e a t e s t o pp o r tuniti e s f or cap
tur ing foo d sinc e in thi s c o n di tio n it h a s co ntrol o f a l a rg e am o unt
,

o f t e r r it or y .

FI G 5 9
. . Spo nt a ne o u sh a nge s o f po s itio ns i n a n u nd i s t u rbe d Hydr a Side
c .

vie w T he e xte nd e d a ni m a l
. co nt r a c t s be nd s to a ne w po s iti o n
a nd th e n e xt e nd s ( Fro m Je nni ngs ) .

SP ON TAN E OU S M OVE M E N T S .
-
All t h em o ve m e nts o f Hydr a
a r e th e r e s ult o f t h e e xp a n s i o n or c o nt r a cti o n o f t h e mu s cl e fib e r s ,

a nd ar e p ro duc e d by t w o kind s o f s timuli int e r n a l or sp o nt a n e

, ,

ous a nd e xt e r n a l
, Sp o nt a n e o u s m o v e m e nts m a y b e o b se r ve d
.

whe n the ani m a l i s a tt ache d a nd undi stur b e d At int e r val s o f .

s e ve ra l minut e s th e b o dy o r t e nt a cl e s o r b o th co nt ra ct s udd e nly

, , ,

a nd ra pidly a nd th e n Sl o wly e xp a nd in a ne w di r e cti o n

,
Hung ry
.

sp e cim e n s a r e m o r e a ctive th a n we ll f e d indi vidual s Th e r e s ult

- .
1
1 3 0 AN IN TRODU CTION To Z OOLOGY

t r a ct s ( Fi g 60 It th e n e xp a nds
.
,
b e nd s o v e r in s o m e o th e r
di r e cti o n a nd a tt a ch e s i t s foo t T h e t e nt a cl e s no w l o ose n
the i r h old a nd a n up r ight p o siti o n i s re ga in e d T h e wh ol e
p r o c e ss h as b e e n like n e d t o th e l ooping l o co m o tio n of a m e a sur
ing w or m At o the r tim e s th e a nim al m o ve s f r o m pl a c e t o pl a ce
.

whil e inve r te d by using i ts t e nt a cl e s a s l e gs Lo co m o ti on m ay .

a l so r e sult f rO m th e gliding o f t h e f oo t a l o ng th e s ub s t r a tum a nd ,

c o n side rabl e di s ta nce s a r e so m e tim e s co ve r e d in thi s w a y .

R E AC TI ON S To E X TE R N AL ST I M U LI — THI GM OTR OP I SM
. .

Hydr a r e a ct s t o v a r i o u s kind s o f sp e ci a l stimuli R e a cti o n t o co n .

t a ct a cco unt s f or i ts t e mp o r a r y fix e d co n di ti o n T h e a tt a chm e nt.

whi l e in th e r e s ting a ttitude i s a r e s ult o f thi s r e a cti o n a nd no t a ,

re sp o n se t o g r a vity s in ce Hyd r a s h a v e t h e l o ngitudin a l a xi s o f t h e

,

b o dy di r e ct e d a t e v e r y p ossibl e a ngl e r e ga r dl e ss o f th e f or ce o f
g r a vity M e cha ni ca l sh ocks s uch a s th e j a rring o f the wa t ch
.
,

gl a ss co ntaining a Sp e cim e n o r th e a git a ti o n o f th e s u r fa ce o f

,

t h e wa t e r ca u se a r a pid co nt r a cti o n o f a p a r t o r a ll o f t h e a nim a l

,
.

This i s foll o we d by a g radu a l e xp a n si o n until th e o ri gin al co ndi

ti o n i s r e gained .

M e ch a nic a l s timuli m a y b e loca li ze d o r non loca li ze d Tha t - .

jus t n o t e d i s o f th e l a tte r typ e Loca l s ti mu la ti on m a y b e a cco m

.

p li s h e d by t o u ching t h e b o dy o r t e nt a cl e s with t h e e nd o f a fi ne

gl a ss r o d T h e r e a cti o ns t o l o ca l s timuli v a r i o u sly a ppli e d h av e

.

b e e n cl assifi e d a s foll o ws
A . Stimul a ti o n b o dy
of

1 We a k : b o dy p a r tly co nt r a ct s
. .

2 M e di um : b o dy co mpl e t e ly co nt r a ct s
. .

3 St.r o ng : b o dy a n d t e nt a cl e s co nt r a ct .

B . Stimul a ti o n o f a t e nt a cl e
I We a k : t e nta cl e s timul a te d co nt r a ct s
. .

2 M e dium : a ll t e nt a cl e s co nt r a ct
. .

3 St.r o ng : t e nt a cl e s a nd b o dy co nt r a ct ( 1 2 3 p ,
.

It sh o uld b e n o t e d th a t th e s timul a ti o n Of o ne t e nt a cl e m ay ca u se
t h e c o nt r a cti o n o f all t h e t e nt a cl e s ( B ,
o r e v e n t h e co nt r a c
 HYDR A AN D C ( ELEN T E RAT E S I N G EN E R AL 1
I 3

ti o n o f b o th te nt a cl e s a nd b o dy ( B Thi s s ho ws tha t the re

,

mus t b e s o m e s or t o f t r a nsmi ssi o n o f s timuli fr o m o ne t e nta cl e

t o a n o th e r a nd to th e b o dy T h e s t r uctu r e o f th e n e r v o u s sys
.

t e m w o uld m a k e thi s p os sibl e ( s e e p 1 2 5 ) .

P H OT OTR OP I S M —
.The r e i s no de fini te r e sp o n se to light al ,

tho ugh the fin a l r e s ult i s quit e d e ci s ive I f a di sh co nta ining Hy

.

dr a s i s pl a ce d s o th a t th e illumin a ti o n i s un e qu a l o n di f fe r e nt sid e s ,

th e a nim a l s will c o ll e ct in t h e b r ight e s t r e gi o n unle ss th e light ,

is too s t ro ng in which ca se the y will c o ng r e g a te in a pl a ce whe re

,

t h e light i s l e ss int e n s e Hyd r a th e re fo r e h as a n o ptimum with

.

r e g ar d to light T h e m o v e m e nt int o o r o u t o f a ce r t a in a re a i s
“
.

a cc o mpli s h e d by a m e th o d o f t r i a l a nd e r r o r ”
Whe n put .

in a da r k pl a ce Hyd r a b e co m e s r e s tl e ss a nd m o v e s a b o ut in no
d e fi ni te di re cti o n ; but if w h ite light i s e n co unte re d i ts l o co m o ti o n ,

b e co m e s l e ss r a pid a nd fin a lly ce a se s a lto g e the r T h e va lue to .

t h e o rg a ni s m o f s uch a r e a c ti o n i s quit e imp o r t a nt s inc e th e sm a ll ,

a nim a l s th a t se r v e a s f o o d f o r it a re a tt r a c t e d to w e ll light e d a r e a s - .

C olor e d li ghts h a v e th e sa m e e ff e ct a s d a r kn e ss ; blu e h o w e v e r , ,

i s p r e f e rr e d by Hyd r a t o whit e .

T H E R M OTR OP I SM T h e r e a cti o ns o f Hydr a to ch a ng e s in

.
- .

t e mp e ra tu re a re al so ind e fini t e a ltho ugh in m a ny ca ses the y

,

e n a bl e th e a nim a l t o e s ca p e f r o m a h e a t e d r e gi o n N o l o co .

m o to r y ch a nge i s p ro du ce d by t e mp e r a tu r e s b e l o w 3 1 C ; a t °
.

thi s te mp e r a tu re h o w e v e r t h e b a s a l di s k i s r e l e a se d a nd the
, ,

a nim a l t a k e s up a ne w p o s iti o n e ith e r a w a y f r o m th e h e a t e d a re a

o r fu r th e r int o it . In th e f o r m e r c a se th e Hydr a e s ca p e s in th e ,

l a tt e r it m ay e s c a p e if s ub s e qu e nt m o ve m e nt s t a k e it a wa y fro m
t h e inju r i o u s h e a t o th e r wi se it p e r i s h e s
,
Hyd r a d o e s no t m ov e
.

f ro m pl ace to pl ace if th e t e mp e r a tu r e i s l o we re d ; it c o ntra cts

l e ss r apidly a nd fin a lly ce a se s all i ts m o ve m e nt s whe n th e f r e e z
,

i ng p o int i s app r o a ch e d I
E LE C T R OTR OP I S M An a tt a ch e d Hyd ra wh e n s ubj e ct e d t o a
.

w e a k c o n st a nt e l e ct r ic cu rr e nt b e nds to w a r d th e a n o de i ts b o dy
, ,

fin a lly b e co ming o r i e nt e d wi th th e b a s a l di sk t o wa r d th e c a th o de
a nd t h e a nt e r i or e nd t o w a r d t h e a n o d e s id e T h e e nti r e a nim a l
.
1 3 2 AN INTR ODU C TION To Z OOLOGY

the n c ontr a cts In a n ani m al a tt a che d by the te nta cl e s a simil ar

.

b e ndi ng occur s but th e b a sal di sk in thi s c ase i s di re cte d to wa r d

,

th e a n o d e . The s e re a cti o n s a re c a u se d by l o c a l c o ntra cti o n s o n

th e a n o d e s id e f or whi ch th e e l e ct r ic cur r e nt i s di r e c tly r e spon
s ibl e ( 1

Hydr a sh o ws no r he otr opi c r e acti ons Wh e n pl a ce d in a cu r .

r e nt o f w a t e r it n e ith e r o r i e nt s it se lf in a d e fi n i te w a y no r m o v e s
e ith e r up o r d o wn s t re a m

G E N E RA L R E MAR K S ON THE B E H AV I OR OF H YDR A It i s e vi .

d e nt fro m the a b o v e o utlin e o f th e r e a cti o n s o f Hydr a to s timuli

th a t th e o nly m o ve m e nts involve d a re p r o duc e d by co ntra cti o n
a nd e xp a n si o n o f t h e b o dy wh e n a tt a ch e d a nd by undi r e ct e d ,

ch ang e s o f p o siti o n B e ing r a di ally symm e tric al th e b o dy m ay

.
,

b e fl e x e d in a ny di r e cti o n .

Local sti mu li s u ch a s the a ppli ca ti o n o f h e a t o r a ch e mic a l to a

,

limit e d a re a of th e b o dy c a use s a co nt r a cti o n o f the p ar t a ffe cte d

,

a nd a b e nding in th a t di r e cti o n Thi s r e s ult s in th e m o ve m e nt

.

o f th e t e nt a cul a r r e gi o n t o w a r d t h e s timuli a nd t h e co nt r a cti o n ,

o f t h e e nti re a nim a l f o ll o w s thu s ca r r ying it o u t o f th e influ e nc e

,

o f th e s timulu s .

N on4locali ze d s ti mu li s uch a s th e j arr ing o f th e v e s s e l c ont a in

,

ing th e ani m al s p ro duce s imme di a t e ly a c ontr a cti o n o f the

, , ,

e nti r e b o dy whi ch in m os t c a se s i s b e n e fici a l Sinc e it re m o v e s it

, , , ,

f ro m an injuri o u s a g e nt If ho we v e r this simpl e c o ntr a cti o n

.
, ,

i s no t e ff e ctiv e a s in th e c a s e o f a c o n s t a nt a pp li c a ti o n o f h e a t
, ,

th e Hydr a usu a lly re s or t s t o s o m e o th e r r e a cti o n e g l o c o m o ti o n ,

. .
,

which Oft e n e na bl e s it to e sc a p e fro m th e injur io us s timulus .

Fin a lly it sh o uld b e r e m e mb e re d th a t th e phy s i ol ogic a l c o ndi

,

ti o n o f th e a nim a l d e te r min e s t o a l a r ge e xt e nt th e kind o f r e a c

ti o ns p ro duc e d no t o nly sp o nta n e o u sly but a l so by e xt e r n al
, ,

s timuli . It de cid e s whe the r Hyd r a sh all c re e p upw ar d to th e

sur fa ce a nd to wa r d th e light or Sh all sink t o th e b o tt o m ; h ow it
,

shall r e act to ch e mi ca l s a nd t o s olid o bj e ct s ; wh e th e r it Sh all

r e m a in qui e t in a ce r t a in p o siti o n o r Sh a ll r e v e r s e thi s p o s iti o n
,

a nd und e r ta k e a l a b o r i o us t o ur o f e xpl ora ti o n ( 1 1 0 p ,

.
1 34 AN INT R ODU C TION TO ZOOLOGY

co nn e ct e d with th a t Of th e p a r e nt Wh e n full gr o wn th e bud .

,

b e c o m e s de t a che d a nd l e a ds a s e p a r a t e e xi s t e nce T h e d e ta il s .

o f th e p r oce ss a r e b r i e fly a s f oll o w s T h e int e r s titi a l c e ll s in a

.

Ce r t a in r e gi o n incr e as e in numb e r a nd v o lum e p r o du cing a Slight ,

o utbulging Of th e e ct o d e r m T h e g ro wing r e gi o n i s l o ca t e d a t
.

t h e p o int wh e r e t h e e dg e s o f t h e p r o t r u si o n m e e t t h e b o dy w a ll .

He r e t h e c e ll s a r e w e ll f e d a nd multiply a ctiv e ly T h e e ct o d e r m .

a nd e nt o d e r m c e ll s o f th e p a r e nt giv e r i se t o t h e c o r r e sp o nding

ce ll s o f t h e bud . Wh e n th e bud i s fully g r o w n th e e ct o de r m ce ll s ,

a t i t s p r o xim a l e nd s e cr e t e a s ti cky su b s t a n ce w hi ch i s u se d l a t e r

f or i t s a tt a chm e nt . T h e e nt o d e r m ce ll s in t h e sa m e r e gi o n th e n
u ni t e se p a r a ting t h e ca vity o f th e bud fr o m tha t o f th e p a r e nt
, .

F in a lly t h e bud b e co m e s d e t a ch e d
,
T h e f oo d s upply d e t e r min e s
.

th e r a t e o f g ro wth o f t h e bud a nd a bud m a y b e e nti r e ly a bs or b e d

,

by a s t a r ving a ni m a l
SE X U AL R E P R ODU C TI ON Wh e th e r or no t th e r e a re d e finit e
.

ge r m c e ll s in th e a dult Hydr a i s s till o p e n t o qu e s ti o n So f ar .

a s i s kn o wn ,
b o th o v a a nd sp e r m a t o zoa a ri se f r o m indi ff e re nt
int e r s titi a l ce ll s
.

SP E R M A OG E N E S I S
T — T h e m a l e ce ll s o f Hyd r a a r e fo r m e d in
.

littl e co ni ca l e l e va ti o n s call e d te s te s whi ch p ro j e ct f r o m th e su r f a ce

o f t h e b o dy ( Fi g 5 4 y t m
.
,
. T h e t e s ti s a r i se s withi n th e e ct o
.
,
.

d e r m fro m int e r s titi a l ce ll s A singl e int e r stiti a l ce ll divide s

.

mi t o ti ca lly ; th e n a dj a ce nt int e r s titi a l ce ll s a l so divid e multi ,

pli ca ti o n co ntin u ing until th e e ct o de r m b e co m e s di st e nd e d An .

inde finit e numb e r o f l o ng mul tinu cl e a t e d cys ts ( Fi g 6 2 A) a r e

'

,
.

f o r m e d withi n th e t e sti s e a ch cys t b e i ng t h e p r o du ct o f a Singl e

,

o r s e v e r a l int e r s titi a l ce ll s E a ch int e rs titi a l ce ll i s a p rim o r di a l

.

g e r m ce ll ; it giv e s r i se by mit osi s t o a v a r i abl e numb e r o f Sp e r

m a t o go ni a whi ch co nt a in t h e s om a ti c numb e r o f ch r o m o som e s
, ,

tw e lv e R e ducti o n in th e numb e r o f ch ro m o so m e s to si x o ccu rs

.

ju s t a fte r t h e sp e r m a t o go ni a ha ve divid e d to fo r m th e p r im ary

sp e r m a to cyt e s ( Fi g 6 2 A b) T h e l a tt e r giv e r i se to s e co nd ar y
.
, ,
.

sp e r m a t o cyt e s (c) whi ch divid e a t o nce p r o du ci ng sp e r m a tids ,

( ) Th s sp m t cyt divi i ns t k pl c with ut h e

‘

c . e e t w o e r a o e s o a e a e o t
 HY D R A AN D C ( ELENT E R AT E S I N G EN ERAL

for m a ti o n of ce ll w alls i e e a ch p rim ar y sp e r m a to cyte d e ve l ops

,
. .

into a fo ur nu cl e a te d c e ll whi ch r e p r e se nt s the fo ur sp e r m a tids

- .

Within thi s ce ll t he sp e r m a tids tra nsfor m into sp e r m a to zo a

( g
F i . 6 2 ,
B ) A .s ingl e cy s t m a y c o nt a in
r e p r e se nt a tiv e s o f a ll o f th e s e c e ll g e n e r

a ti o ns
— sp e r m a t o g o ni a p rim ar y sp e r ,

m a t o cyt e s ,
s e c o nd ar y sp e r m a to cyt e s ,

sp e rm a tid s a nd sp e r m a to z o a T h e
,
.

m a tu r e sp e r m a t o z o a b r e a k o u t o f th e
v e sicl e in whi ch the y a r e for m e d a nd ,

swim ab o ut in th e di s ta l e nd o f the cyst

( g
F i. 6 2 ,
A d );
,
th e y fin a lly re a ch t h e

o ut sid e by w a o f a m i nut e t e mp o r a r y
y
o p e ning in th e e nd o f th e cys t The .

m a tu re sp e r m a t o zoa swim a b o ut in th e
wa t e r s e a r ching f or a n e gg ; the i r a cti v
ity c o ntinu e s fr o m o ne to th re e d a ys .

OOGE N E SI S — T h e e gg i s fi r s t d i s
.

t i ngu i sh e d f ro m th e int e r s titi a l c e ll s o f

th e e ct o d e r m by i t s s lightly g re a t e r s iz e ,

i ts sph e ri ca l s h a p e a nd t h e c o mp a r a
,

t i v e ly l a r g e v o lum e o f i ts nucl e u s As .

th e e gg s g ro w in s iz e t h e n e ighb o r ing
inte rs titi al ce ll s inc r e a se in numb e r by
mitosi s a nd a l so b e co m e l a r ge r T h e
, .

whol e st r u t tu r e m ay a t thi s tim e be

F G 6 P t f t ti f
c all e d an o v a r y ( Figs 63 A ; 54 y .
, ,
.
I . 2 . ar s o a es s o

H yd A i ngl y t ra a s e c s
T h e n ou r i shm e nt o f t h e e gg i s a t fi r s t
.
,

h wi ng p m t g s o s er a o o
simil a r t o th a t o f t h e o th e r e ct o d e r m
ni p im y p m t a, r ar s er a o

ce ll s but l a t e r t h e int e r s titi a l c e ll s n e a r

, yt ( b) nd y c es ,
s e co ar

it a re e ngulfe d th e i r c o nt e nt s b e co ming
, p m t y t nd s er a oc es a

p a r t o f th e o vum T h e nucl e i o f th e se
.
p m ti d ( ) nd s er a s f ,
a

p m t ( d) : B s er a o z oa
int e r s titi al ce ll s fu r ni sh th e y olk o f th e d v l p i ng
,

p m t e e o s er a o
g ro wing e gg U s u ally o nly o ne e gg i s
.

( Af t T nn z oa . er a

de ve l op e d in a singl e o va r y but so m e th i n B i l B l )
, re u er o . u .
1
3 6 AN INTRODU CTION TO Z OOLOGY

tim e s t wo m ay a ri se a nd co mpl e t e the i r de ve l opm e nt side by side .

In m os t ca s e s ho we ve r whe n two o r m o r e e ggs a re co nta in e d i n

, ,

o ne o v a r y th e i r a dj a c e nt w a ll s di sso lv e a nd o ne o f t h e nucl e i su r
,

Viv e s whi l e th e o th e rs di sint e g r a t e As th e o vum g r o ws it b e com e s .

FI G 63
. . An o va ry ( A) , a nd m ature e gg (B) o f
Hydr a . (1, e gg nu c le us ; p s .
, p se u d o po d i u m
of yo u ng e gg ; e m , e gg m e m b r a ne .
( Af t e r
T a nnr e u th e r i n B o B ul ) i l . .

m e b oid in sh ap e sho wing di s tinct p s e udop odi a ( Fi g 63 A

'

a , .
, ,

the se a r e d ra wn in wh e n it h a s r e a che d i ts full siz e T h e e gg is .

now n e a r ly s ph e r ic a l a nd i s s u rr o und e d by a singl e l a ye r o f e ct o

,

d e r m ce ll s ( Fi g 54 . M a tu r a ti o n the n t a k e s pl a c e
,
T wo .

p ol ar b odi e s a r e f o r m e d t h e fi rs t b e ing l a r g e r th a n t h e s e c
,

o nd D u r ing m a tu r a ti o n t h e numb e r o f ch r o m oso m e s i s r e duc e d

.

fr o m th e so m a ti c numb e r twe lve t o si x Thi s o ccu r s a t th e e nd

, ,
.

o f th e gr o wth p e r i o d N o w a n o p e ning a pp e a r s in t h e e ct o d e r m
.

a nd th e e gg i s f o r ce d o u t fin a lly b e co ming f r e e o n a ll sid e s e xc e pt

,

Wh e r e a tta ch e d to t h e a nim al ( Fi g 6 3 B ) .
, .
FI G 6 4
. . R e ge ne r a ti o n a nd gra f ti ng i n Hyd r a. A, s e ve n-h e a d e d Hydr a m a d e
by s p litti ng d i s t a l e nd s l e ngthwi s e ; B , a p i e ce o f Hydr a r e ge ne ra ti ng

ti i l —
a n e n r e a n m a ; 1 5 , s t a ge s i n thi s pr o ce ss ; C , p a r t o f o ne Hydr a
gra f t e d u p o n a no th e r .
( Fr o m M orga n, A a f t e r T r e m bl e y, B a f t e r
M o r gan C a f t e r
, Ki ng )
.
 HY D RA AN D C OELENT ERATES I N GEN ERAL 1
37

FE RTI LI z ATI ON . F e r tiliz a ti o n

usu ally o ccur s within tw o h o urs .
-

Se v e ra l sp e r ms p e n e tr a t e t h e e gg m e mb r a n e but o nl y o ne e nt e rs ,

t h e e gg its e lf If no t f e r tiliz e d within tw e nty fo u r ho urs th e e gg

.
-
,

b e co m e s ste rile T he sp e r m b r ings a nucl e u s c o nt ai ni ng si x

.

chr o m oso m e s into th e e gg Th e m al e a nd fe m al e nucl e i uni te .

,

for ming the fusio n nucl e us .

EM B R Y OL OG Y C le a vage w h ich no w b e gins i s t o t a l a nd r e gu

.
, ,

la r. A w e ll d e fi ne d cl e a v a g e c a vity is p r e s e nt a t th e e nd o f th e
-

thi r d cl e a va g e i e th e e ight c e ll e d st a g e
,
. . Wh e n th e blastu la is
-
.

co mpl e te d it r e s e mbl e s a holl o w sphe r e with a singl e l aye r o f

,

e pithe li a l c e ll s c o mp o sing i t s w a ll Th e s e c e lls m ay b e c all e d th e .

p rimitiv e e cto de r m B y mito tic di vi si o n the y for m e nto de r m

.

c e ll s whi ch d r op into t h e cl e a va g e c a vity c o mpl e te ly filling it , .

T h e ga s tr u la th e r e f or e i s a s olid sph e r e o f c e ll s di f fe r e nti a t e d

, ,

into a singl e o ut e r l aye r th e e ct o d e r m a nd a n i r r e gul ar ce nt r a l

,

m a ss th e e nt o d e r m T h e e ct o d e r m su rr o unds th e g a s tr ul a with
,
.

t w o e nv e l op e s T h e o ut e r is a thi ck chitin o us sh e ll c ov e r e d
.

with sh arp p r o j e cti o n s ; th e inn e r i s a thin ge l a tin o u s m e mb r a n e .

HATC H I N G T he e mb r yo in thi s c o nditi o n s e p a ra t e s f r o m t he

.

p a r e nt a nd f all s t o the b o tto m wh e r e it r e m ains un ch a nge d f or ,

s e v e ra l w e e k s The n int e rs titi a l c e ll s m a k e the i r app e ar a nc e A

. .

sub se que nt re s ting p e r i o d i s f oll o w e d by th e b r e a king a w a y o f

th e o ut e r c hi tin o u s e nv e l o p e a nd t h e e l o ng a ti o n o f th e e sc ap e d
e mb r y o M e so gl e a i s now s e c r e t e d by th e e ct o d e r m a nd e nt o
.

d e r m ce lls ; a ci r cl e t o f t e nt a cl e s a r is e s a t o ne e nd a nd a m o uth
a pp e a r s in th e i r mid s t T h e y o ung Hydr a thus f o r m e d s oo n
.

g ro ws into th e a dul t co nditi o n .

R e ge ne r a ti on — An a cco unt o f th e ph e n o m e n o n o f r e g e n e ra
.

ti o n i s app r op ri a t e a t thi s pl a ce s ince the p o w e r o f a nim als to ,

re s t o r e l os t p a r t s w a s fi r s t di sco v e r e d in Hydr a by T r e mbl e y

in 1 740 Thi s inv e s tig a t o r fo und th a t if Hydr a s w e r e cut int o

.

t w o th r e e o r f o u r pi e ce s e a ch p a r t w o uld g r o w int o a n e nti r e

, , ,

a nim al Oth e r e xp e r im e nt a l r e s ult s o bt a in e d by T r e mbl e y a r e

.

th a t th e hyp o s t o m e to ge th e r with t h e t e nt a cl e s if cut o ff p ro duce , ,

a ne w individu a l ; th a t e ach pi e c e o f a Hyd r a split l o ngitudi nally

I 3 8 AN INT R ODU C TION TO Z OOLOGY

int o tw o o r fo ur p ar t s b e c o m e s a p e r f e ct p olyp ( Fi g 64 A)
, .
, ,

th a t wh e n th e he a d e nd i s split in t w o a nd th e p a r ts se p ar a te d
slightly a tw o h e a de d ani m al r e s ult s ; a nd th a t a sp e cim e n whe n
-

t ur n e d in side o u t i s abl e to r e a dj us t itse lf to ne w co nditi o n s fo r c e d

up o n it .

R e ge ne r a ti on m a y b e d e fin e d a s th e r e pl a cing o f a n e nti r e
o rg a ni s m by a p a r t o f t h e sa m e It t a k e s pl a ce no t o nly in
.

Hydr a but in m a ny o th e r C oe l e nt e r a t e s a nd in so m e o f t h e re p
, ,

r e se nt a ti v e s o f a lm os t e v e r y phylum o f t h e a ni m a l kingd o m .

Hydr a h o w e v e r i s th e sp e ci e s th a t h as b e e n m os t wide ly us e d
, ,

f or e xp e rim e nt a ti o n P i e c e s o f Hydr a th a t m e a s u r e
. mm o r .

m or e in di a m e t e r ar e c ap abl e o f b e co mi ng e nti r e a nim al s ( Fi g .

64 B),
. T h e ti s s u e s in s o m e c ase s r e s t o r e th e l os t p ar t s by a
multiplic a ti o n o f the i r ce ll s ; in o the r case s th e y ar e wo r k e d o v e r ,

di r e ctly into a ne w but s m all e r indi vidu al .

GR AF TI N G .P a r t s o f o ne Hydr a m a y e a sily b e g ra ft e d up o n
a n o th e r ( Fi g 6 4 C )
. In thi s w a y m a ny biz a r re e ffe ct s h ave b e e n
,
.

p r o du ce d P a r ts o f tw o Hydr as o f t wo sp e ci e s h av e al so b e e n
.

succ e ss fully unite d .

GE N E R A L R E M AR K S ON R E GE N E R AT I ON Sp ace will no t pe r .

mit a de t a il e d a cco unt o f th e m a ny int e r e s ting qu e s ti o n s inv olv e d

in th e phe n o m e n a o f r e g e ne r a ti o n but e n o ugh h a s b e e n give n to
,

indi ca t e th e n a tu r e o f th e p ro ce ss T h e b e n e fit t o th e a nim al o f
.

th e a bility t o r e g e n e r a t e l o s t p a r t s i s o bvi o u s t o all Such a n .

a nim a l in m a ny ca se s will s u cce e d i n t h e s t r uggl e f or e xi s t e n ce

, ,

und e r a dve rse c o nditi o ns T h e r e ge n e r a ti o n o f th e e a r thw o r m

.

a nd t h e cr a yfi s h a r e c o n s id e re d in C h a pt e r s X a nd XI It will .

s u ffi c e h e r e the r e fo r e to sa y a f e w w o r ds c o n ce r ning r e g e ne r a
, ,

ti o n in ge n e ral R e g e n e r a ti o n t a k e s pl a ce co ntinu ally in all

.

a nim a l s ; f or e x a mpl e ne w ce ll s a r e p r o du ce d in t h e e pid e r mi s o f

,

m a n t o t a k e th e pl a ce o f th o se th a t a r e no l o ng e r a bl e t o p e r fo r m
the i r p rop e r fun cti o ns B o th int e r n al a nd e xt e r n a l f a ctors h ave
.

a n influ e n ce up o n t h e r a t e o f r e g e n e ra ti o n a nd up o n t h e ch a r a ct e r

of th e ne w p a r t Te mp e r a tu r e fo o d light g r a vity a nd co nt a ct
.
, , , ,

a r e s o m e o f t h e e xt e r na l f a ct o r s I n m a n v a ri o u s tis s u e s a r e
.
,
1 40 AN IN T RODU CTION TO Z OOLOGY

FI G 6 5 P a rt o f a co l o ni a l Hyd ro z oa n, Obe li a
. .
, ( A) , a nd a f re e -sw i m m i ng
m e d u sa ( B ) f r o m a no th e r h yd r o i d co l o ny I ,
t
e c ode rm

2, e tod e r m ; 3 m o u th ; 4
n , ,
l nt e r on
coe e 5, coe no s arc ; 6 , pe r i sa r c ;
7 h yd ro th e ca ; 8 b l a s t os tyl e
, , ; 9 , m e d u sa b u d ; I O, go no the ca .
( Fr o m
Shi pl e y a nd M a cB rid e ) .
 HY D R A AN D C (ELEN T E R AT E S I N GEN ER AL 1 41

fr o m th e e xtr e m e ri m T h e middle l a ye r th e m e s ogl e a i s in

.
, ,

th e m r e m a r k ably thick a nd r e se mbl e s j e lly h e nce the i r n a me

, ,
.

Hyd r a a nd J e llyfi s h e s C o m pa r e d — Alth o ugh th e m e dus ae .

up o n s up e rfici al e x a m in a tio n app e a r to be v e ry di ffe re nt fro m th e

p olyps or zooids the y a re c o nstr ucte d o n the s a me g e ne r al
,

FI G 6 6
. . D i a gr a m s
h o wi ng th e s i m il a r iti e s o f a po lyp ( A) a nd a m ed us a
s
‘ ’
f

'

ci r c c i r cu l a r ca na l ; e ct
.
, e c t o d e r m ; e nd
.
,
e nt o d e r m ; e nt ca v ga s t ro
,
. .
,

v a s cu l a r ca vity ; hyp hypo s t o m e m nb m a nu br i u m ms gt m e so gl e a

.
, .
,
.
,

mtk m o u th ; m
.
,
ne r v e r i ngs ; r ad r a d i a l ca na l ; v v e l um
.
, .
, ( Fr o m .
,
.

P a rk e r a nd Ha s w e ll ) .

pl a n a s th e l a tt e r Figu r e 6 6 illu s t r a te s in a di a gra mm a tic

.

fa shi o n th e r e se mbl a nc e s b e tw e e n th e p olyp ( A) a nd th e m e dusa ,

( B ) by m e a n s o f l o ngitudin al s e cti o n s If th e m e dus a w e re .

g r a sp e d a t th e c e nt e r o f the ab oral s u r fa ce a nd el o ng a te d a hydr a .

like fo r m w o uld r e s ult B o th h a v e s imil a r p a r t s the m o s t no

.
,

ti ce a ble di ff e r e nc e b e ing th e e n o r m o u s qu a ntity o f m e so gl e a

p re se nt in th e m e dus a .

M e ta ge n e s i s — I n s o m e Hyd r o zo a th e r e a r e tw o kinds o f
.

individu al s b e l o ngi ng t o th e sam e sp e ci e s ; one o f th e se in th e ,

fo r m o f a p o lyp give s r i se a se xua lly by budding to th e se cond

,

fo r m th e m e dusa whi ch p r o du ce s e ggs a nd sp e r m s T h e f e r

, ,
.

ti li z e d e gg d e v e l op s int o th e p o lyp T h e p o lyp o r hyd ro id s t a ge

.
, ,
1 42 AN INT RODU CTION TO ZOOLOGY

is m ore p r on o unce d in so m e sp e ci e s th a n in o the r s e g Hydr a , . .

,

h as no m e du sa s t a g e a t a ll ; wh e r e a s c e r t a in sp e ci e s h a ve no
p olyp stage but p ass the i r e nti r e e xi s te nc e a s m e dus ae Vari o us .

co nditi o n s m ay be illu s tra te d by di ffe re nt Hyd r o zo a In th e .

f oll o wing t abl e 0 r e p r e se nts t h e f e r tiliz e d o vum ; H a p olyp ;

, ,

M a m e dus a ; m an inco n spicu o us o r d e g e n e r a t e m e d usa a nd

, ,
,

h an incon spicu o us or de g e n e ra t e p olyp

,

2 . O H— m — O H — m — O (S e r tu lar i a) .

3 . O H M O H M O (Obe li a) .

4 . O h M O h M O ( Li r i ope ) .

5 . O M O M O

Th e lte r n a tio n o f a se xu al with a n a s e xu al ge n e ra ti on as in

a
,

e x a mpl e s 2 3 a nd 4 j u s t li s t e d i s kn o wn a s me ta e ne si s This
, , , , g .

phe n o m e n o n o ccu r s in o the r g ro up s o f th e a nim al ki ngdo m but ,

fi nd s i ts b e s t e x a mpl e s a m o ng th e C oe l e nte r a te s .

Di vi s i o n o f Lab or a m on g C oe l e n te r a te s N o t o nly h a v e th e
.

s o m a tic ce lls o f th e C oe l e nte ra t e s b e co m e di ffe r e nti a t e d into e cto

de r m a nd e nt o d e r m in e a ch o f whi ch ce ll s m ay b e r e cognize d
,

h a ving p a r ti cul a r functi o n s to p e r for m ; but i n ce r t a in g ro ups

col o ni a l sp e ci e s a re fo und in whi ch th e v a ri o us m e mb e r s o f th e
c ol o ny a re s o sp e ci aliz e d f o r c e r ta in kind s o f w o r k th a t th e y are ,

incap abl e of ca rr ying o n o th e r p ro ce sse s P e rhaps th e b e s t .

e x a mpl e o f s uch a co n di ti o n i s P hys a li a th e P o r tugue se M a n

,

o Wa r
f —
( g
F i . P hy s a li a i s a c o l o ni a l H yd ro z oa n co n si s t
ing of a l arge fl o a t (1m ) with a s ai l -like cr e s t ( cr ) fro m whi ch a .

n u mb e r o f p olyp s h a ng do wn into th e wa te r So m e o f the se .

p olyps a re nut ritiv e o th e rs a r e ta ctile ; so m e co nta in b a tt e ri e s

,

o f n e m a t o cy s ts o th e r s a re m a l e r e p ro ductiv e z o o id s
,
a nd still ,

o th e rs gi ve r i se t o e gg p ro ducing m e dus a
-

C or als . One g ro up o f C oe l e nt e ra t e s th e co r a l s i s o f e sp e ci al
, ,

inte r e s t b e ca u se o f i ts e co n o mi c imp o r ta nce T h e co r a l s ar e .

f o und p rin cip ally in th e t r opic s The y liv e n e a r th e sh o r e whi ch

.
,

no t inf r e qu e ntly is built up o f cor a l sk e l e t o n s T h e h ar d p a r ts

.
 FI G 6 8
. . Asso ci a ti o n of a h e r m it f
c a b a nd a

C oe l e nt e r a t e ( Fro m P a r k e r a nd Has
w e ll ) .

FI G 6 7
. . P hys a li a , th e Por tu
gue se m a n-o f -w a r . cr .
,

cr e s t; p,.
po yp ;l pm ,
pne u m a t o ph o r e .
( Fr o m
P ar k e r a nd H a s w e ll t
af e r

Huxl e y ) .
 HY DR A AN D C (ELEN TE R AT ES I N G EN ER AL 1 43

of coral s a re co mp ose d o f ca lcium ca r b o n a t e e xc re t e d by the e cto

d e r m ce ll s o f th e p olyp s So num e ro u s a re the se p olyp s tha t m a ny
.

i sl a nds in th e P a cific Oce a n a nd m a ny r e e fs n e a r o the r i sl a nds co n

s i s t e nti r e ly o f c o r a l r o ck T h e p r e ci o u s R e d C or a l i s f o und o nl y
.

in the M e di te rra ne a n Se a .

Sym b i os i s Symbi osi s m e a n s a n intim a t e a nd a dva nt a g e o us

.

a ss o ci a ti o n b e tw e e n t w o kinds o f o r g a ni s ms T h e m os t co mm o n
.

e x a mpl e i s th e li ch e n whi ch c o nsist s o f t w o pl a nt s a n a lg a a nd a

, ,

fungus Hydr a vi rid i s th e gr e e n Hydr a d e r ive s i ts co l or f ro m a

.
, ,

g re a t numb e r o f uni ce llul a r g re e n pl a nts the Z ooe hlor e llw which , ,

o ccupy t h e b a sa l p o r ti o n o f th e e nt o d e r m ce ll s The se g re e n .

a lg ae m a nuf a ctu r e st a r ch in th e p r e s e n ce o f light a nd du ring , ,

thi s p ro ce s s lib e r a te o xyg e n whi ch i s o f a dv a nt a ge to the p olyp

,
.

P r ob a bly th e Hydr a a l so o bt a in s fo o d f ro m th e se a lg ae T he .

s e cu rity a nd ca r b o n di o xid e fu r ni s h e d by t h e p ro t e cting c e ll s o f

th e p o lyp co mp e ns a t e th e a lg ae f o r th e f o o d a nd o xyg e n th e y
p ro vide .

A m o st c o mpli ca t e d illu s t r a ti o n o f s ymbi o si s i s th a t o f ce r t ain

he r mi t cr a bs with se a a ne m one s T h e h e r mit c ra b live s in the
.

she ll o f a l arg e s alt wa t e r sn ail As soo n a s a s ui ta bl e sh e ll i s

- .

fo und th e he r mit cr ab t a k e s p o sse s si o n It th e n hunts ab o ut

, .

until it finds a se a a n e m o n e whi ch it pl a c e s up o n th e s h e ll jus t

,

a b o v e th e o p e ni ng Oft e n th e a ne m o n e co mpl e t e ly co v e r s th e
.

h e r mit crab s h o us e Th e a dva nt a ge t o th e s e a a n e m o ne in thi s

’
.

p ar tn e r ship li e s in th e g r e a t e r cha nc e s it h a s f o r p rop e r f oo d

c o nditi o ns sin ce it i s ca rr i e d ab o ut f r o m pl a ce to pl ace by th e
,

c r ab T h e b e n e fit s to th e l a tt e r a r e o f a p e culi a r ch a r a ct e r
. .

F igur e 6 8 sh o w s a C oe l e nt e r a t e c o l o ny H yd r a cti nza co n

’

, ,

sisting o f nut r itiv e p o lyps with t e nt a cl e s r e p r o du ctiv e individu a l s

,

b e a r ing a ci r cl e o f m e du s o id bud s s pin e -lik e p r o t e ctiv e m e mb e r s

, ,

a nd b o r d e ring th e e dg e o f th e s h e ll a r o w o f th r e a dlik e d e f e n s iv e
,

p olyp s p ro vid e d with s tinging ce ll s Whe n th e h e rm it cra b i s .

a tt ack e d th e se s tinging c e ll s a r e s h o t int o t h e e n e my w hi ch is

, ,

thus f re que ntly dr iv e n a w a y In thi s w ay th e C oe l e nte ra te b e ne fits

.

its associ ate .

 C HAP T ER IX

S P ON G E S , F LAT WOR M S ,
AN D R OU ND WORM S 1

1 . SP ON GE S GR AN TI A

(Gr a nti a e i lza ta Fl e m )

'

Gr a nti a ( Fi g . simpl e sp onge i n

6 9) i s a

h abiting th e salt w a te r a l o ng th e co a s t o f th e
N e w E ngl a nd s t a t e s j u s t b e l o w t h e lo w tide -

m a rk He r e it i s fo und a tt a che d by o ne e nd
,

t o r ock s a nd o th e r s o lid o bj e ct s Unlik e .

Hydr a , Gr a nti a is p e r m a ne ntly a ttache d ,

n e v e r m o ving fr o m pl a c e to pl ace a s a n a dult .

I ts di s t r ibuti o n in sp a c e i s e ffe ct e d du ring

th e e a r ly e mb r y o nic st a g e s a t w hi ch tim e ,

cili a a r e p r e se nt e na bling it to s wim a b o ut ,

.

Gr a nti a v a r i e s in le ngth fr o m o ne h a lf a n
in ch to alm o s t a n inch a nd r e se mbl e s in ,

sh ap e a sl e nde r v a s e th a t b ulg e s slightly

FI G 6 9 A
. im pl
.
n e ar
s
t h e ce nt e r
e
T h e di st a l e nd . o f th e

p n
sg o ( Af t ae nim
.
a l o p e n s
er t o t h e e xt e r i or by a l ar g e e x

M i n hi n i n L n cu r r e nt or e th e os cnlnm Thi p e ni ng i s
c
p a
,
s o .

k t
’
T
es er s
surro unde d o n a ll s ide s by a ci r cl e t o f l o ng
re a

ti ) se
st r aight n e e dl e s call e d spi cu le s Sm all e r
.

s pi cul e s p ro t r ud e f r o m o th e r p a r ts o f th e b o dy giving the ,

1
Si nce it was i m possi bl e t o i ncl u d e i n this boo k d e t aile d d i scu ssi ons o f
t yp e s f rom e v e ry phyl u m o f t h e a ni m a l k i ngd om m any gro u ps are no t re p ,

re se nt e d Ih thi s ch ap t e r t h re e a nim a l s ar e b r i e fl y d e s cr i be d i n o r d e r th a t
.

th e s t e p f r o m a s i m p l e C oe l e nt e r a t e li k e Hyd r a t o a co m p l e x Anne lid li k e

, , ,

t h e e ar thw o r m m ay no t b e t o o a b r u p t
,
T he s p o nge s fla t w o r m s a nd
.
, ,

r o u nd w orm s po ss e ss ce r t a i n o r ga ns th a t h e l p th e s t u d e nt t o u nd e rs t a nd th e

s t r u c t u re a nd f u ncti o ns o f s i m il ar o r ga ns i n m o re com l e x anim al s

p .
1 46 AN l N TR ODU C T I ON T O Z OOLOGY

of w a te r whi ch e nt e r s t h e incu r r e nt c a n al p ass e s thr o ugh the

, ,

p r OS Opy e s int o t h e r a di a l c a n a l a nd th e nc e int o t h e cl o a c a l

l ,

c a vity fin ally e s caping fr o m th e b o dy by w ay o f th e o s culum

, .

T h e s u r f a c e a re a o f
th e e pith e lium co v e r
ing th e b o dy a nd ,

lining th e int e r n al
c a viti e s i s e n orm o u sly ,

i ncre ase d by the ca n al

m 3e
sys te m
.
.

Gr a nti a is a n a nim a l
p osse ssing a n o ut e r
d e r m a l l a ye r o f ce ll s ,

a n inn e r g a s t r a l e Di

th e lium a nd a middl e ,

r e gi o n c o nt a ini ng c e ll s

of se v e r al v a ri e ti e s
( Fi g . Th e de r
m al e pi the li u m ( g
F i .

) v e r s t h e
7 1 ,
e ct . co

e nti r e o ut e r s u r f a c e o f

t h e b o dy a nd lin e s ,

th e incu r r e nt c a n al s .

It i s co mp ose d e x
t e r na lly o f a singl e
l aye r o f fla t c e ll s .

3 Wh e r e v e r pro sopyle s
nd «
occu r co nn e cti ng t h e
FI G 7
. P t f th b d y f G nti
1 . ar o t e t I n cu rr e nt with th e
o o ra a . ec .
,
ec o

d m ;er nd nt d m ; m e m d m ;
e o er es e so er
r a d i a l ca n a l s a sm gle
.
, .
,

pi pi l
s .
( F m D hl g n nd K pn )
,
s cu e s . ro a re a e er .
’

l arge de r mal cell ,

t e r m e d a porocyt e i s p re se nt T h e por ocyte s a re de r iv e d fr o m

,
.

c e ll s o f the de r m al e pithe lium The y a r e l arg e a nd gra nul a r a nd .

,

fr e qu e ntly e xhi bit a m e b o id m o v e m e nt s T h e p ro sopyl e i s an .

intr a ce ll u l a r p e r for a ti o n o f th e po ro cyte C e ll s call e d scle ro .

,
 SP ON GE S, F LAT WOR M S ,
AN D R OU N D WORM S 1 47

bla s ts , whi ch p r o duce spic u l e s a r e al so co n side r e d co n s titu e nt s ,

o f th e d e r m a l la y e r .

T h e i nne r e pi the li u m lin e s th e cl o a c a l c a vity a nd th e r a dia l

c a n als In the l a tt e r it c o nsi s ts of o ne l a ye r of c o ll ar e d flage l
.

l a t e d c e lls ( Fi g 7 1 t e r m e d choanocyte s N o c oll ar c e lls

.
,
.

a re p re se nt in th e
e pith e lium l i ni ng t h e

cl o a c a l ca vity The .

fl ag e ll a o f t h e coll ar
ce lls c r e a t e th e cu r
r e nt o f w a t e r whi ch
is c ontinu a lly flo wing
thr o ugh th e b o dy w a ll
int o th e cl o a ca l ca vity
a nd out of th e o s cu

lum .

T h e m iddle r e gi on o f

the b o dy w all is no t
so de finit e no r fi r m
in s t r u ctu r e a s a r e th e
o ut e r a nd inn e r e i
p

F I G 7 2 Se cti o n o f a po r ti o n o f Gr a nti a
. . I , o pe n .

i ngs o f i ncu r r e nt ca na l s ; 2 , i ncur r e nt ca na l ;

3 , p ro so py l e ; 4 r a d i a l ca na l ; 5 ch oa no
, ,

cy e s t ; 6 s pi cu l e s ; 7 o p e ni ng o f r a d i a l ca na l
, ,
.

( Fro m Shi p l e y a nd M a cB r i d e a f t e r De nd y ) .

FI G 73 De v e l o p
th e li a but n e v e r th e l e s s it i s co n sid e r e d
. .

i ng p i l f
s cu e s o
, , ,

a p n g s a di s tin
o c t c e llul a
e r l a y e r A m e bo id wan .

( F m D hl d e r i ng ce lls whi ch ing e s t f oo d o r a ct a s

ro a ,
.

g n nd K p
re
s t or a g e c e ll s a r e f o und h e r e
a e a s w e ll a s , ,
n ft A
t h e r e pr odu cti ve ce lls whi ch a lw a ys a r i s e
er a er .

M )
aas
in th e middle l a y e r
.

so ft b o dy w a ll o f Gr a nti a ( F i
T he g 7 2 ) i s s upp o r t e d a nd pr o .

t e cte d by a s ke l e t o n co mp o se d o f a g re a t numb e r o f spi cu le s o f

 1 48 AN INT RODU CTION TO Z OOLOGY

c a r b on a t e of lim e F o ur v ari e ti e s o f spicul e s a re always p r e se nt

.
,

( )
1 l o ng st r a ight m o n a x o n r o d s gu ar di ng t h e o scu l um ( )
2 sh o r t,

straight m o n a xo n r o d s s urr o un di ng th e incurr e nt p or e s (3 ) tri ,

r a di a t e s picul e s a lw a y s f o und e mb e dd e d in t h e b o dy w a ll a nd ,

(4) T - sh a p e d Spicul e s lining t h e cl o a c a l c a vity ; f o u r a n d fi ve

r a y e d spicul e s m ay a l so b e p r e s e nt Spicule s are built up withi n

.

c e ll s c all e d scle r oblas ts ( Fi g . which f or m p ar t of th e inn e r

stra tum o f th e d e r m a l l a y e r A sl e nde r or g a nic a xi a l thr e a d is
.

fi r s t built up within th e c e ll ; ar o und thi s i s d e p o sit e d th e c alc are

o us m a tt e r ; t h e wh o l e Spicule i s th e n insh e a th e d by a n e nv e l op e

o f o r g a ni c m a tt e r lik e th a t c o mp o sing th e a xi a l thr e a d .

N u tri ti on .
— Gr a nti a live s up o n t h e minute o r g a ni sms a nd
sm all p a r ticl e s of o r g a nic m a tt e r tha t a r e dr a wn into th e incu r
r e nt c a n a ls by t h e cu r re nt o f w a t e r p r o du ce d by t h e b e a ting o f

th e co ll ar c e ll cili a
- . So m e o f th e f oo d p ar ticl e s a r e p r o b a bly
ing e s t e d by th e poro cyte s ; but th e m a j o rity o f th e m a r e e ngulfe d
by th e c o ll ar c e ll s Di ge sti on a s in the P r o to z oa i s int r a ce llul a r
.
, , ,

foo d va cuol e s b e ing fo r m e d T h e di s t ributi o n o f th e nut rim e nt

.

i s a cco mpli s h e d by t h e p a ssa ge o f dig e s t e d f oo d f r o m ce ll t o ce ll ,

a id e d by t h e a m e b o id w a nd e r ing ce lls o f t h e middl e l a y e r .

Excr e tor y m a tte r i s di s ch arg e d th ro ugh th e g e n e r a l b o dy su r f a c e ,

a ssi s t e d p r o b a bly by t h e a m e b o id w a nd e ring c e lls a nd p o s sibly ,

by th e c oll ar c e ll s a l so Re spi r a ti on like wi se t a k e s pl a ce in th e

.
,

a b se nc e o f sp e ci a l o r g a ns th r o ugh t h e c e lls o f th e b o dy w a ll
,
.

Sp o ng e s d o no t p o ss e ss diff e r e nti a t e d ne r vou s organs but a r e ,

a bl e to r e s p o nd t o ce r t a in s timuli T h e p o r e s a nd o scul a a r e
.

s u rr o und e d by c o nt r a ctil e ce ll s c a ll e d myocyte s

,
wh i ch a re abl e
,

t o cl os e th e se o p e ning s . App a r e ntly the s e ce lls r e sp o nd t o di r e ct

s timul a ti o n s in ce no n e r v o u s ti s s u e i s p r e se nt
,
Th e y th e r e fo r e
.
, ,

r e p re s e nt wh a t m ay b e co nsid e r e d t h e v e r y b e ginning o f a n e u r o

mus cul ar m e ch a ni s m ( 1 3 0 pp 5 9 ,
.

R e pr o d u cti o n. R e p r o ducti o n in Gr a nti a t a k e s pl a c e by b o th

s e xual a nd a s e xu a l m e tho ds In th e l a tt e r ca se a bu d a ri se s
.
,

n e ar th e p oint of a tt a chm e nt fin ally b e co m e s fr e e a nd t ake s up a

, ,

s e p a r a t e e xi st e n ce
.
1 5 0 AN IN T R ODU CT ION TO ZOOLOGY

o th e r a nimal s ca n b e di s cu ss e d s u cc e ssfully Th e char a cte r o f .

th e o bj e ct to which s p o ng e s a re a tt a ch e d c a u se s th e m t o a ss um e
e xc e e di ngly i rr e gul ar s h a p e s t h e ro ck s b e ing f re q u e ntly inc r u s t e d
,

FI G 7 4 De v e l op m e nt o f a s i m pl e s po nge (S ycon r a pha nus )

. . . a , o v um ;

b, c, o v u m s e gm e nt e d ; d , b l a s t u l a ; e , a m p hi b l a s t u l a ; f , co m m e nce m e nt
o f i nva gi na ti o n ; g, g a s t r u l a a tt a ch e d ; h ,
i , y o u ng p s o nge.
( F r o m
P a rk e r a nd Ha s w e ll t
af e r Sch u lz e .
)
 SP ON GE S, F LAT WOR M S ,
AN D R OU N D WORM S 1 5 1

by inde fi ni te m a ss e s o f sp o ngy tissu e Thi s m a k e s it diffi cult to

.

de cide wh a t co nstitut e s a n individu a l sp o ng e P e rh aps th e b e s t .

w a y t o se p a ra t e o ne f r o m a n o th e r i s t o c o n sid e r a ll o f th e tissu e
s u r r o unding o ne o scul um a s a singl e individu a l .

03 0

FI G 7 5. . T yp e s t ms of
of ca na l sys e

spo nge s T h e a rro w s i nd i ca t e t h e

.

d i r e cti o n o f t h e cu r re nt o f w a t e r .

T h e thi ck b l a ck li ne r e p r e se nt s t h e

ga s t r a l l a ye r t h e d o tt e d po r ti o n t h e
, ,

d e r m a l l a ye r ap p a p py e ;
o . l . .
,

e xcu rr e nt ca na l ;
ex . c .
, fl c . .
,

fla ge lla t e d ch a m b e r ; G C ga s t r a l ,

ca vit y ( cl o a ca ) ; in e i ncu rre nt . .

,

ca na l ; asc o sc u l u m ;
.
,
os t o s ti a ; .
,

pr p pr o so pyl e
. .
, (Af t e r M i nchi n .

i n La nk e s te r s T r e a ti se ) ’
.

C anal Sy s t e m .
— Th e c a n al
syst e ms o f sp o ng e s m ay b e
g ro up e d und e r th r e e typ e s ; the
fi r s t ( Fi g 7 5 A) co n si s t s o f i n
.
,

cu r r e nt p o re s ( p) a g a st r al c a vity ,

(C C) a nd a n o sculum
,
 1 5 2 AN INTRODU CTION TO ZOOLOGY

Th e s e cond typ e ( Fi g 7 5 B) i s m ore co mplic at e d ; th e w a t e r flo ws

.
,

thro ugh t h e d e r m al p o r e s (o sti a) int o th e incurr e nt c a n al s

th e n thr o ugh th e ch a mb e r p or e s (prosopyle s pr p ) into th e r a di a l ,
. .

c a n al s (f l fro m h e re it i s p rop e ll e d by th e fl ag e ll a of t h e
.

choa no cyte s int o th e g ast ra l cavity (GC ) fin ally p as sing o u t ,

thro ugh th e o scul um In t h e thi r d typ e ( Fi g 7 5 C ) the r e .

,

a r e th re e di s tinct p a r t s ( I ) th e w a t e r p a sse s th ro ugh th e d e r m al

,

o sti a ( os t ) a nd by w a y o f incu rr e nt c a n a l s (i nc ) r e a ch e s ( 2 ) a
. .

numb e r of sm all ch a mb e r s (f l c ) lin e d with choa n ocyt e s the nc e it

. .
,

is c arri e d thr o ugh ( 3 ) a n e x cu rr e nt syst e m (cxc ) to th e g ast ral .

c a vity (GC) a nd fin ally ou t of th e o s culum

,
.

Th e s ke le tons o f sp o ng e s a re c omp o s e d o f spicul e s and spongi n .

Th e f or m e r a r e s e c r e t e d by c e lls o f t h e d e r m a l l ay e r a nd c on si s t ,

o f c a lcit e a nd sil ic a Th e sp o ngin i s a n o r g a nic s ub s t a nc e

. .

R e pr o d u cti on R e p ro ducti o n i s e ith e r a se xu al or s e xu al

. .

By th e a se xu a l m e th o d th e re a r e p ro duc e d bud s a nd g e mmul e s .

B ud s m a y b e se t f r e e t o t a k e up a s e p ara t e e xist e nc e o r m ay ,

re m a in a tt a ch e d t o t h e p a r e nt sp o ng e aiding in th e f o r m a ti o n o f
,

a c o mpl e x a sse mbl a g e o f individu a l s Ge mmu le s o ccu r in th e .

f re sh wa t e r sp o nge S pongi lla and se v e ral m arin e sp e ci e s A

-
, ,
.

numb e r o f c e ll s in th e middl e l aye r o f th e b o dy w all g a the r int o

a b a ll a nd b e c o m e s u r r o und e d by p r o t e cting spicul e s The se .

g e mmul e s a r e for m e d in t h e a utumn ju s t b e f or e th e de a th o f t h e

a dul t sp o ng e In th e sp ring the y d e ve l op into ne w sp onge s
. .

The y a re o f v alue in c arrying th e r a ce thr o ugh a p e ri o d of a dve r se

c o nditi o ns s u ch as th e wint e r se a so n S e xu al r e pr odu cti on t ake s
, .

pl a c e e ss e nti ally as de scrib e d f or S ycon o n p age 1 4 9 .

P i e ce s 0 sp o nge s a r e c a p abl e o f r e ge ne r a ti ng e nti r e a nim als

? .

This ch ara ct e ri s tic e n abl e s sp o nge g r owe r s t o pl a nt a b e d o f

-

sp onge s by sca tte ring s m all pi e ce s o v e r th e b o tto m of th e se a in

f a vorabl e pl a c e s Afte r a p e ri o d o f s e ve r al ye ar s a nim a ls o f
"
.
,
”
co mm e r ci a l v alue m ay b e g a the re d a nd a n e w lot of slips ,

se t ou t .

Em b r yology . p rincip al st ag e s
T he of sp onge e mb ry ol ogy
m ay b e p r e s e nt e d b ri e fly as fo ll o ws :
1
54 AN I NT RODU C T ION TO Z OOLOGY

st r ucture s diffi cult to de t e r min e in a living anim al In o r de r to .

s tudy P la na r i a succ e s s f ul ly in th e l a b o r a t o r y th e so ft co nt r a ctil e , ,

b o dy i s u s u ally pl a ce d o n a
s lid e a nd the n p re sse d o u t ,

slightly with a co v e r gl ass .

A p a ir of e ye s pots ( Fi g 7 6 I ) .
,

a re p r e s e nt o n th e d o r s a l s u r

f a ce n e a r th e a nt e r i or e nd .

T h e m ou th i s in a p e culi a r p osi
”

tio n n e a r th e m iddl e o f the

v e ntr a l s ur fa c e F rom it th e .

muscul a r pr obosci s (3 ) m ay
e xte nd P o s te ri o r to th e .

m o uth i s a sm all e r o p e ning ,

t h e ge ni ta l p or e Th e su r

b o dy i s cove r e d with f a ce of th e
ci li a whi ch p ro p e l t h e a nim a l ,

th ro ugh th e w a t e r Thi s i s .

no t th e o nly m e th o d o f l o co ‘

m o tio n sinc e mu scul a r con ,

t r acti on i s al so e ffe ctive .

S tr u ctu r e A s tudy o f th e .

s t r u ctu r e o f th e a dult a nd o f

th e e a rly e mb r y o nic s t a ge s

s h o w s P la na r i a t o b e a tr i plo

blasti c a nim a l p o sse ssing th e

th r e e g e r m l a ye r s e cto de rm , ,

FI G 7 7 A n t m y f P l n n
. . a
m o
e so d e r m a nd e nt o d e r m f r o m
o a a a .
, ,

n b in ;
o ,
ra
y ; g v y; , ,
whi
6, e
c h se
e v e r a l
,
sy
o
s t e m
ar
s o f o r
1’ 2,

i b n h
s, f i nt ti n ; ln l t
ra c l g a ns h a ve
es o es b e e n de r iv e d
e ,
a e ra .

n v ;er m th ; P1 p h ynx ; Th e r e a r e w e ll d e v e l op e d mus

e 772, ou 1, ar -

d
o vi d t ; t t ti ;
,
o uct ; cula r ,
n e r vo u s di ge s tive e x
es s u , u e r us
, , ,
v y l k gl nd ; vd
o d f an ; s va s e e re s
r e p r o d u c ti v e
,
a nd
,
cr e t or y
O mni : Q w gi n ; E
e S " m
s a a , ,
C0
,

m n g nit l p
o ( F
e m L n a
s y s t e m s ; th
or e
e
,
se a r e c o n
ro s t r u c t e d a

k s t s T a ti s af t e r V G ff )
e er
’
re in such a way as to functi o n
e . ra .
 SP ON GE S, FLAT WOR M S ,
AN D R OU N D WOR M S 1
55

witho ut th e coo r din a ti o n of a cir cul a to ry syste m ,

re spir a to ry
sy s te m c oe l o m and a nus
, ,
.

DI G E STI V E SY STE M . dig e s tive sy s te m ( Fi g 7 7) c o n sis ts

Th e .

o f a mou th a pha rynx lying in a muscul ar s he a th a nd ,

a n i nte sti ne o f thr e e m a in t r unks (i i i ) a nd a l a r g e numb e r o f

, 2, 3

s m a ll l a t e ra l e xt e n s i o n s T h e mu s cul a r ph a r ynx ca n b e e x
.

t e nde d as a p r ob o sci s ( Fi g 7 6 thi s f a cilit a t e s the c apture o f

.
,

f o o d Dige s ti o n i s b o th i nte r ce llu la r a nd i ntr ace llu la r i e p a r t

.
,
. .

o f th e f o o d i s di g e s t e d in t h e int e s tin a l t r unk s by s e c r e ti o n s f ro m

ce ll s in the i r w a ll s ; wh e r e a s o the r foo d p a r ticl e s a re e ngulfe d

by p se udop o di a thr ust o u t by c e lls li ni ng th e inte s tin e a nd a re ,

dig e s te d in side o f th e c e ll s in v a cu ol e s T h e dige s t e d foo d i s a b .

s o rb e d by the w a lls o f t h e int e s tin a l t r unk s a nd Since b ra n ch e s , ,

f ro m the se p e n e tr a te all p a r t s o f t h e b o dy no ci rcul a to ry sy s te m ,

i s n e c e ssary to c ar r y nut rim e nt f ro m o ne pl a c e t o a n o th e r As .

in Hydr a no a nu s i s p re se nt the fa ce s b e ing e j e cte d thro ugh th e

, ,

m o uth .

E X C RE TOR Y SY S TE M Th e e xc re t o r y sy s t e m co mp r i se s a p a i r
.

o f l o n gitudin a l mu ch co il e d tub e s o ne o n e a ch Sid e o f th e b o dy ;

,
-
,

the se a r e c o nn e cte d n e a r th e a nte ri or e nd by a tr a nsve rse tub e ,

a nd o p e n t o th e e xt e r i o r by t w o s m a ll p o r e s o n t h e d orsa l s u r f a c e .

T h e l o ngitu di n al a nd t r a n sv e rse t r unk s giv e o ff num e r o u s fin e r

tub e s which r a mify thr o ugh all p ar ts o f th e b o dy u s u ally e nding ,

in a f la me ce ll T he fl a m e ce ll ( Fi g 7 8 ) i s l a r ge a nd holl o w
. .
,

with a bun ch o f fli cke ring cili a (c) e xt e nding into th e ce nt ra l ca v

ity (e ) Sinc e it co mmuni c a te s o nly with th e e xcre to r y tubul e s
.

it i s co n side r e d e xcr e t ory in functi o n tho ugh it m ay a l so ca r ry ,

o n r e spi ra t o r y a ctiviti e s .

M U SC U LAR SY S TE M T h e p o w e r o f ch a nging th e s h a p e o f its

.

b o dy whi ch m ay b e o b se rv e d whe n P lanar i a m o ve s fr o m pl a ce

,
‘

to pl a c e li e s p r incip a lly i n th r e e s e ts o f mu s cl e s a ci r cul ar l ay e r

, ,

j us t b e ne a th th e e ct o de r m e xte r n al a nd int e r n a l l a ye r s o f l o ngi

,

t u di nal muscl e fib e r s a nd a se t o f o bliqu e fib e r s lying in th e m e s o

,

de r m .

NE RV OUS SYS TE M . P la na r i a po s se s s e s a w e ll d e ve l ope d n e r v -

1 5 6 AN INT RODU C TI ON To Z OOLOGY

ous syste m co nsis ting of a bil ob e d m ass of tissue j us t b e n e a th the

,

e ye sp o ts call e d th e br a i n ( Fi g 7 7 cn) a nd two l a te ra l longi tu

,
.
,

di nal ne rve cor d s (ln) c o nn e ct e d

by t ra nsv e r s e n e rv e s From th e .

b r a in n e r ve s p as s to va ri o us p ar ts
,

o f th e a nt e r i or e nd o f the b o dy ,

imp ar ting to this r e gi on a highly '

s e n sitive n a tu re .

R E P R ODU C TI VE SY S TE M — . Re
p ro ducti o n i s by fi ssi on or by th e
se xu a l m e th o d E a ch in d ividu al.

p oss e ss e s b o th m al e a nd fe m a l e
o r g a ns i e IS, he r m aphr odi ti c
. . .

T h e m ale orga ns m a y b e l o c a t e d
e asily in F igu r e 7 7 ; th e y co n si s t

o f num e ro u s sph e r ic a l te s te s ( t ) .

co nn e cte d by sm all tub e s c all e d

FI G 7 8
c, c
Flam e CC “
ili a ; e ope ni ng i nto e xcre
Of P 10mm “d f
va sa

e e r e ns
d ef e r e nti a
fr o m ea
(
ch
vd .

S ide
th e
of
va s

th e
,
b o d Y j oi ns th e Ci n u s o r
,

P e ni s 3
t o ry t u b u l e ( Fr om La nk e s
mu s cul ar gan nte rs
.

a or WhI Ch e
t e r s Tre a ti se )
’
.

th e ge ni tal cloaca . A s e mi nal

ve s i cle li e sb a s e of th e p e ni s a l so a numb e r of uni c e ll ular
at th e ,

p ros tate gla nd s Sp e r m a t o zo a o r igin a t e in t h e t e st e s a nd p a ss

.
, ,

by w ay o f th e vas a def e r e nti a into th e s e min al v e sicl e whe re

, ,

the y r e m ain until n e e de d f o r fe r tiliza ti o n T h e fe male re pr o .

ducti ve orga ns c o mp r i se t w o ova r i e s (g) tw o l o ng ovi du cts (od ) ,

.

with m a ny yolk gl a nds (v) e nt e r ing the m a vagi na whi ch ,

Op e ns int o t h e g e ni t a l cl oaca a nd t h e u te rus whi ch i s a l so co n

, ,

ne ct e d with thi s c a vity Th e e ggs or igin a t e in th e o va r y p a ss

.
,

d own th e o viduct coll e cting yolk fro m th e yo lk gl a nds o n th e

,

way a nd fin ally r e a ch th e ut e r u s
,
He r e fe r tiliz a ti o n o c cur s and
.
,

cocoons a r e f or m e d e a ch c o nt a ining f r o m fo u r to m o r e th a n tw e nty

,

e ggs su rro und e d by se v e r a l hund r e d yo lk ce lls

,
T h e d e ve lop .

me nt of th e e gg i s ill us tr at e d and e xpl ain e d in F igur e 7 9 .

FI G 8 0 R e ge ne r a ti o n o f P la na r i a m a cu la ta
. . A, no r m a l w o r m ;
.

a nt e r i o r h a l f ; C , C , r e ge ne r a ti o n o f po s t e r i or h a l f ; D , crossp i e ce o f w o r m ; D ,
1 1

D , D , D , r e ge ne ra ti o n o f sa m e E o ld h e a d E , E , E , r e ge ne ra ti o n o f sa m e F ,
2 3 4
,
1 2 3

tio n o f he ad t e r io r he a d ( Fro m Mo rga n )

I
P ,
re ge ne r a ne w on po s e nd o f o ld . .
 SP ON GE S, FLAT WORM S , AN D R OU N D WOR M S 1 57

FI G 7 9 De ve l o pm e nt o f P la nar i a la cte a
. . .
,
I gg e ( )
0 yy o l
s u r ro u nd e
d b k (v)
2 , f o u r b l a s t o m e r e s f r o m s e gm e nt e d e gg ; 3 , l a t e r s t a ge ; m o r e b l a s t o

m e r e s ( bl) 4 , m u ch l a t e r s t a ge , d i ff e r e nti a ti o n o f bl a s to m e r e s i nt o e ct o
d e r m ( e p) , e nt o d e r m ( hy) , a pr o vi s io na l p h a rynx (ph) , w a nde r i ng
a nd

ce ll s ( to) ; 5 , ce ll u l a r d iff e r e nti a ti o n m o r e a d v a nce d ; e p e ct o d e r m ;

,

e nt, pr mi itive gu t ; hy e nto d e r m ; ph ph a rynx ;

, ,
6 e m b r yo ch a nge s
,

s h a pe t o a fl a tt e ne d o v o i d e nt p r i m itive gu t ; m
, , ,
m ou th ; ph p h a r ,

yu x .
( Fro m La nk e s t e r a f t e r Ha lle z ) .

R e ge n e r a ti o n . P l a n a r i a ns
ho w r e m ar k abl e p o we r s o f re
S

g e n e ra tio n If a n individu a l i s cu t in tw o ( Fi g 8 0 A) th e a n
. .
, ,

te ri or e nd will r e g e n e r a t e a ne w t ail ( B whi l e th e p os te ri o r ,

p ar t de ve l op s a ne w h e a d ( C A cr o sspi e c e ( D) will r e g e n e r
,

a t e b o th a h e a d a t th e a nt e r i o r e nd a nd a ne w t a il a t th e p os t e r i o r
,

e nd ( D T h e h e a d a l o n e Of a P l a n a r i a n will g r o w int o a n
'
—
D) 4
.

e nti r e a nim a l ( E — E) P i e ce s cu t fro m v a r i o u s p ar ts o f th e

3
.

b ody will al so r e ge n e ra te co mpl e t e ly N o diffi culty i s e xp e ri .

e nce d in g r a fting pi e ce s f r o m o ne a nim a l up o n a n o th e r a nd m a ny ,

cu rio us m on s t e r s h av e b e e n p r o duce d in thi s w ay .

1 5 8 AN INT R ODU CTION TO Z OOLOGY

FLAT WORMS IN GE N ER AL

Fl a t wo rms a re usu ally se p a ra te d into th re e cl a ss e s .

C l a ss 1 — Turb e ll a ri a F l a t w or m s with cili a t e d e pide rmi s

. .

a nd di g e s tiv e c a vity ; m o s tly no n p ara sitic E x a mpl e P la na ri a

- .
,
.

C l a ss 2 — Tr e m a to da Fl a t w or ms witho ut cili a te d e pi
. .

de r mi s ; di g e stive app a ra tus w e ll de v e l op e d ; e cto or e ndo

p arasitic E x a mpl e live r fluke of she e p
.
,
.

C l as s 3 C e s to d a
. F l a t w or ms with o ut cili a t e d e pid e r mi s
.

and dig e stiv e a pp ara tus ; e nd op ara sitic E x a mpl e t ap e w or m .

,
.

P la na ri a sh o ws th e p rincip a l f e a tu re s ch ar a ct e r i s tic o f fla t
w orm s ; but the re mu s t n e ce ss a rily be wide dive rsity in str uctu re
a m o ng th e m e mb e rs o f a phylum c o mp os e d o f b o th f r e e -livi n
g
a nd p aras itic f o r m s T h e fr e e li ving fla t w or m s such a s P la na ri a
.
-
, ,

p rob a bly a re m ore n e a rly like the a nce s t ors o f thi s phylum tha n
t h e p ar a sitic sp e ci e s sinc e th e l a tt e r h a v e und o ubt e dly b e c o m e
,

d e ge n e ra t e with r e sp e ct to c e r t ain s t r uctur e s and m ore sp e ci al ,

i z e d with r e sp e ct t o o th e r s b e c a us e o f th e i r m o difi e d h abits o f

,

life Fro m a study o f P la nar i a th e re fore w e can g ain s o m e ide a

.
, ,

o f wh a t kind o f a n a nim a l g a v e r i se t o t h e fla t w o r m s .

In th e fi rs t pl a c e de fini t e bi late r al symme try is e xhibit e d he re .

Fl a t wo r m s f b u s Sho w a n a dva nc e in thi s r e sp e ct o v e r th e m o re

simpl e ra di a l symm e t r y o f C oe l e nt e ra te s A s e cond p oint to be .

n o t e d is t he p re se nc e o f a dis tinct m e so de r m b e twe e n the e cto

de r m a nd e nto de r m Thi s m e so d e r m c on sists of mus cl e ce lls
.

a nd c o nn e ctiv e ti s s u e As in th e C oe l e nt e r a t e s ho we v e r th e re
.
, ,

is but o ne b ody c avity r e p re se nt e d by the dige stive sys t e m

, ,

tho ugh the g e nita l sacs m ay re p re s e nt a s e c ond c a vity kno wn ,

as the coelom which is w e ll d e ve l op e d in m or e c ompl e x

,

a nim a ls .

T h e d i ge sti ve a ppa r a tu s is no t a simpl e blind sa c a s in Hydr a , ,

but c o n si st s of s e ve r a l l a r ge b r a n che s e a ch with m a ny sm all e r ,

side p o uche s e nt e ring it th e whol e b e ing m o difi e d t o tr an sp or t

,

nutrim e nt to a ll p ar t s o f th e b o dy a ci r cul a t ory syst e m b e ing ,

l a cking An a nus h o we v e r i s a b se nt th e ing e s ti o n of f oo d a nd

.
, , ,
1 60 AN INT R ODU CTI ON To ZOOLOG Y

E xcr e tory orga ns mplica t e d r e pr oducti ve syste m m ake

a nd a co

the i r app e ara nce in thi s phylum T h e mu s cle fi be rs o f Hydr a .

a r e s impl e sp e ci a liz a ti o n s o f c e ll s th a t p e r f o r m o th e r fun cti o n s ;

but in P lana ri a c e ll s a re se t a side f or no o the r purp ose tha n to

,

give a hi gh de g r e e of c o nt r a ctility to the b o dy The se c e lls .

f o m di s tinct b a nds o f cir cul a l o ngi udin a l a nd o blique mu scl e s

r r t
, , .

Ta bl e V c o ntr a s ts th e s t r u ctu re s o f Hydr a a nd P la na r i a In .

this wa y a cl e a r id e a o f th e a dva nce in co mpl e xity e xhi bit e d

by th e l a tt e r m ay b e g a ine d .

3 . R OUN D W OR M S - ASCARI S

( As ca r i s lu mbr i coid e s Linn .

)
Ext e rnal F e atu r e s . und w or m p a r as itic i n th e
As ca r i s i s a ro

int e s tin e s of pig s ho r s e s a nd m a n T h e se x e s a r e s e p a r a

, ,
.

Th e f e m a l e b e ing t h e l a r g e r m e a s u r e s fr o m fi v e to e l e v e n in ch e s
, ,

in le ngth and ab o ut one f o u r th o f a n in ch in di a me te r T h e b ody .

i s light b r o wn in color ; it h a s a d o rs a l a nd a v e nt r a l whit e n ar r o w

s t r ip e r unning its e nti r e l e ngth a nd a b ro a d e r la te r a l li ne i s p r e s e nt

o n e ith e r S id e . T h e a nt e r i o r e nd p osse sse s a m ou th ope ni ng ,

s u rr o und e d by o ne d o r s a l a nd t w o v e nt r a l li ps N e a r th e po s .

t e ri o r e nd i s th e a nal ope ni ng f r o m whi ch in th e m al e e xt e nds

'

, , ,

p e n i al se tte o f u s e du r ing co pul a ti o n T h e m a l e ca

. n b e di s tin
g u i sh e d f r o m t h e f e m al e by t h e p re se nce of a b e nd in th e p os te ri or
p ar t o f th e b ody .

Inte r n al An atomy If a n anim a l i s cut Op e n a l o ng the d or s al

.

line ( Fi g . it will b e f o und to co nt ain a s t r aight a lim e nt a ry

c an al a nd c e r t ain o the r o rg a ns lying in a ce nt ra l ca vity th e
, , ,

coelom a ca vity m e t with no w f o r t h e fi rs t tim e

,
T h e a li me nta ry .

ca na l (2 ) is v e r y s impl e s inc e th e fo o d i s t a k e n f r o m m a t e r i a l
,

a l r e a dy di ge s t e d by th e h o s t wh ose int e s tin e th e w o r m inh a bit s .

It op e ns a t th e p o s t e r i o r e nd thro ugh th e anu s w hi ch i s no t p r e se nt ,

in m e mb e rs o f th e phyl a al re a dy discu sse d A mus cul a r pha rynx .

( I ) dr a w s th e fluids int o th e l o ng non mu s cular i nte s ti ne

-
 SP ON GE S, FLAT WOR M S ,
AN D R OU N D WORM S 1 61

t h r o ugh th e w all s o f whi ch th e

nutr im e nt i s a b so rb e d Jus t b e .

f or e th e a n a l Op e ning i s r e a ch e d ,

th e int e s tin e g r a du a lly b e c o m e s

Sm a ll e r ; thi s p or ti o n is kn o wn a s

the r e ctu m .

The e xcr e tor

y sys te m c o n si s t s of

two l o ngitudi n a l c a n al s ( Fi g 8 1 .
,

o ne in e a ch l a t e r a l lin e ; th e se

Op e n t o th e o ut s id e by a singl e

p o r e (8) situa t e d n e ar the a nt e ri o r

“
e nd in th e v e nt r a l b o dy w a ll .

A r ing o f ne rvou s ti ss u e s u r
r o und s t h e ph a r ynx a nd giv e s o ff

t w o l a r g e n e r ve co r d s o ne d orsa l
, ,

t h e o th e r v e nt r a l a nd a numb e r
,

of o th e r s m a ll e r st r a nds a nd
co nn e cti o n s .

T he m a le re produ cti ve or ga ns

a re a S ingl e co il e d th r e a d like -

te s ti s f r o m whi ch a vas d ef e r e ns
,

l e a d s to a wid e r tub e th e s e mi nal ,

ve s i cle ; thi s i s fo ll o w e d by th e
Sh o r t mus cul a r ej acu la tor y du ct
whi ch op e n s int o th e r e ctum .

In th e f e ma le li e s a Y sh ap e d -

re p ro d ucti ve syste m E a ch .

b ra nch o f th e Y co n si sts o f a
co il e d th r e a d li k e ova r
y ( Fi g 8 1
-
.
,

3 ) whi ch i s co ntinu o u s with a

l a r ge r ca na l th e u te r u s
, The
ut e ri o f th e two b r a n che s u ni te
into a sho r t mus cul a r tub e th e ,

va gi na whi ch op e n s to th e o u t
s id e th r o ugh t h e n i ta l a e r tu r e
g e
p
F e r ti li za ti on t a k e s pl ace in
t h e ut e r u s . T h e e gg i s th e n s u r
r o und e d by a S h e ll o f chitin a nd ,
1 62 AN INT RODU C TION TO ZOOLOGY

p ass e s o u t thr o ugh th e ge ni ta l p o r e . T he chitin o us e ggshe ll

p re ve nts the dige sti o n o f the e gg withi n th e inte s tine o f th e host .

FI G 8 2 T r a ns v e r se
. . se ctio n of As ca r i s lu mbr i coid e s . en, . cu ti cl e ; dl
.
,

d or sa l li ne ; de r . c .
,
e i i
p d e r m s ; e x v , e xcr e o ry u b e ; i nt , n e s
pl/mi . . t t . i t
ti ne ; la t l. .
,
li
la t e r a l ne ; m , m u s cu a r a ye r . ovy , o a r y ; l l
u t , u e r us ; . v . t
v v,
. . ve nt r a l li ne ( Fr o m P a r k e r a nd Ha s e a f e r Vo g a nd u ng )
. w ll t t Y .

Th e l ti o ns o f the va ri o u s org a ns t o one a n o the r a s w e ll a s

re a ,

th e s t r uctu r e o f t h e b o dy w a ll a nd th e ch ar a ct e r o f th e c oe l o m , ,

a r e s h o wn in F igu r e 8 2 whi ch i s a t r a n sv e r s e s e cti o n o f a f e m a l e

,

sp e cime n o f As ca r i s lu mbr i coid e s T h e b o dy o f th e wor m sho uld .

b e c o n sid e r e d a s c o n si s ting o f t w o tub e s o ne t h e int e s tin e (i nt ) , , ,

lying within the o the r th e b o dy wa ll ; whil e b e tw e e n the m i s a

,

c a vity th e coelom in whi ch li e the r e p ro ductive o rga n s (ovy a nd

, , .

T h e body wall i s mp o se d o f s e v e ra l l aye r s a n o ute r chitino us

co ,

cuticl e a thi n l a y e r o f e ct o d e r m (d e r e pthm ) ju s t b e n e a th it . .

,

a nd a thi ck s t r a tum o f l o ngitudin a l mu scl e fib e rs m e so de rmal

in o rigin lining th e c oe l o m Thi cke nings o f th e e cto d e r m form
, .

t h e d orsa l (d l) v e nt r a l (v v) a nd l a t e r a l (la t l) li n e s
.
,
In e a ch
. . .
 C HAP T E R X

HE EAR TH WORM AN D AN N ELID S IN G ENERAL

1 . TH E EAR TH WOR M

( Lu mbr i cu s te rre s tr i s L innae us)

THE e ar thworm ,
live s in th e gro und whe r e
Lu m br i cu s te r r e s tr i s , 1

th e soil i s no t to o d ry o r sa ndy c o ming to th e s u r f a ce o nl y a t ,

night o r a fte r a r a in T he burr o ws a re cylind ri ca l a nd p e n e t ra te

.

t o a d e pth o f f r o m s i x in ch e s to se v e ra l f e e t E vid e n ce s o f th e .

p r e se n ce o f e ar thwor m s a r e fo und in the littl e he ap s o f bl a ck

e a r th c all e d c as ting s whi ch s t re w th e g ro und b e ing e sp e
, , ,

ci a lly n o tic e a bl e e a r ly i n t h e m o r ni ng The se c a s ting s a re r e a lly .

th e f ae c e s c a s t o u t f ro m th e a lim e nt a r y c ana l o f th e w o r m D ar .

win ( 1 3 9) e s tim a t e d th a t m o r e th a n e ighte e n t o n s o f e a r thy c a s t

ings a r e ca rr i e d t o the s ur fa ce in a Singl e ye a r o n o ne a cr e o f
gr o und a nd in twe nty ye a r s a l a ye r thr e e in che s thick w o uld b e
,

tra n sfe rr e d f ro m th e s ub so il t o th e s ur fa c e Thi s co ntinu o us .

hon e ycombing o f the so il m a ke s th e l a nd m o r e p or o us a nd in su r e s ,

th e b e tt e r p e n e t r a ti o n o f m o i s tu re a nd th e co ntinu o u s w o r king ,

o v e r o f th e s u r f a c e l a y e rs o f e a r th a l so h e lp s t o m a k e t h e s o il m or e

f e r ti l e .

Exte rn al F e a tu r e s Lu m br i cu s te r r e str i s r e a ch e s a le ngth o f

.

fr o m si x in che s to a s mu ch a s a fo o t It i s c ap abl e o f p r o n o un ce d .

e xt e n s i o n s a nd co nt ra cti o n s s o t h a t t h e l e ngth o f a n in di vidu a l

‘

va ri e s a s doe s a l so i ts di a m e te r Altho ugh i ts b o dy i s cylind r o id

, .

it i s p ossibl e to r e cogniz e d or s al v e nt r al a nd l a te r al s u rfa ce s , ,

.

M ove m e n t t a ke s pl a ce in a de fini te di re cti o n th e a dva n cing ,

1
T hi s i s o ne of m a ny spe c i e s of e ar thw o rm s I n m any p a r t s o f thi s
.

co u n ry t the i
s pe c e s Allolobophor a ( He lodr ilu s ) longa o r o ne o f th e spe ci e s o f
D i pl o ca r d i a are m or e a b u nd a n t i n c u ltiva t e d so il ,

1 64
 T HE EART HWOR M AN D AN N ELI DS I N GEN ERAL 1 65

p or tio n b e ing kn o wn a s th e a nt e r i or th e bi nd e r p or tio n a s th e

,

p os t e ri o r e nd T h e p r incip a l a xis o f th e b o dy i s a nte r o p o s te ri o r

’
.
-
,

a nd s i nce t h e chi e f o r g a n s b o th e xt e r n a l a nd int e r n a l li e h a lf o n

,

o ne Side a nd h a lf o n t h e o th e r s id e o f a m e di a n pl a n e t h e w o r m is ,

s a id to be bil a te r a lly symme t rica l .

A n o tic e a bl e f e a tu r e o f t h e e a r thw or m is i ts divi si o n int o a l a r g e

numb e r o f simil a r r ings by g r oo ve s e xt e nding a r o und th e b o dy a t
sho r t inte r va l s Th e s e rings ar e t e r m e d
.

s e gm e nts , s om i te s , or m e ta me r e s , a nd th e
b o dy i s sa id to b e s e gm e nte d or to h a v e
a m e t a m e r ic s t r uctu re T h e e a r thw o r m
.

a nd m a ny o f its n e a r re l a tiv e s di ff e r in

thi s r e sp e ct f r o m th e un se gm e nt e d fla t
w o r m s a nd r o u n d w o r m s s u c h a s ,

P la nar i a a nd Asca r i s T h e s o mit e s a r e

.

no t e x a ctly a lik e . T h e a nt e r i or l o b e
e xt e ndi ng a b o v e a nd b e y o nd t h e m o uth ,

a nd b a ckw a r d o n t h e d o r s a l s u r f a ce i n

t e r se cti ng t h e fi r s t se gm e nt i s no t a t r u e
s o mit e ; it i s kn o wn a s t h e r os tom i u m t fL m
p FI G 84 T w . . . o se ae o u

I t h a s b e e n fo u nd th a t no t o nly e xt e r n a l M hi ghl y m gni a ts , a

fi d ( F m P k
st ru ctu re s but a l so int e r n al o r g a n s b e a r
, ,
r e ' ro ar e

nd H w ll ) a as e
a co n s t a nt r e l a t i o n to t h e s e gm e nt s ;
.

f o r this r e a so n th e so mit e s h a v e b e e n numb e r e d b e ginning with ,

th e o ne ju s t b a ck o f th e p r o s t o mium In m a tu r e wo r m s t h e
.

six o r s e ve n s o mit e s XXXI o r XXXII t o XXX VII a r e swo ll e n

, ,

o n th e i r d o rs a l a nd l a t e r a l s u r f a ce s p r o du cing a sa ddl e Sh a p e d -
,

e nl a r g e m e nt kn o wn a s th e cli te llu m o f u se du r ing r e p ro du cti o n .

,

If a n app a r e ntly s mo o th e ar thw o r m i s d ra wn thr o ugh th e

finge r s fro m i ts a nte r ior e nd t o wa r d th e t a il it fe e l s ro ugh to th e
,

to uch This i s c a use d by small f sh ap e d chitin o u s b ri s tl e s call e d

.
-
,

s e lm ( F i g. fo u r p ai rs of which e xt e nd o utwa r d fro m e pide r m a l

s acs in e ve ry so mite e x ce pt th e fi rs t a nd l a s t E a ch p ai r m ay b e .

m o ve d by re t r acto r a nd p ro t r acto r mu scl e s T h e a rra nge m e nt .

of th e s e t ae is s h o wn ha lf o f th e se ctio n in Figure
1 66 AN INTRODU CTION TO Z OOLOGY

85, se t
. Ne w se tae
p ro duce d by any o ne o f a numb e r of
m ay b e
c e lls lying a t th e b ott om o f th e e pide r m al sacs On m a tu r e
w o r ms th e se t ae o n s o mite XX VI ar e e nl arge d a nd m
.

o difi e d a s

s e xu al s e t ae .

s u ba rea

FI G 8 5 T r a ns v e r se
. . tio n th ro u gh th e m i dd l e r e gio n o f th e b o dy o f Lu m
se c

br i cu s . ci r c m u s .ci r cu l a r m u s cl e fi b e r s ; cce l
.
,
coe l o m ; d or s v d o r sa l .
,
. .
,

ve sse l ; e pid e pi d e r m i s ; e xt ne ph ne ph r i d io por e ; he p chl o ra goge n

.
, . .
,
.
,

ce ll s ; long m u s . l o ngi ud i na l m u s cl e s ; ne ph ne p h r i d i u m ; ne phr osl

.
,
t
,
.
,

ne ph r os t o m e n co ne r ve co r d ; s e t s e t ae ; s u b n ve s s s u b ne u ra l
. .
,
.
,
. . .
,
-

v e s se l ; typh typ hl o so l e ; ve nt v v e nt ra l ve sse l ( Fro m P a r k e r a nd

.
, . .
, .

Hasw e ll a f t e r M a r sh all a nd H u r s t ) .

Th e ut e r cove ring of th e b o dy i s a thi n t r an sp are nt m e mb rane

o ,

th e cu ti cle ( Fi g 8 5 which i s se cr e t e d by the ce ll s lying just

.
,

b e n e ath it Th e cuticl e p r o te ct s th e b o dy wall fro m physical

.
1 68 AN INT RODU C TION TO Z OOLOGY

coe l .
) c o mmu ni c a t e s with xt e r i or by m e a n s o f dors a l por e s
th e e .

One of the se i s l o ca te d in th e mid dorsa l lin e a t th e a nte ri o r

-

e dge o f e ach so mi t e f r o m VIII o r IX t o th e p o s t e r i o r e nd o f th e

b o dy .

G e n e r al In te r n al An a to my If a sp e cim e n is cut o p e n fro m

.

th e a nt e r i o r to th e p o s t e r i o r e nd by a n in ci si o n p a ssing th ro ugh
t h e b o dy w a ll a t r ifl e to o ne sid e o f th e mid d o r s a l lin e a g e n e ra l -
,

vi e w o f th e int e r n a l s t r u ctu r e s m a y b e o bt a in e d ( Fi g As in .

Ascar i s th e b o dy i s e sse nti a lly a d ou ble tu be ( Fi g 8

, th e body .

wa ll co n s tituting th e o ut e r t h e s t ra ight a li m e nta ry ca na l th e

, ,

inn e r ; b e tw e e n the tw o i s a c a vity th e coelom T h e e xte rn a l , .

s e gm e nt a tio n cor re sp o nds to a n inte r n a l di vi si on o f the c oe l omic

c a vity into comp ar tm e nt s by m e a n s o f p a r titi o n s call e d s e pta , ,

whi ch li e b e n e a th th e g ro o ve s Th e se se pt a ar e ab se nt in Asca ri s
. .

T h e alim e nt a r y ca n a l p a sse s th ro ugh th e ce nt e r o f th e b o dy a nd i s ,

s u sp e nd e d i n th e coe l o m by th e p ar titi o ns Se pt a a re a b se nt .

be tw e e n so mit e s I a nd II a nd in compl e t e b e tw e e n somit e s III

,

a nd I V a nd X VII a nd X VIII
,
T h e w a ll s o f th e c oe l o m a r e lin e d
.

with a n e pithe lium t e r m e d th e pe r i tone u m Th e c oe l omic cavity

,
.

i s fill e d with a col o rl e s s fluid whi ch fl o w s fr o m o ne co mp a r tm e nt

t o a n o th e r wh e n th e b o dy o f th e w o r m co nt ra ct s In so mite s IX .

to X VI a r e th e r e pr odu cti ve or ga ns ( Fi g 93 ) r unning al o ng t h e .

upp e r sur face o f the a lim e nta ry can al i s th e d orsal blood ve ss e l

( g 5
F i 8. d or s ,
a n d ju
. s t b e n e a th it li e t h e ve ntr a l blood ve s s e ls

(ve nt . v.
) a ndne r ve cor d (n .

B o dy Wal —
l t t a of the b o dy w all are a s foll o ws
T he s ra
.

( g
F i ( )
I a n o ut e r n o nc e llul a r co v e r ing t h e c u ti cle ( );
cu t ,

( )
2 a n e pi d e r m i s (p )
e i d co mp o s e d o f t w o c e llul a r l a y e rs a n o ut e r ,

s t ra tum co n si s ting o f gl a nd int e r s titi a l ,

a nd se n s e c e ll s a nd a n
, ,

inn e r stra tum o f ve ry sm all c e ll s ; ( 3 ) a l ay e r o f ci r cu lar mu scle

b e rs (ci rc m u s ) r unning a ro und th e b o dy e ach fib e r b e ing l ong
fi . .
, ,

p ointe d a t b o th e nds a nd l ongitudin ally st r i a t e d ; (4) a thick

,

l aye r of mu scl e fib e rs r unning l e ngthwi se o f th e b o dy and app e a r

i ng in cr oss s e cti o n to b e a rr a ng e d in fe a th e rlik e g r o up s (long .

mus ) ; 5) th e i nn e r m o s t l aye r the c oe l omi c e pithe lium o r pe rit o

,
1 70 AN INTR ODU CT I ON TO Z OOLOGY

XII T h e fi r s t p ai r ar e p o uche s pu sh e d ou t fro m th e alime nt a ry

.

ca n al a nd Op e ning di re ctly into th e oe soph agus T h e o the r tw o .

p ai rs are swe lling s of th e oe sophage a l wa ll ; the y h ave a numb e r o f

sm all cavitie s whi ch op e n di r e ctly th r o ugh th e e pithe lium int o
th e oe soph a gu s in so mit e X I V C a rb o n a t e o f lim e i s p r o du ce d
.

by the se gl and s a nd p o u r e d into th e alime nta r y ca n al whe re it

, ,

p ro b ably n e ut r aliz e s a cid foo d s

N utr iti on .F OO D — T h e fo o d o f t h e e ar thw o r m c o n si s t s
.

p r in cip ally o f pie ce s o f l e av e s a nd o th e r v e ge ta tio n p a r ticl e s o f ,

a nim a l m a tt er a nd s o il ,
This m a te ri al i s ga th e r e d a t night At
. .

this tim e th e wo r m s a re a ctive ; the y cr a wl o u t int o th e ai r and , ,

with the i r t ail s holding fa s t to th e top s o f th e i r bu r ro w s e xpl o r e ,

th e n e i ghb o r h oo d .

I N GE STI ON — F oo d p ar ti cl e s a r e d ra wn int o th e bucc al c a vity

.

by s ucti o n p r o duce d whe n th e pha ryng e a l cavity i s e nl a rge d .

This i s a cco mpli sh e d by th e co nt r a cti o n o f the muscl e s whi ch

e xt e nd f ro m t h e ph a r ynx t o t h e b o dy w a ll .

D I GE S TI ON In th e ph a rynx th e fo o d r e c e iv e s a s e c r e tio n f ro m
.
,

th e ph a r yng e a l gl and s ; it th e n p a sse s thro ugh th e oe s oph agu s

t o th e cr0p wh e re it i s s t o r e d t e mp o r a r i ly
,
In th e m e a ntim e .

t h e se cr e ti o n f r o m th e cal cif e ro u s gl a nd s i n th e oe so ph ag e a l w all s

i s a dd e d n e ut r alizing t h e acid s
,
T h e g izz a r d i s a g rinding o r
.

g a n ; in it t h e f oo d i s b ro k e n up int o minut e f r a gm e nt s by b e ing

sque e ze d a nd r o ll e d ab o ut Solid p a r ti cl e s s u ch a s r o ugh p e bble s
.
, ,

whi ch are fr e que ntly swal lo we d p r o b ably a i d in this grinding ,

p ro ce ss T he foo d the n p a sse s o n t o the int e s tin e whe re m o st o f

.
,

th e dig e s ti o n a nd a b s o rpti o n t ak e s pl a ce .

Dig e s ti o n in th e e a r thwo r m i s v e r y sim il a r t o th a t o f high e r

a nim a l s . T h e dig e stiv e fluid s act up o n pr ote id s ca r bohydr a te s , ,

a nd f a ts ; in the m a re sp e ci a l ch e mical co mp o unds call e d f e r ,

me nts o r e nzyme s whi ch b r e a k up co mpl e x m o l e cul e s with o ut

,

the mse lve s b e co mi ng cha ng e d ch e mi cally T h e th re e m o s t

‘

imp o r ta nt e nzym e s a re ( 1 ) tryps i n whi ch di ssolve s p ro t e id , ,

( )2 di a s tase whi,
ch b r e a k s up m o l e c ul e s o f c a r b o hyd r a t e s a n d ,

(3 ) s te a p s i n whi c h
, a c t s up o n f a t s T h e se th . r e e e nzym e s a re
 T HE EART H WOR M I
AN D AN N EL D S I N GEN E R AL 1 7 1

p rob ably p re se nt in th e di g e s tiv e fl uids of th e e a r th wo r m T he .

p ro te ids a re cha ng e d into p e pto ne s the ca rb o hyd ra t e s into a

,

sug a r c o mp o und a nd th e fa t s a r e divid e d into glyce r in a nd f a tty

,

a cid s.

AB SORP TI ON T h e foo d is no w r e a dy f or a b so rpti o n

. Thi s i s .

a cco mpli s h e d th ro ugh t h e w a ll o f th e int e s tin e by a p ro ce ss

kn o wn a s os mosi s a s s i s te d by a n a m e b o id a ctivity o f s o m e o f
,

th e e pith e li a l c e ll s Osm osi s is th e p a ssa ge o f a liquid th ro ugh

.

a m e mb r a n e .

C I R C U LA T I O N Up o n re a ching the bl oo d th e ab so rb e d foo d i s

.
,

c a rri e d to v a ri o us p a r ts o f th e b o dy by Ci rcul a tio n th e de t ail s o f ,

whi ch a re d e s crib e d in a n o th e r pl a ce (pp 1 7 4 Ab s o rb e d

.

fo o d a l so m a k e s i ts w a y int o th e coe l o mi c ca vity a nd i s ca rrie d

di re ctly to tho se ti ss ue s b a the d by the cce lom i c f lu i d In o ne .

c e ll e d a nim al s a nd in s u ch M e ta z o o ns a s Hyd r a P la na r i a a nd
, , ,

As ca r i s no ci r cul a t o r y sy s t e m i s n e ce ss ar y s in ce th e fo o d e ith e r i s
,

dige s te d withi n the ce ll s o r co m e s into dir e ct co nt a ct with the m ;

but in l a rg e co mpl e x a ni m als a Sp e ci al sys te m o f o rg a ns must be
,

p ro vide d to e n abl e the p ro p e r dis t r ibuti o n o f nut rime nt .

AS SI M I LATI ON a s in t h e typ e s a l r e a dy d e s c rib e d i s th e a dditi o n

, ,

o f ne w p a r ti cl e s a m o ng t h e p re e xi s ting p a r ticl e s o f p r o t o pl a s m .

Va s cular Sys te m — T h e bl oo d o f th e e a r thwo r m i s co nt a in e d

.

in a co mpli ca te d sy s te m o f tub e s which r a mify to all p a r ts o f the

b o dy A numb e r o f th e se tub e s a r e l a rg e a nd ce nt ra lly l o ca t e d ;
.

th e se give o ff b r a n che s which like wi se b r a n ch fin ally e nding in ,

e x ce e di ngly thin tubul e s th e ca pilla r i e s T h e fun cti o ns o f thi s

.
,

s y s t e m o f tub e s a r e t o ca r r y n o u r i s hm e nt f r o m th e a lim e nt a ry

ca n a l t o a ll p a r t s o f th e b o dy t o t r a n sp o r t w a st e p ro du ct s a nd t o
, ,

co nve y th e bl oo d to a p o int n e a r th e s u rfa ce o f th e b o dy whe re

o xyg e n m a y b e o bt a in e d a nd s uppli e d t o t h e ti ss u e s .

B L OOD . T h e bl o o d o f th e e a r thw or m co n si s t s o f a plas ma in

whi ch a r e s usp e nde d a g r e a t numb e r o f col o r l e ss ce ll s call e d ,

cor pu s cle s . I t s re d col o r i s d ue t o a pigm e nt t e r m e d hee moglobi n ,

whi ch i s di ssolve d in th e pl a sm a In ve r te b ra te s th e h a m oglobi n

.

i s l o ca t e d in th e bl oo d co rpu scl e s .
1 7 2 AN IN TR ODU C T ION TO ZOOLOGY

B L OOD VE S SE L S . F o ll o wing
cus to m whi ch i s so fi r mly
— a ,

e s t a bli sh e d a s t o m a k e i ts a b a nd o nm e nt in a dvi sa bl e w e sh a ll call

t h e bl o o d tub e s by th e in a pp ro p r i a t e n a m e
“ ve sse l s ”
Lo ngi
,

88 . D i agr a m s ho wing th e a rr a nge m e nt o f th e b loo d ve s se l s i n th e e a r th

s

w o r m A l o ngit u di na l vi e w o f th e v e s se l s i n so m it e s VII I I X a nd
.
, , ,

X ; B t ra ns ve rse se cti on o f sa m e r e gio n ; C l o ngit u d i na l vi e w o f th e

, ,

ve sse l s i n th e i nt e s ti na l r e gio n ; D t r a ns ve r s e se cti o n th ro u gh t h e

,

i nt e s ti na l re gio n ; of i a ff e r e nt i nt e s ti na l v e sse l ; c v pa r i e t a l v e sse l ;

. .
, . .
,

ef. i e ff e r e nt i nt e s ti na l v e s s e l ; ht h e a r t ; i i nt e s ti ne ; i t i nt e s ti no
.
,
.
, ,
.
,

t e gu m e nt a ry ; u l l a t e r a l ne u r a l v e sse l ; a s oe so ph a gu s ; s s e pta s b
.
,
-
,
. ,
.
,

ve nt ra l v e ss e l ; s n s ub ne u r a l v e sse l ; s p d o r sa l v e sse l ; ty typhlo so l a r

.
,
- .
,
.
,

v e sse l ( Fro m B o ur ne a f t e r B e nh a m )
. .
1 74 AN IN T RODU C TI ON TO Z OOLOGY

n e phridia and b o dy wall (D) The dor s al t r unk al so co mmuni .

ca te s wi th th e inte s tine by m e a n s of b r a n ch e s T wo p ai rs of .

v e sse l s p a ss fr o m it t o th e w all s o f th e int e s tine in e ve ry se gm e nt

(fe . i in. C ) T h e bl o o d i s co ll e ct e d fro m th e int e s tin e by two p a i r s
.

o f v e s s e l s whi ch e nt e r a l o ngitudi n a l typhlos ola r tub e ( t in C )

y . .

T h e l a tt e r i s c o nn e ct e d with t h e d or sa l t r unk by th r e e o r fou r

Sh or t tub e s in e ach s o mit e Th e v e nt r a l t r unk in th e int e stin al
.

r e gi o n giv e s o ff in e v e r y s e gm e nt a p a i r o f v e ss e l s e a ch o f which

divide s se nding a b r a nch to the n e phridium and one to th e b ody

,

w all (s b in D) . .

STR U C T UR E OF B L OOD VE S SE L S — Th e d or s a l t r u nk a nd h e ar t s
.

d e t e r m in e th e di r e ctio n o f th e bl oo d flow sin ce the y furnish th e ,

p owe r by m e a n s o f the i r mu scul a r wall s T h e wall o f th e do rsal .

t r unk is co mp o se d o f fo u r l a ye r s : ( 1 ) a n inn e r e pithe lium of thi n

c e ll s ( 2) a c o nne ctiv e ti ss ue l a ye r ; (3 ) a we ll de ve l op e d st ra tum
,
-

o f ci r cul a r mu scl e fib e r s a nd ( 4 ) a n o ut e r c o v e r ing o f chlo r ogoge n

,

c e ll s T h e w all s o f th e h e a r t s a r e Simi l ar in s tr uctur e P a i r s o f

. .

fo r wardly di re cte d va lve s a r e Situ a t e d in th e do rsa l t r unk j ust

b e hi nd th e Op e nings o f the p a ri e tal v e sse l s Th e va lve s do no t .

p re ve nt th e flo w o f th e bl oo d in a n a nt e ri o r dir e cti on but th e ,

do r s al t r unk wh e n co n st r i ct e d i s co mpl e te ly cl ose d by the m

, , ,

m a king th e b ackw a r d flo w imp o ssibl e P ai rs o f va lve s are al so .

p re se nt in th e d o r sal t r unk jus t in fro nt o f th e op e nings o f th e

he a r ts Othe r v alve s o ccu r in th e v e sse l s di re ctly co nne cte d with
.

th e d o r s a l t r unk .

T h e v e nt r a l t r unk d o e s no t p o sse s s a l a y e r o f ci r cul a r mu scl e

fi b e r s a nd th e r e fo r e h a s no t th e p o we r t o c o nt r act
, , ,
It i s .

s t r e ngth e n e d by a thick l a y e r o f fib r o u s c o nn e ctiv e ti ss u e The .

s u b n e u r a l a nd l a t e r a l n e u ra l t r unk s a s w e ll a s a ll t h e s m alle r
- -
,

br a nch e s h a v e s imply a n inn e r e pithe li um a nd a n o ut e r co nn e c

,
~

tive ti ss ue l a ye r
C I R C U LATI ON T h e flo w o f bl o o d in th e bl o o d v e sse l s i s no t
.

s e gm e nt al but syste mi c B l o o d i s fo r ce d fo rwa r d by w ave like

,
.
-

co nt r a cti o n s o f th e d o r s a l t r u nk b e ginning a t t h e p o s t e ri o r e nd
,

and t r a v e ling quickly a nt e r i o r ly The se co nt ra ctio n s a re sai d

.
 T HE E ART HWORM AN D AN N EL DS I N I GEN ER AL 1
75

to b e p h
e r i s talti c, a nd
a v e b e e n lik e n e d t o th e a cti o n o f t h e fing e r s

in th e ope r a ti o n o f mi lking T h e v alve s in th e w alls o f t h e dor sal

l

t runk p re v e nt th e r e tu r n of bl oo d fro m th e a nte r ior e nd In .

so mit e s VI I t o XI t h e bl oo d p a s se s f ro m t h e d or s a l t r u nk int o

th e h e a r t s a nd i s f or c e d by th e m b o th f o r w a r d a nd b a ckw a r d
,

in th e ve nt ral t runk Th e v alve s in th e he ar t al so p re ve nt th e

.

b a ckw a r d flo w F r o m the v e ntral tr unk th e bl oo d p asse s to th e

.

b ody wall a nd n e phridi a Bl o o d i s r e tu r n e d fr o m the b o dy w all

.

t o th e l a t e r a l n e u r a l t r unks
- T h e flo w in th e su b n e u r a l t r unk is
.
-

t owar d the p oste rior e nd the n upw a r d thr o ugh th e p a ri e t al

,

ve sse l s into th e dorsal t r unk T he ant e ri or r e gio n r e ce ive s bl oo d

.

f ro m th e do r sal a nd v e nt r al t r unks T h e bl o o d whi ch i s c a rr ie d .

to th e b o dy w a ll a nd int e gum e nt r e c e iv e s o xyg e n thr o ugh th e

cuticl e a nd i s the n r e tur ne d to th e dorsal t r unk by w ay o f the
,

s u b n e u ra l t r unk a nd th e int e s tin al c o nn e c tiv e s

- B e c a u se o f i ts .

p ro ximity to the sub n e u ral t r unk th e n e rv o us sy ste m r e ce ive s a

-
,

co ntinu o u s s upply o f th e f re she s t bl oo d ( 1 4 3 ,

R e s pi ra ti o n — T h e e a r thw o r m p o sse s se s no r e spi ra t or y sys

.

te m but o bt a in s o xyg e n a nd g e t s r i d o f car b o n di o xi d e th r o ugh

,

th e m o i s t o ut e r m e mb ra n e M a ny ca pill a ri e s li e jus t b e n e a th th e
.

cuti cl e m aki ng the e x ch ang e of g ase s e a sy T he o xyge n is co m

, .

b i ne d with the h a m oglobi n .

Excr e tory Or ga n s M o s t o f th e e xc r e t ory m a tt e r i s c arri e d

.

o ut sid e o f th e b o dy by a numb e r o f co il e d tub e s t e r m e d ne hr id i a

p ,

( Fi g 8 5 m ph ) a p ai r o f whi ch a r e p r e se nt in e v e r y so m it e e xce pt
.
, ,

th e fi rs t thr e e a nd t h e l a s t A n e ph r idium occupi e s p a r t o f tw o

.

s u cce s siv e s o mit e s ; in o ne i s a cili a t e d funn e l th e ne phr os to me ,

( Fi g 8 5
.
, whi ch i s conne ct e d by a thi n ci li ate d tu be
with the m aj o r p o r ti o n o f th e st ructu re in th e s o mit e p os te ri o r
to it .
Th re e l oop s m ak e up th e co il e d p o r ti o n o f th e n e phridi um .

T he thi n tub e m e nti o n e d a b o v e i s a singl e ro w o f h oll o w cyli n

d ri cal ce ll s pl a c e d e nd t o e nd ; it e xt e nd s th ro ugh o ne l o op a nd a
h alf a nd co nn e cts wi th a l a rge r tub e b ro wn in col o r a nd cilia t e d
,

th ro ugho ut i t s ce nt r al ca vity This p or ti o n kn o wn a s th e m iddle

.
,

tube Op e n s int o a thir d tu be whi ch i s wid e r but wi th o ut cili a

, , ,
.
1 7 6 AN INT R ODU CTION To ZOOLOGY

Le a di ng fro m
thi r d tub e to th e e xte rn al ap e r ture i s a l arge
th e
mu s cu lar d u ct T h e s t r u ctu r e o f t he n e ph r os t o m e a nd o f th e
.

thi n tub e jus t b e hind it i s o f p a r ti cul a r int e r e st T h e n e phr o

’

s to m e ( Fi g 8 9) c o nsi s t s o f a l ar g e c r e s c e ntic c e nt ra l c e ll
. su r

r o und e d by a l a y e r o f

cili a t e d m argin al ce ll s
the se j oin a num
be r of g roov e d c e lls (cf )
whi ch le a d to th e thin
cili a t e d tub e T h e ce n .

t ral cavity o f this cil

iate d tube i s no t
s u rro und e d by ce ll s ,

th a t is inte rce llul ar , ,

but i s int r ace llul ar p ass ,

ing di re ctly thr o ugh the

ce ll s T h e tub e thu s .

re se mbl e s a d r ainpip e ,

th e se cti o n s o f which
a r e re p r e s e nt e d by s i n

g le ce ll s pl a ce d e nd t o

e nd .

FI G 8 9
. . N e p h ro s to m e of th e e ar thwor m . c c
. .
,
E X C RE TI ON . A ne
ce t r a l ce ll ; cf
n .
, groo v e d ce ll s ; p
c .
,
c ili a t e d
ph r i di u m fun cti on s in
m ar gi na l ce ll s ( Fro m Da hl gre n a nd
th e fo ll owing m ann e r
.

K e pne r ) ili a on the n e ph The

'

r o st o m e a nd in th e thi n tub e a nd middl e tub e c r e a t e a cu r r e nt

th rough the muscul a r duct whi ch l e a ds t o th e e xt e rio r Solid .

w ast e p a r ti cl e s which m ay b e floa ting ab out in th e c oe l o mic fluid

a r e d r a wn int o t h e n e ph r idium by thi s cu r r e nt a nd p a s s t o th e ,

o ut sid e Wast e m a tte r in s oluti o n i s t ak e n fro m th e bl ood by

“
.

”
th e gl a nd s o f th e wide tub e and store d in th e large s ac like -

mus cul ar duct until e xc re te d .

N e r vo u s Sys te m T h e n e r vo u s sy st e m di ffe r s fro m th a t o f

.

P la na ri a a nd As ca r i s in b e ing m ore co nc e nt r a t e d At th e an .
1 7 8 AN IN TRODU CTION TO Z OOLOGY

p p e ri
h e r a l ne rvou s s s te m ( F i g 9 0 , 9 a nd
y . T h e s up r a -
ph a ryn
ge a l g a ngli a s upply th e p ro s t o mium with tw o l a rg e n e rve s whi ch
giv e o ff m a ny b ra nche s (8 ) th e y a l so se nd n e rve s into s omit e s
II a nd III One n e r ve e xt e nds o u t f ro m e a ch ci r cum pha ryng e a l
.
-

co nn e ctiv e . In e a ch so mi te fr o m I V t o th e p os t e ri o r e nd o f the
b o dy th re e p a i r s o f n e rv e s a ri se tw o p airs fr o m th e g a nglio ni c
, ,

m ass a nd one p ai r f ro m the side s o f the n e rv e co r d jus t b e hind

t he se ptum whi ch se p a r a t e s th e s o mit e f r o m th e o ne p re ce di ng .

E a ch e nl a r g e m e nt o f th e v e nt r a l n e rv e c o r d r e a lly co n si s t s o f
t wo g angli a whi ch a r e cl o se ly fu s e d t o g e th e r In t r ansv e r se
.

s e cti o n th e se fu se d g a ngli a a r e se e n t o b e s u rr o und e d by a n o ut e r

thi n l aye r o f e pithe lium th e p e rito n e um a nd a n inn e r muscul ar

, ,

s h e a th co nt a ini ng bl o o d v e ss e l s a nd co nn e ctiv e ti s s u e a s w e ll a s

muscl e fib e r s N e a r the d o rsa l s u r fa ce a re th re e l a rg e a r e a s

.
,

e a ch s u r r o und e d by a thi ck d o ubl e Sh e a th a nd co nt a ining a bundl e

o f n e r v e fib e r s
“
The se a re ca lle d ne u ro ch o rd s o r gi a nt fi be r s
. .
”

La rg e p e a r sh a p e d ne rve ce lls a re vi s ibl e n e a r th e p e r iph e r y in th e

-

l a te ra l a nd v e ntr a l p a r t s o f th e g anglio n .

Th e n e r v e s o f th e p e r iph e r a l n e r vo u s sys t e m a r e e ith e r e ffe r e nt

o r a ff e r e nt Efi e re nt n e rv e fib e r s a r e e xt e n si o n s f r o m ce ll s in th e
‘

ga ngli a o f th e ce ntra l n e rvo u s sys t e m Th e y p ass o u t t o th e

.

muscl e s o r o th e r o rg a n s a nd sin ce impul se s se nt a l ong the m g ive

, ,

ri s e t o m o v e m e nt s th e ce ll s o f w hi ch t h e v a r e a p a r t a r e sa id to b e
,

m otor ne rve ce lls T h e a fie r e nt fib e r s o rigin a t e f ro m n e r v e ce lls

.

in th e e pide r mi s whi ch a re se ns ory in functi on and e xt e nd into ,

th e v e nt r a l n e r v e c or d .

T h e fu ncti ons o f n e r vo u s ti ss u e a re pe r ce pti on condu cti on a nd

, ,

s ti mu la ti on Th e se a re u su a lly p e r fo rm e d by n e rv e c e ll s ,

c all e d n e ur o n s Th e ne u r on the or y s upp ose s th a t the r e i s no

.

ne r v e fib e r inde p e nd e nt o f n e r v e c e ll a nd th a t th e c e ll with
a ll its p ro l o ng a ti o n s i s a unit o r a n e u r on ; th a t th e s e unit s a r e

no t uni t e d t o o ne a n o th e r a n a t o mi ca lly but a ct t og e th e r phy si o

,

l ogica lly by co nta ct ; tha t th e e nti r e n e r vo us sys te m c onsi sts o f

sup e rimp ose d n e uro n s ; (13 6 p ,
.

T h e r ef le x c arri e d ou t e i th e r co n sci ousl o r unconsci o u sly i s

y
 THE EART HWORM AN D AN NELI DS I N GEN ER AL 1 79

co n side re d th e physio l ogi ca l unit o f n e r vo u s a ctivity T h e a p .

p a r a tu s r e qu ir e d f or a simpl e r e fl e x in th e b o dy o f a n e a r thwo rm
i s r e p r e s e nt e d in F igu r e 9 1 A p r im a r y se n so r y n e u r o n
.

lying at th e s u r fa c e o f th e b o dy s e nds a fib e r (sf ) into th e v e nt r a l

,
.

n e r ve cor d whe r e it b r a nche s ou t ; the se b r a n che s a r e in physi o

l o gi ca l co ntinuity with b r a nch e s fr o m a p rim a r y m o to r ne u ro n
( )
i nc .ly i ng in t h e g a ngli o n o f t h e v e nt r a l n e r v e co r d T h e s e co nd .

V
9

FI G 9 1
. . T r a ns v e r se
ti o n o f th e v e nt r a l ne r ve ch a i n a nd s u r ro u nd i ng
se c

s t r u c t u r e s o f a n e a r thw o r m cm . ci r cu l a r m u scl e s ; e p
. e pi d e r m i s ; .
, ,

[ m l o ngit u d i na l m u scl e s ; mc m o t o r ce ll b o d y ; mf
.
, .
, m o t o r ne r-
. ve
,

fi be r ; s e se nso ry ce ll b o dy ; sf s e nso ry ne r v e fi be r ; vg v e nt r a l ga n
.
, .
, .
,

gli o n ( Fr o m P a rk e r i n P op S ci M onthly m o d ifi e d a f t e r R e tz i us )
. . .
,
.

n e u ro n (mc ) se nd s fib e rs (mf ) int o a r e a cting o rg a n whi ch in thi s

. .
,

ca s e i s a mu s cl e The se fib e r s e xt e nding t o th e r e a cting o rga n a re

.

ca ll e d m o t o r fib e r s tho se l e a ding t o the v e ntr a l n e r ve co r d

a r e t e r m e d se n s o ry fib e r s T h e fi r s t n e u r o n o r r e ce ptor r e, ,

ce i v e s t he s timulu s a nd p r o duc e s t h e n e r v e impul s e t h e s e c o nd

;
ne u r o n t h e adj u s tor r e ce ive s di r e ct s a nd m o di fi e s t h e impul s e ;
, , , ,

a nd t h e mu scl e o r o the r o rg a n stimul a t e d t o a cti vity i s the

e
j e ctor W.ithi n t h e v e nt r a l n e r v e co r d a r e a s soci a ti on ne u r ons

who se fib e rs se rve t o co nne ct s t r uctu re s within o ne g a ngli o n o r t wo

s u cce e ding g a ngli a The se Sho r t n e ur o n s o ve rl ap o ne a n o the r
.
,

a nd a r e d o ubtl e ss r e sp o n sibl e f o r th e mu s cul a r w a v e s whi ch p ass

fro m th e a nte ri o r to the p o st e r io r e nd o f th e w o r m during l o co

m o tio n T he th re e gi a nt fi be rs which li e in th e dor sal p art o f
.
,
1 80 AN I N T R ODU C T ON T o I Z OOLOGY

th e v e ntr al n e r ve co r d th ro ugho ut almos t i ts e ntir e l e ngth are ,

co nne ct e d by m e a n s o f fib ri l s with n e rv e ce ll s in th e g angli a

a nd p r o b a bly di s t r ibut e th e impul se th a t c a u s e s a w o rm t o co n

t r a ct i t s e nti r e b o dy wh e n s t r o ngly
pe r .

stim ul a te d
S e n s e Or ga n s — T h e se n sitive .

n e ss o f Lu mbr i cu s to ligh t a nd
o th e r s timuli i s d ue t o th e p r e se nc e
an 0 .

of a g r e a t numb e r o f e pide rm al
se nse or g a n s The s e a re groups o f .

s e n se ce ll s (Fi g 9 2 sn c ) co nne ct e d .
,
. .

with the ce nt ra l n e r vo us syste m by

m e ans o f n e rv e fib e r s a nd

co mmunic a ting with th e o ut side

nv . w orld th r o ugh s e n se h air s (pe r ) .

whi ch p e n e tr a te th e cuti cl e M or e .

o f th e s e se n s e o r g a n s occu r a t t h e

a nt e r i o r a nd p os t e r i o r e nd s th a n in

a ny o th e r r e gio n o f th e b o dy ( se e

F i g 90 T h e e pid e r mi s o f the .
,

e a r thw o r m i s a l s o s uppli e d with

e f fe re nt n e r v e fib e r s whi ch p e n e t ra t e

b e tw e e n th e e pid e rm al ce ll s for ming

a sub -e pid e r m a l n e tw o r k ( 1 44 ,

R e pro d u cti o n F E M ALE OR .

GA N S ( F ig —
T h e fe m al e r e .
.

F G 9
I T t l n v
2 nd ng n
ac i e er e e i s I
p ro du ct i v e o rga n s a p ai r o f
. .

a re
t h i nt g m nt f t h
e th
e u e o e e ar

v m p . or n y h i
.
o v a
er
r i
.
e s
,
( )
O se
I n s o m
so r
i t e XIIIa rs
t w o ov i ,
~

j ti th h ti d t ( )
O D in mit XIII n d X I V
“
p n g
ro e c g t h u c s ro u so e s e cua ,

1 ; w3 6 w ; m ne a nd tw o p a i rs o f s e mi na l r e ce ptacle s
e 6 » se se

ll
ce ( F m D hl g n nd S o r s e r ma the cw lying in s o mit e s
s ro a re a
( ) p
.

K pn ) e er
IX and X T h e o va ri e s a re small
'

p e ar sh ap e d b od ie s Th e y li e o ne o n e ithe r side o f th e mid

- .

v e nt r a l line in so mit e XIII a nd a r e a tt ach e d by th e ir l a rge r ,

e nd s to th e v e nt r a l p ar t o f th e a nt e ri o r se ptum T h e o viducts .
1 82 AN IN TR ODU C T ION To ZOOLOGY

w all ,
o ne p ai r in s o mite IX th e o the r in somi t e X The y op e n , .

t o th e o ut sid e th r o ugh th e sp e r m a th e ca l p or e s b e tw e e n so mit e s

IX a nd X and X and XI ,
.

M ALE OR G AN S ( Fi g — T h e m a l e r e p ro ductiv e o r g a ns a r e
.

t wo p a i r s o f gl ov e sh a p e d te s te s ( T) o ne p a i r in so mit e X th e o th e r
-
, ,

in so mit e XI The y o ccupy p o sitio n s in th e so mit e s Simil ar to

.

th a t o f th e ov a ri e s B e hind e ach te sti s i s a ci li ate d fu nne l (S F )

.

shap e d like a r o se tt e ; this i s th e o p e ni ng o f th e sp e r m du ct o r

vas def e r e ns ( VD) Thi s duct p a sse s thr o ugh th e se ptum j us t
.

b ack o f th e funn e l fo r m s se v e r a l conv olutio n s a nd the n e xt e nds

, ,

b ackw a r d n e ar th e v e ntr a l s u r fa ce T h e two sp e r m ducts a r i sing .

o n e ith e r sid e o f th e mid v e nt ra l lin e unit e in s o mit e XII a nd th e n

-

r u n b a ck a s a Singl e tub e o p e ning t o t h e o ut sid e th r o ugh th e

,

sp e r miduc al p o re o n somit e X V In a se xu a lly m a ture e ar th .

w o r m th e te s te s a nd funn e l sh ap e d inn e r op e ni ngs o f th e sp e r m

,
-

ducts a r e incl ose d by l a rge whit e sa c s th e s e mi na l ve s i cle s ( A , ,

B C ) lying in s o mit e s IX t o XII

, ,
The r e a r e thr e e p a i r s o f .

the se sp e r m sacs o ne in so mite IX ( A) one i n somi t e XI (C )

, , ,

a nd th e thi r d in s o mit e XII In s omit e s X a nd XI a re ce nt r al

.

r e se r v o i r s ( B ) .

C OP U L ATI ON M a ny o f th e e v e nts whi ch p r e c e de th e l a ying o f

.

e gg s by t h e e a r thw o r m h a v e no t ye t b e e n l e a r n e d R e pr o d u c .

ti o n i s b e s t kn o wn i n th e c ommo n m anur e w o r m A llolobophor a ,

re tid a h e br e e di ng s e a s on b e gin s e a r ly in th e sp r ing a nd con

f T
.

t i nu e s until l a t e in th e a utumn Se xu a lly m a tu r e w o r m s p osse ss .

a mu ch sw o ll e n clit e ll a r r e gi o n Altho ugh b o th e gg s a nd sp e r ms .

a re p r o du ce d by e v e r y individu a l s e lf f e r t i li z a ti o n d o e s no t t a k e ,
-

pl a c e but th e e gg s o f o ne wo r m a re fe r tiliz e d by th e Sp e rm o f
,

a n o th e r T h e sp e r m a r e t r a n s fe rr e d fr o m o ne w o r m t o a n o th e r
.

during a p roce ss ca ll e d copul a ti o n whi ch usu ally t a ke s pl a ce ,

ju st b e n e a th th e s u r f a ce o f th e e a r th T w o wor m s co m e tog e the r .

with the i r he a ds in o pp o sit e di r e cti on s a nd the i r v e nt r al sur fa ce s ,

o pp ose d a s s h o wn in F igu re 94 A
,
E a ch se cr e t e s a b o ut it se lf ,
.

a tub e o f slim e whi ch e xt e nd s f r o m a b o ut t h e e ighth t o th e thi r ty

»

si xth s o mite Fo u r b andli k e thicke nings p rob ably of th e Slime

.
,
 T HE E ART HWOR M I
AN D AN N E L DS I N GEN E R AL 1 83

tub e s e nci rcl e b oth w or ms at th e a nt e r i or and p oste r i or e dg e s o f

,

th e clit e llar re gi o n s The s e h old th e wor ms fir mly tog e the r and

.

m ay l a t e r a i d in cl osing th e e nd s of th e co co ons T h e slim e tub e .

p ro te cts th e co
co o n a s i t de v e l
o ps a nd co nfin e s
,

t h e se mi n a l fluid
a nd sp e r m a to

p h o r e s .D u r ing
co pul a ti o n sp e r m s a re no t e x ch a ng e d b e tw e e n

t h e sp e r m a th e c a o f th e t w o w o r m s but th e ,

sp e rm a th e c a o f e a ch a r e l o a d e d f r o m t h e
Spe r m a d u ca l p o r e o f t h e o th e r o r s p e r m a t o ,

p h o re s f o r m e d in t h e s p e r m a th e c a o f o ne i n

dividua l a re a tt a ch e d to th e s kin o f th e o th e r .

C OC OON S . C o c oo n s a re f o r m e d e ith e r
du ring copul a ti o n o r a ft e r the two w o r ms
,

h ave s e p ara t e d In th e fo rm e r c ase a b and

.

is se cre te d a b o ut th e clit e ll ar r e gio n o f one

wo rm and thr e e o r m or e opp o sit e s o mit e s of
th e o th e r wo r m At fi r s t th e c o co o n s a re
.

p e rfe ctly whit e ( Fi g 94 B ) but aft e r de p osi F I G 94 A th n . .

,
e a
,
.
,
t i gm nt s e r or se e
tio n th e e xpo sur e t o th e a i r ch a ng e s the m to f tw l t o o co u a
p
a y e ll o w c ol or . Whe n copul a ti o n is co mpl e te d i ng thw m , e ar or s .

th e w or m s sl o wly withd r a w b ackw a r d f r o m th e Sli m t b m e u es e

Slim e tub e a nd coco o n As th e co coo n i s

.
i l th p i c rc e e a r

f m t h 8 th t
slipp e d o ve r th e h e a d i ts e nd s co nt ra ct ro e o
, ,
th 33d g e se
fo r ming a tur nip sh ap e d cap sul e l ying withi n
-
m nt ,
B e . co
,

th e slim e tub e ( F i g 94 B) . T h e co coo n ,

.
n f hl y coo re s
,

a ft e r d e p o siti o n c o nt a in s o n a n a v e r a g e fou r
, d p it d f n
, e os e , o a

e gg s ,
e a ch o f which h a s b e e n p e n e t ra t e d by thw m e ar or su r

nd d by n
fro m o ne t o a s m a ny as nin e sp e rm a to z o a .
rou e o e

h lf f li m a o a s e
T h e nu cl e us o f a s ingl e s p e r m a t o z oo n unit e s
t b (Af t r u e . e
with the e gg nucl e u s ; th e r e m aining spe rm a F t in J rn oo ou .

to zoa di si nt e g r a t e ( 1 4 1 , M ph ) or .
1 84 AN I N TR OD U C T ON T o I ZOOLOGY

B e fo re co n side ring th e e mb ryo l o gy of the e a r thw orm th e ,

de v e l opm e nt o f the e gg s and Sp e r m a to zoa a nd th e hi sto ry of the se

p re vi o us to fe r tiliz a ti o n mu s t b e de scr ib e d .

SP E R M ATOGE N E SI S Du r ing th e d e v e l opm e nt o f th e sp e r m a t o

.

z oa ,
th e p rim or di a l g e r m ce ll s se p ara t e f ro m th e t e s t e s a nd li e ,

in the c a viti e s o f th e s e min a l v e sicl e s T he nucl e u s o f e ach ge r m

.

c e ll di vide s into 2 4 8 or 1 6 d a ughte r nu cl e i which b e co m e

, , , ,

a rra ng e d in a singl e l a y e r n e a r th e su r f a c e o f th e c e ll C e ll w a lls .

no w app e ar e xt e nding inw ar d f r o m th e p e r iph e r y ,t h e re s ult b e ing

,

a c o l o ny o f c e ll s a tt a ch e d by cyt o pl a s mic p e d e s t a l s to a c e nt ra l

non-nucl e a t e d m ass Th e ce ll s o f th e co l o ny th e n divide i ncr e a s

.
,

ing th e numb e r to 3 2 64 1 2 8 o r m o re A di sso ci a ti on o f th e

, ,
.

c ol ony int o se ve r a l p ar t s now t ak e s pl a ce e a ch p a r t cont a ining ,

a numb e r o f sp e r m a t o g o ni a Su ch a sp e r m a t ogo ni a l g r o up fin ally

.

b e com e s a sphe ric al m or ul a o f 3 2 p rim ar y sp e rm a tocyt e s whi ch ,

a r e s till f a s t e n e d by cyt opl a s mic th r e a d s t o a ce nt ra l b o dy c all e d

th e blas tophor e . E a ch g r o up o f p r im ary sp e r m a t o cyt e s giv e s

ris e to 64 s e c o nd a ry sp e r m a t o cyt e s a nd th e se divid e to f or m 1 2 8
,

sp e r m a tid s. T h e l a tt e r th e n m e t a m o r ph o se int o sp e r m a t o z oa .

T h e numb e r o f ch r o m oso m e s in th e sp e r m a t o z oa i s sixt e e n ; this

i s o ne h alf the numb e r c o nt a in e d in th e so m a ti c ce ll s th e re du e ,

tio n h aving ta k e n pl ace dur ing m a tu ra ti o n by th e union of th e

ch romoso m e s tw o by tw o in th e se co nd a ry sp e rm a to cyt e s and a ,

sub se qu e nt s e p ar a ti o n wh e n th e sp e rm a tid s w e re form e d ( 1 3 8 ,

D u ring th e l a te r s t age s o f copul a ti o n the sp e rm a toz oa a r e

drawn int t o h e r o e tt
s e -like funn e l s ( Fi g 93 S F ) o f the vas ad ej e .
,

r e nti e,
p as s th ro ugh the se ducts a nd a r e t ran sfe rr e d to th e,

sp e r m a th e c a o f th e o th e r w o r m He r e th e y are sto r e d unti l th e

.

co c o o n p asse s o v e r th e op e ning s o f th e sp e r m a th e ca du ring th e

withdra wa l o f th e wo rm wh e n s om e o f th e m p a ss into th e nutri

,

e nt fluid w hi ch h as b e e n se cr e te d into th e c avity o f th e co coon .

OOGE N E SI S . T h e e gg s d e v e l op in th e o v a r i e s ( Fi g 93
-
.
,

Th e se are p e a r Sh ap e d b o die s co mp o se d o f e gg c e ll s in va ri ous

-

st age s of growth T h e b asa l p or tion of e ach ovary con si st s of

.
 K
.

95 . St a ge s i n t h e l o gy o f t h e e a r thw o r m A t wo ce ll e d s t age ;
e m b r yo .
,
-

B f o u r ce ll e d s t age
,
- C Si x ce ll e d s t a ge
,
- D e i ght ce ll e d s t a ge
,
- E bl a s ,

t u l a i n se cti o n ; F b l a s t u l a fl a tt e ne d ; G s t age sh o wi ng m e sob l a s ti c

, ,

ba nd s ; H ga s t r u l a ; J o l d e r ga s t r ul a ; K l o ngit u d i na l s e cti o n o f l a t e r
, , ,

s t a ge bl b l a s t o coe l ; bp b l a s t opo r e ; c
. .
,
.
,
c e ct o d e r m ; e u
.
,
e nt o d e r m ;
.
,

m a c m a cro m e re ; m e s m e so d e r m
.
,
.
, m i c m i cro m e r e ; mm m e so m e r e s
.
,
.
,
.

( Fro m B o u r ne a f t e r Wilso n ) .
 T HE EAR T H WOR M AN D ANN EL DS I N GEN E R AL I 1 85

p rimitive ge r m c e ll s ; f ro m h e r e to th e p o inte d e nd o f the ov a ry

the ce ll s in cr e a se in s iz e th o s e t o w ar d th e e xt r e m e e nd b e ing
,

re co gniz a bl e a s e gg s ,
e a ch o f whi ch i s s u rr o und e d by a f olli cl e

o f s m a ll nut r itiv e c e ll s .Egg s se p a r a t e f ro m th e e nd o f th e o v a r ie s

a nd p as s int o th e b o dy c a vity F ro m h e r e th e y e nt e r th e cili a t e d
.

funn e l o f th e o vidu ct the nc e into th e e gg s ac ( Fi g 93 R) whe r e

, .
, ,

p a r t o f the m a tu ra ti o n p r o c e s se s o ccu r F r o m h e r e the y e ith e r .

p a ss o u t into th e ca vi ty o f th e Slim e tub e a nd a r e co nv e y e d fro m

t h e e xt e rn a l op e ning s o f th e o vidu ct o n s o mit e X I V to th e co
coo n o r e nt e r th e c o c oo n wh e n it p a ss e s o v e r thi s s o mi t e du r ing
,

de p o sitio n
F E R TI LI Z ATI ON . T h e e gg s a re p e n e t ra t e d by sp e r m a t o zo a
-

a ft e r th e cocoo n i s s h e d At thi s tim e th e y h a v e no t co mpl e t e d

.

the i r m a tu ra ti o n p ro ce sse s ( 1 4 1 ,

E m b ryo l ogy . T h e e gg s o f t h e e a r thw o r m a re holoblas ti c ,

but cle a vage i s un e qu a l th e fi r s t divi sio n r e sulting in o ne l arg e

,

a nd o ne sm a ll ce ll ( F i g 95 A )
. T h e se c
,
o nd c l
. e a v a g e di vid e s

th e sm a ll ce ll int o t w o e qu a l p a r t s a nd cut s o ff a s m all ce ll f ro m the

l a rge r ( B ) B e ca u se o f th e i r ( Eff e r e n ce in siz e th e l ar g e ce ll s a r e
.
,

ca ll e d m acr om e r e s (m ac ) a nd t h e s m a ll c e ll s
. mi cr ome r e s ,

C l e a v a g e b e co m e s i r re gul a r a ft e r th e se c o nd di vi si o n B o th th e .

m acro m e re s a nd mi cr o m e r e s co nti nu e to divid e the numb e r o f ,

ce ll s in cr e a sing to si x ( C ) e i ght ( D) e tc So o n a c a v ity th e

,
.
,
,

bla s tocoe l a pp e ar s b e tw e e n th e mi cr o m e r e s a nd m a cro m e re s ,

a nd a h o ll o w sph e r e r e co gniz a bl e a s a bla s tu la r e s ult s ( E) T wo .

o f th e l a r g e r ce ll s o f th e bl a s tul a lying s i d e by s i d e b e h a v e di f f e r

e ntly f ro m th e o th e r s Th e y p ro j e ct d o wn into th e bl a s to coe l

. ,

a nd by r e p e a t e d d ivi s i o n s
,
giv e ri se to t w o ro w s o f sm all c e lls
,

( m e s a
. n d m m . in F a n d G ) B e c a u se o f
. t h e f a c t th a t t h e se t w o

ce ll s giv e r i s e t o th e m e so d e r m th e y a r e t e r m e d m e s om e r e s a nd
, ,

th e tw o r o w s o f ce ll s d e r iv e d f r o m th e m th e m e s oblas ti c ba nds ,
.

Whi l e the m e so m e r e s a r e dividing th e bl as tul a b e co m e s fla t ,

t e n e d ( F) th e l a rg e r ce ll s fo r m a pl a te o f cl e ar c olumn a r ce ll s
,

a nd t h e s m a ll c e ll s sp r e a d ou t int o a t hi n d o m e s h ap e d -

e pith e lium
1 86 AN IN TR ODU CTION To Z OOLOGY

B il a t e r a l
symme try is a l re a dy e st abli she d a t thi s e arly p e ri o d ;
th e m e sobl a s tic b a nd s li e al o ng wh a t will b e co m e th e l o ngitudin a l
a xis o f th e futu r e w or m a nd t h e m e so m e r e s o ccupy t h e p o s t e r i or
,

e nd . A gastr u la i s no w fo r m e d by th e inv agin a ti o n o f th e pl a t e

o f l arg e c e lls ( H) ; th e e dg e s o f th e ca v ity thu s p ro d u ce d f old in

until o nly a slit re m a in s (J) Thi s slit i s the blastopor e .

a nd th e c a vity is t h e e nte r on which l a t e r b e c o m e s p a r t o f t h e

,

a lim e nt a r y c a n a l Soo n th e s lit cl ose s e x ce pt a t o ne e nd wh e r e a

.
,
.

p or e th e futur e mou th r e m a ins T h e th re e ge r m l aye rs a r e a t

, , .

thi s tim e quite di stinct a nd a re w e ll shown in a l ongitudin al

,

se cti o n o f th e g a str ul a ( K) T h e l a r g e cl e a r ce ll s whi ch i nv agi

.

nat e d lin e th e e nt e r o n b e co ming th e e ntod e r m

, th e d o m e
shap e d e pithe lium o f sm a ll ce lls cov e r s th e o ute r s u rface a nd r e pr e
se nt s the e ctode r m a nd b e tw e e n th e se t w o l a y e r s a re th e tw o

ro w s o f m e sobl a stic b a nds which giv e r i s e to th e m e sode r m (me s )

( 1 5 7)
The d e ta i le d histor y o f the se ge r m laye rs i s too l ong and com
plic a te d to b e discu sse d in a b oo k o f thi s ch ar a cte r T he de ve l op .

m e nt of the m e so de rm a l l a y e r s and coe l o m Sho uld how e v e r be , ,

m e ntio n e d Th e m e s o de r m b e co m e s s e p ar a t e d into two l aye r s

.

b e twe e n whi ch a cavity the coelom i s fo r m e d Th e o ute r l ay e r

, ,
.
,

c all e d th e s oma tople u r e cling s to th e e cto de r m a nd give s ri se to

, ,

th e mu s cl e s o f th e b o dy w a ll ; t h e inn e r l a y e r c a ll e d th e s pla nch ,

nople u r e re m a in s a tt ach e d to th e e nt e ro n a nd giv e s r i s e t o th e

,

mu scl e s o f th e a lim e nt a ry c a n a l All o th e r s tr uctu r e s o f m e so

.

de rm al origin a r e de riv e d fro m th e se tw o l aye r s Aft e r th e e s .

t abli sh m e nt of th e g e r m l a y e r s a s d e s c rib e d a b o v e th e e mb ryo ,

e l o ng a t e s a nd fin a lly b e co m e s v e rmi fo r m
, e sc aping f r o m th e ,

c o coo n in a b o ut tw o o r thre e w e e k s .

B e h avi or E X TE R N AL STI M U LI
. T h e e xt e r n a l s timuli th a t
.

have b e e n m o s t fr e que ntly e mpl o ye d in s tudying th e b e ha vi or o f

e a r thw o r m s a r e th o se d e a ling with t hi gm o t ropi s m ch e m o t r opi sm , ,

a nd ph o

T HI GMQfl lg
t o t ro pi s r

I SM
n( 13 5 ,
1 4 5,
M e ch a ni ca l
1 5 0 ,
1 5 2 ,53
1 5 5,
,

s timu la ti o n ,
1

if ntinuo us a nd
. co

no t too st rong ca lls for th

, a p o sitive re a cti o n ; th e w o r ms liv e
1 88 AN IN T RODU CTION To ZOOLOGY

s e e m to Sh o w that th e wo r m s are s tim ul a t e d in diff e r e nt de gr e e s

by th e m sinc e th e int e rv a l of tim e b e twe e n th e applic a ti on of th e
,

s tim ul us a nd t h e r e a cti o n di f fe r s f o r e ach sub s t a n ce b e ing s h o r t e r ,

f o r sodi um chl orid e a nd s u cc e ssiv e ly l o nge r f or th e o th e r s in th e

,

o rde r n a me d In m an the se sub s t a nc e s all t a st e pra cti

c a lly a like b e ca u se of th e p r e se n ce o f chl orin e
, .

P H OTOTR OP I SM N o d e fi ni t e vi su a l o rg a n s h av e b e e n d i s
.
-

co v e r e d i n e a r thw o r m s but n e v e r th e l e ss th e s e a nim al s a r e v e ry

,

se n sitiv e t o light a s i s p r o v e d by t h e f act th a t a s udd e n illumin a

,

tio n at ni ght w ill o ft e n c a use th e m to d ash lik e a r abbit into

th e i r bur ro ws One inv e stig a to r cl aim s t o h av e fo und ce ll s in
.

th e e ct o d e r m e sp e ci a lly i n th e p r os t o mium a nd p o s t e r i or e nd
, ,

which a ct a s vi su al o rg an s T h e e nti r e s u r fa c e o f th e
b o dy ho w e v e r i s se n sitive to light a lth o ugh th e a nt e r i or r e gi o n
, , ,

i s m o r e s e n siti v e th a n t h e t a i l a nd th e middle l e ss than e ith e r o f

,

t h e o th e rs If a n a nim al which i s lying al o ng th e g ro und with

.
,

i ts t ail clinging t o th e t op o f th e bu rr o w i s s udd e nly illumin a t e d

, ,

it p ro mptly withd ra w s int o i ts h ol e ; if not in t o uch wi th i ts b u r '

row ,
it will cr awl a w a y fro m th e so u r ce o f light Ve r y slight .

diff e r e nc e s in the int e n sity o f th e light a re di stingui she d sin ce , ,

if a choic e o f tw o illumina t e d r e gi o n s i s give n th a t m o re faintly ,

light e d i s in th e m aj o rity o f ca se s se le ct e d Thu s f a r we have

, ,
.

c on side r e d light as ca u sing a n e g a tive r e sp on se ; but a p ositive

re a cti o n t o f a int l i ght h a s b e e n d e m o n s t r a t e d f o r A llolobophor a

f a tid a Th i s p o sitiv e ph o t o t r opi s m t o f a int light m ay

a cco unt f or t h e e m e r g e n ce o f t h e w o r m s f r o m th e i r bu r r o w s a t

n ght E xp e r im e nt s with l ight s o f diffe r e nt co l ors Sho w tha t r e d

i
.

i s p r e fe rr e d to a ny o th e r g r e e n b e ing n e xt a nd blu e l a s t a nd that

, , ,

th e i n t e n sity of th e c o l o re d r ays de t e r min e s th e e ff e ct

C OMB I N ATI ON S AN D I N TE R F E RE N C E OF STI M U LI — I t has b e e n
.

sh own tha t c o nt a ct a nd che mi ca l stimuli m ay co mbin e a s in ,

th e ca se o f th e bu rr o wing r e a cti o n o f A llolobophor a f a tid a In .

o th e r inst a n ce s s timuli int e r f e r e w i th o ne a n o th e r ; f o r e x a mpl e ,

light ca ll s fo rth no r e a cti o n if th e a nim a ls are fe e ding o r ,

 ti thw m A h d n d f fi m e n t s
e ge ne ra o n i n t h e e a r g
o r e a e o v e s e
FI G .
9 6 . R .
,

t t i i f w m B t il r e e n e r a t e d f r om
e
r g e n e r a e d f r o m t h e p o s e r or p e e c o a o r ,
a g
t i i f w orm ; C t il r e e n e r a t e d f r o m a n a n t e i
r or
h
t e p o s e r o r p e ce o a ,
a g
i e o f a w o r m ( F r o m M o r ga n )
p c e
.
.
1
.

90 AN IN TRODU CTION TO ZOOLOGY

Th e ra e t g e n e ra tive g ro wth d e p e nds up o n the a m o unt of

of r e

old tis s u e re m ov e d Thus if only a f e w se gm e nt s o f the p oste ri or

.
,

e n d ar e cut o ff a n e w t a il,
r e g e n e r a t e s v e r y s l o wly ; if m o r e is

r e m o v e d th e ne w tissu e is a dd e d m o r e rapidly
,
In fa ct the ra te .
,

o f gr o wth inc r e a s e s up t o a ce r t a in p o int a s t h e a m o unt r e m ov e d

incre ase s T he f a ctors r e gul a ting the r a te o f r e ge n e ra ti o n h av e

.

no t ye t b e e n f u lly d e t e r min e d a lth o ugh s e v e r a l p o ssibl e e xpl a n a

,

ti ons h ave b e e n sugge ste d .

G r a f ti ng P i e c e s o f e a r thw or ms m a y b e g ra ft e d up o n
.

o th e r w or m s witho ut much d i fli cu lty Se ve ra l re s ults are .

sho wn in F igure 9 7 Th r e e pi e ce s m ay b e so uni t e d a s to

.

p r o duc e a v e r y l ong w o r m ( A) ; th e t a i l o f o ne an i m a l m ay

be g ra ft e d up o n th e sid e of a n o th e r p r o du cing a do ubl e ,

t ail e d w or m ( B ) ; o r th e a nt e ri o r e nd o f o ne in di vidu al m ay
b e unit e d with th a t o f a n o th e r ( C ) In a ll the s e e xp e rim e nt s
.

th e p ar ts w e r e h e ld t o g e th e r by th re a ds un til th e y b e c a m e
unit e d .

2 . AN N E I I DS IN GE N E R AL

T h e tw o chie f cl asse s o f Ann e lid s a re th e C h a topod a o r Ea r th

wo r ms a nd m a rin e Ann e lid s a nd th e Hi r udi n e a or Le e che s ,
.

C l ass C h a topod a m ay b e divid e d into tw o subcl a sse s th e Oli ,

go ch a t a t o,
whi ch t h e e a r thw o r m b e l o ng s a nd t h e P o lych a t a ,
.

Tabl e VI co ntrasts th e ch ar acte r s o f the se tw o divi si ons and ,

F igu re 9 8 sh o ws o ne o f th e m os t c o mm o n P o lych a ts th e m a r in e ,

a nn e lid N e r ci s
,
.

Th e m e mb e rs o f th e C l as s Hi r udin e a h av e n e ith e r se t a nor

p arap o di a The y m ove by m e a n s o f a v e nt r al s uck e r n e ar th e
.

p ost e ri or e nd a nd a s uck e r m o uth o n th e v e ntr a l s urf a ce at the

,

ante ri or e nd Th e y are fl a tt e n e d do r so v e nt r ally T h e gr oov e s

.
- .

o n th e o ut sid e o f th e b o dy d o no t cor r e sp o nd in numb e r o r p o si

tio n to the s e pt a within the r e b e ing se ve r al e xt e r n al rings to e a ch

,

int e r nal se gm e nt T h e coe l o m i s no t a s l arge a s in th e C h a tOpods

.
,

b e ing r e duce d to the caviti e s in w h ich th e g e r m ce lls li e and a ,

FI G 9 7
. . Gr a f ti ng i n th e e ar thwor m . A, i
u n on o f thr e e p i e ce s t o m a k e a

lo ng wo r m ; B ,
i
u n on o f t w o p i e ce s t o m a k e a d o u b l e t a il e d w or m ;
-

C a nt e r i or a nd
, po s t e r i or p i e ce s u ni t e d t o m a k e a s ho r t wo rm .
( Fro m
M o rga n ) .
1 9 2 AN IN TR ODU C TION To Z OOLOGY

T ABLE VI

THE C H ARA C TE R S OF OLI G OC HE TS AND P OLY C HE T S C ON TR ASTE D

OLI GOCHE I A
' ‘

P OLY CHE TA

No i l lo com o to r
sp e c a p ro tr u s i ons P a rapo di a m a ny se ta .

(p a r a po d i a ) ; f e w s e t a .

No o th e r t
e x e r na l a ppe nd age s . Ext e r na l a pp e nd a ge s i n th e f or m
'

of

t
a n e nna , g ill s ,
a nd c rri i (se e F ig .

He r m ap h r o d ite . Se xe s u s ua ll y se p ara t e .

De v e l o p m e nt d i r e ct . A m e t a m o rp ho s i s i n d e v e lo p m e nt .

Aq u a ti c ( f r e s h -w a t e r ) or t e rre s t ri a l . Aq u a ti c (m ar i ne ) .
 C HAP TE R XI

THE C RAYFI S H AND AR TH R OP OD S I N G EN ERAL

I . THE C R AYF I SH

( C a m ba r u s vi r i li s Gi rar d) 1

C RA Y F I SHE S inh abit fre sh-wa t e r l a k e s p o nds and st re am s

, , .

The sp e ci e s C a mba r u s vi r i li s i s co mm o n in so m e o f th e c e nt ra l
s t a t e s a nd C a mba ru s a i ni s in th e e a s t e r n p a r t o f t h e c o unt r y
fi .

T h e l ob s t e r is so n e a r ly lik e th e cra yfi s h in s t r u ctu r e th a t th e

a n a t o mi ca l p o r ti o n o f thi s ch a pt e r m a y b e a ppli e d a l so in l arg e

p a r t to this anim al In E u r op e th e m os t c o mm o n c r ayfish is

.

Astacu s f lu vi a ti li s a Sp e ci e s m a d e f a m o us by Huxl e y s cl a ssic a l

’
,
”
wo rk T h e C r ayfi sh .

Th e c ra yfish C a m ba r u s vi r i li s i s usu a lly fo und c o nce a l e d und e r

, ,

rock s o r l o g s a t th e b o tt o m o f p o nds a nd st r e a m s He r e it li e s .

with i ts he a d t o war d the e nt r a nce to its hiding pl ace Wh e n .

c ra wling ab o ut o r s wimming in the op e n w a t e r i ts h ar d sh e ll ,

h e lp s p r o te ct it fro m fi sh while i t s col or which r e s e mbl e s the

, ,

b o tto m te nds t o m a k e its de t e cti o n difficult C r a yfi sh e s m ay b e

,
.

captu re d e asily by ha nd with a ne t or by fi shi ng f o r the m wi th a

, ,

s t ring b a it e d with a pi e c e o f m e a t Th e y th r ive in an a quar ium

.
,

a nd th e i r e nti r e l i f e hi s t o ry m a b e o b se r v e d in t h e l a b o r a t o ry
y . .

Th e y e a r ly d e c r e a se i n th e numb e r o f l o b s t e rs a v a il a bl e f or f oo d ,

1
Th e l t e li f e hi s to ry a nd th e d e t a il s o f th e a na t o m y a r e no t k nown f o r
co m p e ,

a ny s i ngl e spe c i e s o f t h e ge nu s C a m ba r u s T h e f o ll o wi ng a cco u nt th e r e f o re

.
, ,

m u s t ne ce ssa r il y b e a co m po s it e co nt a i ni ng no t o nl y o b s e r v a ti o ns o n v a r i o u s
s p e ci e s o f C a m ba r u s b u t a l s o o n th e E u ro p e a n c r a y fi s h As ta cu s ( P ota m obi u s )
, ,

fl i a ti li s T h e d i ff e r e nce s b e tw e e n th e s e cr a yfi sh e s a r e h o w e v e r so sli ght

u v .
, ,

a s t o b e o f littl e i m p o r t a nce e xce p t i n a d e t a il e d m o no gr a h

p .

O 1 93
1 94 AN IN TR ODU CTION TO Z OOLOGY

a nd ste adi ly incre asing de m and f or cr ayfi she s will undo ubt
th e ,

e dly soo n m a k e it w o r th whil e t o ra i s e th e l a tt e r f o r m ar k e t

Exte rnal F e a tur e s — E X OSK E LE T ON

. T h e o ut sid e o f th e b ody
.
-

o f th e c r a yfi sh i s co v e re d by an e xt r e m e ly h a r d chitin o u s cu ti cle

imp r e gn a t e d with lim e salts This e x osk e l e to n i s thinn e r a nd

.

fl e xibl e a t the j o ints all o wing m o ve me nt A d e lica te cuticl e

,
.

o f th e s a m e subst a nc e ( chiti n) w a s n o t e d in t h e e a r thwor m

( p .

RE GI ON S OF T HE B ODY Unlik e th e e ar thwor m t h e b o dy o f

.
,

the cr ayfi s h sh o w s tw o di s tinct r e gi o n s a n a nt e r i or r igid p or ti o n

, ,

th e ce phalothor a x and a p os t e ri or s e r i e s of s e gm e nts th e a bdome n

, , .

T ra ns v e r se tio n thr o u gh th e a b dom e n o f th e cra yfis h

.

FI G . 1 00 . se c .

d o r sal abdo m i na l ar t e r y ; E M e xt e nso r m u s cl e s o f th e a b do m e n ;

,

EP e pi m e ro n F M fle xo r m u s cl e s o f abd o m e n ; M m u scl e s o f a ppe nd

, , ,

a ge ; N e nd opo di t e ; N G ne r v e ga ngli o n ; P
, , p r o t o p,
o d it e ; P L .

p l e uro n ; P R i nt
,
e s ti ne ; S s t e r,
nu m ; T t e r g
,
u m V v e n tr a l a b ,
d o m i na l

a r t e r y ; X e xopo di t e
, ( Fr o m M a r sh all a nd Hur s t )
. .

Th e ntire b o dy i s se gm e nt e d but th e j o int s h ave b e e n oblit e rat e d

e ,

o n t h e d or s al s u r f ace of t h e ce ph a l o th o r ax .
 Y I
T HE C R A F SH AN D ART HROP ODS I N GEN E RAL 1 95

STR U C TUR E OF A SE GME N T — typic a l se gme nt i s shown i n

. A
c ross s e ctio n in Figur e 1 00 It con sists of the foll o wing p a r t s :
.

a c o nv e x dor s a l pl a t e th e te r gu m ( T) ( 2 ) a v e nt r a l t r a nsv e r s e b ar
, , ,

th e s te r nu m (S ) ( 3 ) pl a te s p ro j e cting d o wn a t t h e sid e s t h e
, ,

p le u r a ( P L ) a nd , ( 4 ) sm a ll e r pl a t e s b e tw e e n t h e pl e u ra a nd the
b as is of th e limb th e e pi me r a ( E P )
,
.

C E P H AL OTH OR AX T h e c e ph a l o th o ra x c o n si s t s o f s e gm e nts
.
-

I XIII whi ch a r e incl ose d d o r s ally a nd l a t e ra lly by a cuticul a r

—
,

s hi e ld th e ca r a pace
,
An ind e nt a ti o n t e r m e d th e ce rvi cal gr oove
.
, ,

r uns a c ro ss t h e mid d o r s a l re gi o n o f th e c a ra p a ce
- a nd o bliqu e ly ,

forw ar d on e ithe r side s e p ara ting th e ce phali c or h e a d r e gion

,

f ro m the p o st e ri or thor aci c p or ti o n The “ i nt e r i o r p o int e d e xt e n

.

sio n of th e c ar ap a ce i s kn o wn a s the r ostr u m B e n e a th this o n .

e ith e r sid e i s a n e ye a t t h e e nd o f a m o v a bl e p e duncl e T h e m ou th .

is situa t e d on th e v e nt ra l s u rf a c e n e ar the p o st e ri or e nd of the

he a d re gio n It i s p a r tly o b s cu r e d by th e n e ighb o r ing app e nd
.

age s . Th e c a ra p a c e o f th e tho r a x i s se p a ra t e d by br a nchi o

car di ac gr oove s int o th re e p ar ts a m e di a n d o r s a l l o ngitudin a l
,

strip th e ar e ola and two l arge c onve x fl aps o ne o n e ithe r side

, , , ,

th e br a nchi os te gi te s whi ch p r o t e ct th e gill s b e n e a th th e m

,
.

AB DOME N — I n th e a bd o m e n th e re ar e six s e gm e nt s and a te r ,

.

min al e xte nsi o n th e te lson b e aring on i t s v e nt ra l s u r fa ce th e l ongi

, ,

t u di nal a n a l op e ning Whe the r or no t th e t e l son is a t r ue se g

.

m e nt is still in di sput e ; w e shall a dopt th e Vi e w th a t it i s no t .

Th e fi r st a bd o min al se gm e nt (X I V) i s s m a ll e r th a n th e o th e r s a nd
l acks the pl e ur a Se gm e nts X V X IX a re lik e th e typ e d e scrib e d
.
—

a b ove .

AP P E N DAGE S — With th e p o ssibl e e xc e pti o n o f the fi r st ah

.

d o mi nal s e gm e nt in t h e fe m a l e e v e r y s e gm e nt o f th e b o dy b e a rs ,

a p ai r o f j o int e d a pp e nd a g e s The se a r e a ll v ari a ti o n s o f a com

.

m o n typ e ( Fi g . c o n si s ting o f a b a s al se gm e nt th e pr otopo ,

di te ( P ) whi ch b e a r s tw o b ra n ch e s a n inn e r e nd opodi te ( N )

, , ,

a nd a n o ut e r e xo odi te ( X ) B ginning t t h nt e ri o r e nd t h e
p e a . e a ,

a pp e nd a g e s a r e a rr a ng e d a s fo ll o w s ( Fi In f ro nt o f th e
g .

m o u th a re I
( ) t h e a n te n n u l e s a n d II
( ) th e a n te n
,
na ; t h e m o uth
1 96 I N T R OD U C T ON T O Z OOLOGY I

posse sse s ( III ) a p ai r of man

di ble s b e hind whi ch a r e (I V)
,

th e fi r st a nd ( V) th e s e c o nd
,

-e x m axi lla ; t h e th o r a cic re gio n

b e a r s ( VI ) th e fi rs t (VI I ) the ,

s e co nd a nd (VIII) th e thi r d
,

m a xi lli pe d e s ( IX ) th e pi n ,

che rs or che li ed s ,
p a nd (x
X III ) fo u r the r p ai rs of o

wa lki ng le gs ; b e n e a th the
abd o m e n a r e six (X I V XIX)
—

p ai r s o f swi mme r e ts so m e of ,

which a r e much modifi e d .

Tabl e VI I give s b rie f d e

scr i p ti o ns o f th e di ff e r e nt ap

p e ndag e s and shows the ,

m o difi ca ti o ns d u e to dif
f e r e nce s in fun cti o n T he .

functi ons of so me of the ap

p e ndage s a re still in do ubt .

Thr e e ki nd s of appendage s
ca n b e di stingui she d in the
a dult cr a yfi sh ; ( 1 ) th e f oli a
ce au s e g th e s e co nd m a xill a
1 02 . h app e nd age s
Typ e s o f cra yfi s . ,
. .

A s e co nd m a xill a f o li a ce o u s t yp e
, , ( Fi g. 1 0 2, A) , ( )
2 th e bi r a
— ba so po di t e ; 5 e nd op o d it e
1 4 , , m ou s , e g th e swimm e re t s ( Fi g
. . .

6 e xop o d it e ; e p s ca p h o gna thit e

1 0 2 , B ) , a nd (3 ) th e u ni r a
. .
, ,

( Fr o m C a m b r i dge N a t u r a l Hi s
t o ry ) B s wi m m e r e t b i r a m o u s
.
m ou s , g ew a.lking l
. e gs
th e
, ,

t y p e ; cx bs p r o t o p o d i t e ;
. .
,
( g
F i . 1 0 2 C ) A ll o,
f th e se .
.

ex .e xo po d it e ;
,
en e nd o p o d it e .
,
. app e nd a g e s h ave do ubtl e ss
( Fro m La nk e s te r s T r e a ti s e ) b e e n de r iv e d fr o m a Singl e
’
.

C se co ri d w a l k i ng le g u ni ra m o u s
, ,
typ e th e m o difica tion s b e ing
,
typ e ; e xp bp p ro topo d it e ; i p
d u e t o th e fun cti o n s pe r
. .
, .
,

mp cp pp dp se gm e nt s o f e n
.
,
.
,
.
,
.
,
.

d o po di te ; e p e pi po d it e ( Fro m .
, .
forme d b y the m The .

C a m br i d ge N a t ura l Hi s t ory ) . bira mo us typ e prob ably

1 98 AN IN TRODU CTION TO ZOOLOGY
 Y I
T HE C R A F SH AN D ART HR OP ODS I N GEN E R AL 1 99
 200 AN INTRODU C TION TO Z OOLOGY

re p r e s e nts th e c o nditi on fro m which the o the r typ e s de ve l op e d .

Th e uni ra m o u s w a lking l e g s f or e x a mpl e p a s s t h ro ugh a

, ,

bi ra mo us s t age during th e ir e mb ryol ogic a l de ve l opm e nt Ag ain .

,

th e b i r a m o u s e mb ry o ni c m a xillip e d e s a r e co nv e r t e d int o th e
fo li a ce o u s typ e by th e e xp ansi o n o f the i r b a sal se gm e nts ( 1 66)
Ge ne r a l I nte n al Anatomy
r .
— T h e b o dy o f th e c r a yfi sh
contain s all o f the imp or tant sys te m s of o rg an s ch ar a ct e ri stic o f
t h e hi gh e r a ni m a l s T h e c oe l o m i s no t a l ar g e ca vity a s in th e e a r th
.
,

w o r m but i s r e str ict e d to th e c a viti e s of th e r e p r o ductive o rg an s

, .

C e r t ain o f th e o rg a n s a r e m e t a m e ric ally a rr a nge d e g th e ne rv ,

. .

ou s sys te m ; o th e r s lik e th e e xcr e tory organs are co nce ntra t e d ,

int o a sm all sp a ce T h e sys t e m s of orga n s a nd the ir functi on s

.

will be p r e se nt e d in th e fo ll o wing or de r : ( 1 ) dige stiv e ( 2) v as ,

cular ( 3 ) r e spi r a t o ry (4 ) e xc r e t o r y
, , 5
, ) n e r v o u s ( )
6 s
,
e n s e o r g a n s ,

( 7) mu s cula r a nd ( 8) re p r o du ctiv e
,
.

Di ge s ti ve Sys te m — .T h e a lim e nt ar y c a n a l o f C a mbar u s co n

si st s o f th e foll o wing p art s ( Fi g 1 0 1 ) .

( )
1 T h e m ou th (4 ) o p e ns o n t h e v e nt r a l su r f a c e b e tw e e n th e
j aw s
( 2 ) T h e oe so h
p g a u s ( 2 0) is a S h or t tub e l e a ding f r o m th e m o uth
t o th e s t o m a ch .

(3 ) T h e s to m ac h i s a l a rg e c avity divid e d by a c o n s t r i cti o n

into an a nt e ri or cardi ac ch a mb e r (2 1 ) and a s m a ll e r p o st e rior

l r i c ch a mb e r In t h e s t o m ach a r e a numb e r o f chitin o u s
py o

o s si cl e s o f u s e in ch e w ing the f oo d a nd co ll e ctiv e ly kn o wn a s th e

,

g as tr i c m i l l
. T h e m o s t imp o r t a nt o f th e s e are ( a ) a m e di a n c a r

dia c o ssi cl e (b) a m e di a n u r oca rdi a c o ssicl e (c) two l a te ra l pt e ro

, ,

ca r di a c o ss i cl e s ,( )
d a p a i r o f l a t e r a l zyg o car di a c o ss i c l e s ( e ) ,
a

pyl ori c ossi cl e and (f ) a p r e pyl o ric o ssi cl e T h e o ssi cl e s a r e abl e

,
.

t o m o v e o ne up o n a n o th e r a nd b e ing co nn e ct e d with p o w e r ful

, ,

mu scl e s are e ffe ctive in g rinding up th e foo d On e ithe r side o f

,
.

th e pyl o ri c ch a mb e r e nt e rs a du ct o f th e dig e s tiv e gl a nds a nd ,

a b o ve i s th e Op e ning o f th e s m all ca cum .

At c e r t a in tim e s t w o ca l ca r e o u s b o di e s kn o wn a s gastr oli ths , ,

a r e p r e s e nt in t h e l a t e r a l w a ll s o f t h e ca r di ac ch a mb e r o f th e
20 2 AN INT R ODU CTION TO Z OOLOGY

susp e nde d a numb e r o f a m e b oid c e ll s the blood corpu scle s or ,

a m e b ocyt e s T h e p rin cip a l f u ncti ons o f th e bl oo d a re th e

.

t ra n sp or t a ti o n o f fo o d m a t e r i al s f ro m o ne p ar t o f th e b o dy t o

a n o th e r o f o xyg e n f ro m t h e gill s t o th e v ar i o us ti s s u e s o f c a rb o n
, ,

dioxide to th e gill s a nd of u re a to th e e xcr e to r y o rg a ns If a

, .

c rayfi sh is w o unde d th e bl oo d o n c omi ng in c o nt a ct with th e ai r

, , ,

thi cke ns fo r ming a cl o t It i s said to coagu la te Thi s cl o g s th e

,
. .

o p e ning a nd p r e v e nt s l o ss o f bl o o d T h e ch e lip e d s a nd o th e r .

w alki ng l e g s o f th e cr a yfi sh h ave a b re a king p oint n e a r the i r

b a se s Wh e n o ne i s injur e d th e a nim al m ay b re a k the limb a t
.

thi s p o int a nd l e sse n th e bl oo d flo w sin ce o nly a sm all sp ace is ,

p r e se nt in th e a pp e nd a g e a t this p a r ticul a r Sp o t a nd c o a gul a ti o n , ,

th e r e fo re t a ke s pl a ce v e ry qui ckly
,
.

B LOOD VE S S E L S —
T h e p rin cip a l bl o o d v e s se ls a r e a h e ar t
.
,

s e ve n m ain a r t e r i e s a nd a numb e r o f sp a ce s c all e d sinu se s

,
.

HE AR T T h e he a r t ( Fi g 1 0 1 2 9) is a mu s cul a r w all e d s a ddl e

. .
,
-
,

sh a p e d sa c lying in th e pe r i ca r di al si nu s in t h e m e di a n d o r s a l p a r t

o f th e th ora x It m ay b e co n sid e r e d a s a dil a t a ti o n o f a do r s al

.

v e sse l r e se mbling th a t o f th e e a r thw or m Si x e l a s tic li ga me nts .

,

t w o a nt e r i o r t w o p o s t e r i or a nd t w o r unning a l o ng th e v e nt r a l
, ,

b or de r of e ach l a t e r a l s u r fa ce fa s t e n it to th e w all s o f th e p e ri
,

c a r di a l sinus Th r e e p ai r s o f v a lvul a r a p e r tur e s c all e d osti a

.
, ,

o ne d o r sa l a nd t w o l a t e r a l a ll o w t h e bl oo d t o e nt e r f ro m t h e
,

s u rr o un di ng s inus

AR TE RI E S F iv e a r t e r i e s a r is e f r o m th e a nt e ri or e nd of th e
.

h e a r t ( Fi g .

( I ) T h e ophthalmi c a r t e r y 3 4) i s a m e di a n d o r s a l tub e p a ssing

f o rw a rd o ve r th e s to m a ch a nd s upplying th e c a r di a c p or ti on, ,

th e oe soph a gu s a nd h e a d , .

( 2 3 ) T h e t w o a nte nna ry a r t e ri e s (3 5 ) a ri se o ne o n e a ch sid e

,

o f th e o phth a l m i c a r t e r y p as s f o r w a r d s o utw a r d s ,
a nd d o wn , ,

w ar d s and b r a n ch s e ndi ng a g a st r ic ar t e r y t o th e c a r di a c p ar t
, ,

o f th e s t o m ach a r t e r i e s t o th e a nt e nn a
, t o th e e x cr e t or y o r g a n s , ,

a nd to t h e mu scl e s a nd o th e r ce ph a li c ti s s u e s .

(4 5) T h e tw o he pati c a r t e r i e s (3 6) l e av e the h e ar t b e l o w
,
 Y I
T HE C R A F SH AN D ART HROP ODS I N GEN ER AL 20
3

th e a nt e nn a ry a r te ri e s Th e y l e a d di re ctly to th e di g e s tiv e
.

gl ands .

A singl e d or s a l a bd o min al a r t e r y a r i se s f ro m th e p o s t e r i o r
e nd o f th e h e a r t .

( 6 ) T h e d o r s a l a bd o mi na l a r te r (
y 3 1 ) i s a m e di a n tub e l e a ding
b a ckwa r d s fr o m th e v e nt ra l p a r t of th e he a r t a nd s upplying th e ,

d orsa l r e gio n o f th e a bd o m e n It b ra n ch e s n e a r i ts p o int o f

.

o r igin giving r i s e t o t h e s te r nal a r t e r y

,
this l e a d s dir e ctly
d o wnw ard a nd p assing b e twe e n th e n e r ve c o r d s c o nn e cting th e
, ,

f o u r th a nd fifth p a i r s o f th or a ci c g a ngli a ( se e p 20 5 ) di vide s int o .

two a rte rie s One o f th e se th e ve ntr a l thor a ci c a r t e ry r un s f or

.
, ,

w ar d b e n e a th th e n e rve ch a in a nd se nds b ra nche s to th e v e nt r al

,

tho r a ci c re gi o n a nd to app e nd age s III t o XIII ; th e o th e r

, ,

t h e ve ntr a l a bd omi na l a r te r y r un s b a ckw a r d b e n e a th th e

n e rv e ch ain a nd se nds b r a nch e s to th e v e nt r al abdo min al re gi o n

,

a nd t o t h e a bd o min a l a pp e nd a g e s .

SI N U SE S .T h e bl oo d p a sse s f r o m th e a r t e r i e s int o Sp a c e s lying

in t he mids t o f th e ti ss u e s c all e d sinu s e s T h e p e ric a r di a l sinu s
,
.

h a s a l r e a dy b e e n m e nti o n e d T h e th o r a x c o nt a ins a l a r g e v e nt r a l
.

bl oo d sp a c e th e s te r na l s i nu s a nd a numb e r o f br a nchi o ca rdi a c

, ,
-

ca na ls e xt e nding f ro m t h e b a s e s o f t h e gill s t o t h e p e r ic a r di a l

Sinu s . A pe ri vi sce r a l s i nu s s u r ro und s t h e a lim e nt ary c a n a l in

the ce pha lothor ax .

B LOOD F LOW . h e a r t by m e an s o f r hythmic al co nt ra e

T he
-
,

ti o n s fo r ce s th e bl oo d th ro ugh th e a r t e ri e s to a ll p a r ts of th e b o dy
, .

Va lve s a r e p r e se nt in e v e r y a r t e r y wh e r e it l e a v e s t h e h e a r t ;
th e y p r e v e nt th e bl oo d f r o m fl o wing b a ck T h e fin e s t b r a nch e s.

o f th e s e a r t e r i e s th e ca pi lla r i e s o p e n int o sp a c e s b e tw e e n t h e
, ,

tissu e s and th e bl oo d e v e ntu ally re a ch e s th e ste rn a l sinu s

, .

F ro m h e re it p asse s int o t h e efle r e nt cha nne ls o f t h e gill s a nd int o

th e gill fil a m e nt s wh e re th e c a r b o nic a cid in so luti o n i s e xch a ng e d
,

f or o xyg e n f r o m th e w a t e r in th e b ra n chi a l ch a mb e r s It the n .

r e tu r n s by w a f h e a e r e nt i ll cha nne ls p a s s e s int o th e b r a n

y o t fi g ,

chi o ca r di a c s inu s e s th e nc e t o th e p e r ic a r di a l s inu s a nd fin a lly

-
, ,

th r o ugh t h e o s ti a int o th e h e a r t T h e v a lv e s o f th e o s ti a a ll o w
.
2 04 AN INTRODU CTION TO Z OOLOGY

th e bl oo d to e nte r th e h e a r t but p re v e nt it fr o m fl owing b a ck int o

,

th e p e r ic a r di a l s inu s .

R e s pi r a tor y S ys te m B e tw e e n th e b r a nchi o st e gi t e s a nd th e
.

b o dy w all a r e th e b r anchi a l ch a mb e r s co nt aining th e gi ll s At .

t h e a nt e r i or e nd o f th e b r a n chi a l ch a mb e r i s a ch a nn e l in whi ch th e
s ca phogna thi te o f th e se co nd m a xill a ( Fi g 1 0 2 A e p ) m o v e s b a ck .
, ,
.

a nd f o r th f o r cing t h e w a t e r o u t th r o ugh th e a nt e r i or o p e ning

, .

Wa t e r fl o ws in th ro ugh the p oste ri or op e ni ng o f th e b ra n chi al

ch a mb e r .

GI LL S . Th e r e a re two r ows o f gills n a m e d a cco r ding to th e i r ,

p o int s o f a tt a chme nt T h e o ut e r m os t th e podobr a nchi a a r e

.
, ,

f a s t e n e d to th e coxopod i te s o f c e r t a in app e nda g e s ( se e Ta bl e

VIII ) a nd t h e inn e r d o ubl e ro w t h e a r thr obr a nchi ce a r i se f ro m
, ,

t h e m e mb ra n e s a t th e b a s e s o f th e se a pp e nd a g e s In A s ta cu s .

th e re is a thi r d r ow th e ple u r obr anchi ce a tt a ch e d t o th e w a ll s o f

, ,

t h e th or a x .T h e numb e r a nd a r r a ng e m e nt o f th e se gill s a r e
sh o wn in Tabl e VIII T h e p o d ob ra nchi a co nsist o f a

T AB LE V III
THE NU M BE R AND P OSI TI ON OF T HE G I LL S OF T HE C RAYFI SH

( C a mba r u s )

SE GME NT ART H R OB R AN C HU E

6 ( p)
e . 1 7 (e ? )

b a s al pl a t e c ove r e d with d e lic a t e s e t a and a c e nt ra l a xi s b e a ring

a thin l o ngitudi n a lly f o ld e d co rr ug a t e d pl a t e o n i t s di st a l e nd
, ,

a nd a f e a th e r -lik e g ro up o f br o nchi a l fi la me nts T h e a r th r ob r a n .

chi a h a ve a c e nt r al s t e m o n e ith e r Sid e o f whi ch e xt e nd s a numb e r

o f fil a m e nt s c a u sing t h e e nti r e s t r u ctu r e t o r e se mbl e a plum e
,
.
2 06 AN IN T R ODU C TION TO ZOOLOGY

at th e e nd m o v abl e stalk s whi ch e xt e nd o u t o ne fro m e ach side

of ,

of th e r o st r um ( Fi g 1 0 1 .
,
T h e e xt e r n a l co nv e x su r f a ce o f
th e e ye i s co v e r e d by a m o difi e d
p or ti o n of th e tra n sp a r e nt cuti cl e ,

c all e d th e cor ne a Thi s co r n e a is .

di vide d by a l a rg e numb e r of fi ne
lin e s into fo u r Side d a r e a s t e r m e d -
,

fa ce ts E a ch f a ce t i s but th e e x
.

t e r n a l p a r t o f a l o ng vi s u al rod
kn o wn a s a n omm a ti di u m ( Fi g .

Se cti o n s sh o w th e co mp o und e ye
t o b e m a d e up o f Simil a r o mm a
ti di a lying s id e by Sid e but s e p a ,

r a t e d f ro m o ne a n o th e r by a l a y e r

o f d a r k pigm e nt ce ll s T h e a ve ra g e .

numb e r o f o mm a tidi a in a singl e e ye

is 2
5 0
0

T wo mm a tidi a a r e sh own I n Fi g
o

u re 1 03 B e ginning a t th e o ut e r
.

su r fa c e e ach o mm a ti di um co n si st s of
,

th e f o ll o wing p a r ts : ( a ) a cor ne al
fa ce t ( b) tw o cor ne age n c e ll s
(2 ) whi ch se cr e t e th e c or n e a (c) a ,

crys ta lli ne cone 5 ) fo r m e d by f o u r

co n e ce ll s o r vi tr e llee
, (d ) t w o
r e ti nu la r ce lls s u rr o undi ng th e con e
(no t sho wn in Fi g ()
6 s e
. v e n
FI G . 1 03 . Lo ngit u d i na l
tio ns o f t wo o m
se c

m a t i d i a o f th e cra y fi s h ; A p i gm e nt ,

a r ra nge d a s i nfl u e nce d by li ght ; B ,

p i gm e nt a r r a nge d as i nfl ue nce d by
d a r k ne ss ; 1 co r ne a ; 2 nu cl e u s o f
, ,

co r ne a ge n ce ll s ; 3 nu cl e u s o f vit r e ll a ;
,

4 nu cl e u s o f p i gm e nt ce ll ; 5 cr ys t a lli ne co ne ; 6 t a p e t u m ce ll ;
, , ,

7 ,
r h a b d o m ; 8 r e,
ti na l ce ll ; 9 b a se
,
m e nt m e m b r a ne ; 1 0 r e ti n al ,

ne r ve fi be r ( Fr om Se d gwi ck a f t e r P a r k e r )
. .
 Y I
T HE C R A F SH AN D AR T HR OP OD S I N G EN E R AL

re ti nul a r c e ll s (8) which f or m a r ha bdom ( 7 ) c o n si s ting o f fo u r

r ha bd om e r e s a nd ( f ) a numb e r o f pi gm e nt ce lls (4
,
F ib e r s ,

f ro m t h e opti c ne rve e nt e r a t the b a se of th e o mm a tidium a nd

c ommunica te with th e inn e r e nds o f th e r e tinul ar c e ll s
VI SI ON .
— T h e e ye s o f th e c r a yfi sh a r e supp o s e d t o p r o duce
a n e r e ct m osa i c o r a ppos i ti on i m age thi s i s illu s t r a te d in Fi g
u r e 1 04 wh e re th e o mm a tidi a a re r e p r e se nt e d by a e a nd th e fib e r s
—

f ro m th e o ptic n e rv e by a e ’— '
T h e r a y s o f light f ro m a ny p oint
a b o r c will all e n co unt e r th e d a r k pigm e nt c e ll s s u r ro u n di ng th e
, , ,

o mm a ti di a a nd b e a b s o rb e d e xc e pt th e r a y whi ch p a s se s di r e c tly
,

thro ugh th e c e nte r o f th e co rne a a s d o r e ; thi s r ay will p e n e t r a t e

t o th e r e tinul a a nd th e nc e t o th e fib e r s f r o m th e o ptic n e r v e
“
.
,

Thu s th e r e tinula of o ne o mm a tidi um r e c e iv e s a singl e re s ulta nt

imp r e ssi o n f ro m th e light whi ch r e a ch e s it B u t th e a dj a c e nt .

o mm a tidi a b e ing di r e ct e d t o a di ff e re nt th o ugh a dj o ining re gi o n

, ,

o f t h e o ut e r w o rld m a y t r a nsmit a di ff e r e nt i mp r e ss i o n a nd t h e
, ,

stimuli f r o m a ll th e o mm a
f idi a whi ch m a k e up a co m
p o und e ye will co rre sp o nd in
g re a te r o r l e ss d e g r e e to th e
wh o l e o f t h e vi s ibl e o ut e r

w o rld whi ch s ubt e nds the i r

s e ve r a l opti c a x e s Th e su m
.

o f th e re s ulting im a g e s whi ch

w e m ay thu s supp ose t o b e F G I .

4 Di g m t
10 . a pl in m
ra io ex a o sa c

t r a n smitt e d to th e b r a in m ay vi i n ( s o t t ) ( F m P k d
se e ex . ro ac ar

ft aL bb k )er u oc
b e co mp a r e d t o a m o sa i c in
.

whi ch th e e ff e ct i s giv e n by a l a r g e numb e r o f se p a r a t e pi e c e s ,

”
o f o ne s iz e a nd e a ch o f u ni f o r m co l o u r ( 3
1 8 p Thi ,
s .

m e tho d o f im a ge f o r m a ti o n i s e sp e ci ally w e ll a d apt e d f o r r e cor d

i ng m o ti o n Sinc e a ny ch a ng e in th e p o s iti o n o f a l a r g e o bj e ct
,

a ff e ct s th e e nti r e 2 5 00 o mm a tidi a .

Wh e n th e pigm e nt s u r r o und s th e o mm a tidi a ( Fi g 1 0 3 A) .

, ,

vi si o n i s a s de scr ib e d ab o ve ; but it h a s b e e n fo und tha t in dim

light th e pigm e nt mig r a t e s p a r tly t o w a r d th e o ut e r a nd p a r tly
2 08 AN
'

INTR ODU CTION TO Z OOLOGY

t ow a r d the b a s a l e nd of th e o mm a ti di a ( F i g 1 03 B ) Wh e n this .
,
.

o c cu rs th e o mm a tidi a no l o ng e r act se p a ra t e ly but a co mbin e d

im ag e i s thr o wn on th e r e tinula r l a y e r “ In thi s m a nn e r a n .

,

e r e ct su pe r pos i ti on i m a ge is f or m e d th e r ays r e fr a ct e d by a l arge

‘
,

numb e r of c r y s t a llin e co n e s b e ing s up e rp ose d a t th e f o cus o n th e

r e tin a a nd a stimulus f a r st ro ng e r in p r op o r ti o n t o t h e int e n s ity
,

o f th e illumin a ti o n th a n th a t o f th e a pp o s iti o n im a g e tho ugh ,

p r ob ably much l e ss di stinct in d e t ail i s giv e n t o th e r e tin u l a,

( 1 83 , p
STATOC Y ST S .t t yst
-
The s a oc s of chitin ous w a ll e d
C a mba r u s a r e -

s a cs situ a te d o ne in th e b a sal se gm e nt of e a ch a nt e nnul e In th e .

b ase o f th e s t a t o cy s t i s a r idg e call e d th e s e ns ory cu s hi on a nd

, ,

th re e s e ts o f ha i rs o v e r t w o hund re d in all e a ch inn e rv a t e d by a

, ,

s ingl e n e r v e fib e r Am o ng th e se h a i r s a r e a numb e r o f l a rg e
.

g ra in s o f sa nd th e s ta toli ths whi ch a r e pl a ce d th e r e by th e cr a y

, ,

fi sh . B e n e a th th e se n s o r y cu shi o n a r e gla nds whi ch se cr e t e a su b

s t a nc e f or th e a tt a chm e nt o f th e s t a t o lith s t o t h e h a i r s

T h e s t a t o cy s t f or m a ny y e ar s w a s c o n sid e re d a n a udit o r y
o rg an a nd it m a y p oss ibly fun c ti o n as s u ch
,
th o ugh re c e nt ,

inv e s tig a ti ons h a ve p ro ve n tha t it i s p r im a r ily a n o rg a n o f

e q u i li br a ti on T h e co nt a ct o f th e st a t o lith s with th e s t a t ocy s t
.

h ai rs d e t e rmi n e s th e o r i e nt a ti o n of th e b o dy whil e swimming ,

since any cha nge in th e p o siti o n o f th e a nim al c a u se s a ch ang e

in th e p ositi o n of th e s t a t o lith s und e r th e influ e n ce o f g ra vity .

Wh e n th e c r a yfi sh ch ang e s its e xo s k e l e to n in th e p r o ce ss of m olt

ing the s t a t ocy s t i s a l so sh e d Individu a l s th a t h a v e just m olt e d
,
.
,

o r h a v e h a d th e i r s t a t ocy s t s re m o v e d l ose much o f th e i r p o w e r s

,

o f o r i e nt a ti o n P e r h a p s t h e m os t c o nvin cing p r oo f o f th e fun c

.

ti o n o f e quilib ra ti o n i s th a t fu r ni sh e d by th e e xp e r im e nt s o f
Kr e i dl This inve s tiga t or pl a c e d shr imp s which h ad ju s t ,

m o lt e d a nd w e re the r e fo r e with o ut st a t olith s in filt e re d w a t e r ,

.

Wh e n s uppli e d with i ro n filings th e ani m al s fill e d th e ir st a t o cys t s

,

with the m A str o ng e l e ctro m a gn e t w as th e n h e ld n e ar th e

.
‘
-

s t a t o cy s t a nd t h e Sh rimp t oo k up a p os iti o n c o rr e sp o n di ng t o th e
,

re s ul t a nt of the t w o pull s th a t o f gr a vity a nd of the m a gn e t

,
.
21 0 AN IN TRODU CTION TO Z OOLOGY

f e mal e r e p r o ductiv e o r g a ns
w e r e f ound in a singl e sp e ci
men .

M ALE R E P R ODU C TI V E OR
G AN S — Th e m al e or g a n s ( Fi g .

1 06) c o n si s t o f a te s ti s a nd two
vasa d ef e r e nti a 3) whi ch o p e n
th r o ugh th e co x0po di t e s o f th e
fifth p a i r o f w alking l e g s (4 ,

T h e t e s ti s li e s ju s t b e
n e a th th e p e ri ca r di a l sinu s It .

i s a so ft whit e b o dy p osse ssing

t w o a nt e r i o r l o b e s ( F i g 1 0 6 1 ) .
,

a nd a m e di a n p os t e ri o r e xt e n

si o n T h e va s a d e f e r e nti a
a r e l o ng co il e d tub e s . M a le tiv e o rga ns o f
r e pr o d u c

th e cr a y fi s h .1 r i ght
,
a nt e r i or

l ob e o f t e s ti s 2 po s t e r i or l o b e o f
,

t e s ti s 3 v a s d e f e re ns 4 e xt e r na l
, ,

Op e ni ng o f v a s d e f e re ns ; 6 b a se ,

o f fi f th w a l k i ng le g ( Fr o m Shi p
.

le y a nd M a cB r i d e ) .

SP E R M ATOGE N E S I S .

p rimitive g e r m c e ll s T he
within th e t e s tis p ass
th ro ugh t wo m a tu ra ti o n
divi si o n s a nd th e n m e t ,

a m o rph o s e into sp e rm a
toz o a Th e se a r e fla t .

t e n e d sph e r oid al b o di e s
wh e n in cl o se d Wi thi nt h e
FI G 1 07 sp mert n f t
a oz ooh y fi h o e cr a s
t e s ti s o r v a s d e f e r e n s but
. . .

( Fr o m And w i n A n t A n )
re s, a . z .
,

if e x a mi n e d in w a t e r or
som e o th e r liquid th e y a re s e e n to uncoil fin ally b e co ming s t ar ,

sh ap e d ( F i g .
 T HE C R A Y FI SH AN D AR T HR OP OD S I N G E N ER AL

Th e sp e r m a to z oa re m ain in th e t e s ti s a nd v a sa d e f e r e nti a
until c o pul a ti o n t a ke s pl a c e As m a ny a s t w o milli o n sp e r m a
.

to z oa a r e c o nt a in e d in th e v a s a d e f e r e nti a o f a singl e sp e cim e n

F E M ALE R E P R ODU C TI V E OR GAN S — The ova ry re se mbl e s

th e t e s tis in f o r m a nd i s simil a rly l o c a te d in th e
,
b o dy
( g
F i . A s h o r t ovi d u ct ( 1 ) l e a d s f ro m n e a r t h e

c e nt e r o f e a ch Side o f th e o va r y to th e e xt e r n al ap e r tu r e in th e
c o xop o di te o f th e thi r d w a lking le g (4 ,

OOG E N E S I S T h e p r imitiv e g e r m c e ll s in th e w a ll s o f th e o v a r y
.
-

g ro w in Size b e co m e s u r
,

r o und e d by a l a y e r o f
sm all c e lls th e follicl e
, ,

which e ve ntu ally b r e a k

d o wn a ll o wing th e e gg s to
,

e scap e into the c e nt r a l

c a vity o f th e o va r y At .

th e tim e o f l a ying th e o v a
p a ss o u t th ro ugh th e o vi
duct .

B r e e d in g H a b i ts Th e .

d e t ail s o f c opul a ti o n e gg ,

l aying and th e l a r v a l
,

s t a g e s o f C a m ba r u s h a v e

o nly r e c e ntly b e e n m a d e

ou t ,
a nd e ve n no w o u r
a c co unt mu s t b e d e riv e d
F IG 8 F m .l 10p d . tiv e g n a e re ro uc e or a s

Of t h yfi h i ghte vi d ti
cr a S I r O uc
fro m ob se rva tions of i ght l b f v y ; 3 l f t l b
2, r
'
,

o e
o e o o ar e
s e ve ral di ffe r e nt Sp e ci e s p n d t, h w nt l vity ; 4
o e e o S o ce ra ca
,

,
ex

sinc e th e e nti r e lif e h i s t n l p ni ng f vi d t ; 5 b

er a o e f o o uc a se o
,

t o ry Of a singl e Sp e ci e s thi d w l k i ng l g ( F m Sh ipl y nd

r a e . ro e a

h a s n e ve r b e e n re c o r d e d M B
.
id ) ac r e '

T h e d e v e l o pm e nt o f th e e gg s o f C a m ba r u s f ro m th e tim e o f
de p o siti o n t o th e tim e o f h a tchi ng h a s li k e wi se n e v e r b e e n
inve stig a te d .
21 2 AN IN T R ODU C TION TO ZOOLOGY

The p rin cip al e v e nt s in the re p ro ducti o n of cra yfi she s m ay b e

e num e r a t e d a s f o ll o w s

( )
I C o pu l a ti o n du r ing which t,
h e s p e r m a t o z o a a re t r a n s fe r r e d

fro m th e v a sa d e fe re n tia o f th e m a l e to th e s e min a l r e ce pt a cl e o f

th e f e m a l e ; ( 2) e gg l a ying ; ( 3 ) t h e e mb r yo ni c d e v e l opm e nt
-

o f th e e gg s ; (4 ) h a tch i ng ; 5) t h e g ro wth o f t h e y o ung cra y

fi sh e s
C OP U LATI ON C opul a ti o n in cr a yfi sh e s in m o s t ca s e s t a k e s
.
, ,

pl a c e in Se pt e mb e r Oct ob e r or N o ve mb e r o f the fi r s t ye ar of
, ,

FI G . 1 09 . M al e cr a y fi sh tr a nsf e rr i ng spe r m a toz oa to th e se m i na l t l

r e ce p a c e

o f th e f e m a l e .
( Fr o m Andr e w s i n Am N a t )
. .

th e i r live s th a t i s wh e n th e y ar e ab o ut fo u r m onths old A se c

, , .

o nd c opul a ting se a s o n i s p a s se d th r o ugh a t th e e nd o f t h e se co nd

summ e r whe n the a nim al s a r e ab o ut se v e nt e e n m o nth s old a nd a

, ,

thi r d c opul a ting se aso n o c cu r s a t th e e nd o f th e thi r d s umm e r

At th e se tim e s a m a l e a pp ro a ch e s a f e m a l e g ra sp s h e r ,

by h e r c e ph alic app e nd ag e s a nd a fte r a s tr uggl e tu rn s h e r o ve r

, , ,

o n h e r b a ck He th e n s t a nd s o v e r h e r in th e p o siti o n sh o wn in
.

F igu re 1 0 9 a nd t r a n s fe rs Sp e r m a t o z o a t o th e se m in a l r e c e pt a cl e
,
.

D u ring thi s p ro c e ss th e sp e r m a t o z o a flo w ou t o f th e o p e ni ng s o f
,

th e v a s a d e f e r e nti a p ass al o ng th e g ro o v e s o n t h e fi r s t a bd o min a l

,

a pp e nd a g e s o f th e m a l e ( Fi g 1 0 1 a nd e nt e r t h e se min a l r e
.
,

ce p t a cle He r e th e y are s t ore d du ring t h e wint e r Th e .

s e mi na l r e ce ptacle i s a c a vity in a f o ld o f cuti cl e lying b e tw e e n th e

21 4 AN IN T R ODU C TION TO Z OOLOGY

no d o ubt s e cre te d by th e c e m e nt gl ands Thi s is b r o ught a b o ut

.

by th e tu r ning o f th e a nim a l fi rs t o n o ne sid e a nd the n on the

o th e r a numb e r o f tim e s F ro m o ne hund re d to ov e r si x hund r e d
.

gre e ni sh e ggs ar e l aid by a singl e fe m al e d e p e nding up o n th e ,

s iz e a nd a ge o f th e a ni m a l Aft e r th e e ggs a r e l a id th e cr a yfi sh
.

r ight s h e r se lf th e a p r o n o f gl a i r b r e a ks d o wn a nd th e a bd o m e n
, ,

is e xt e nde d ( Fi g .

FI G . 1 1 1 . Fe m a le ti ng e ggs b y r a i s i ng a nd s t r a ight e ni ng a bd o m e n a nd
a e ra

w a vi ng s wi m m e r e t s ba ck a nd f o r th ( F ro m And r e w s i n A m N a t )
. . .

F E R TI LI Z ATI ON m e tho d o f fe r tiliz a ti o n h a s n e v e r b e e n

.
— The
dis co ve re d It i s supp o se d th a t a s th e e gg s a r e l aid th e y p a ss
.

o v e r th e o p e ning o f t h e s e min a l r e ce pt a cl e a nd a r e th e n p e n e
,

f r a te d by th e sp e r m a t o z o a whi ch w e re pl a c e d th e r e by th e m a l e

th e p r e c e ding a utumn
E m b ryo l o gy Whil e th e e gg s a r e d e v e l oping the y ar e pro
.

t e ct e d by th e a bd o m e n o f th e f e m al e a nd a r e a e r a te d a nd k e pt
,

fr e e fro m di r t by the w aving o f th e swimm e re ts b a ck a nd forth

( g
F i . T h e e mb r y o l o gy o f C a m ba r u s h a s n e v e r b e e n i nv e s

t i ga t e d but it p r ob a bly r e se mbl e s cl o se ly th e d e v e l o pm e nt o f th e

,

co mm o n E u r op e a n c ra yfi sh A s tacu s f lu vi a ti li s
,
.

T h e f e r tiliz e d e gg o f As ta cu s co n sis t s o f a l a rge numb e r o f

lk h r e s e mb e dd e d in c topla s m n d nt ins a nu cle u s
y o s p e y a c,
o a

co mp o se d o f th e e gg nu cl e u s a nd t h e sp e r m a t o z o o n c o mbin e d .

Thi s nu cl e us p roce e ds t o fo r m tw o d aught e r nucl e i T h e e gg .

doe s no t b e com e divi de d int o tw o p ar t s by ce ll w alls but th e ,

d a ughte r nu cl e i se p ara t e fro m one a n o the r and divide aga in

 Y FI SH
'

T HE C R A AN D AR T H R OP OD S I N GEN ER AL 21 5

( Fi g . 1 1 2, This divi sio n c o ntinu e s until the re a r e a l a rge

A) .

numb e r o f nucl e i s c a tt e re d a b o ut a mo ng th e yo lk sphe r e s Fi .

n a lly all o f th e nucl e i mig ra te t o w ar d th e p e r iphe r y o f th e e gg and

c o me to li e jus t b e ne a th th e s ur fa ce ( Fi g 1 1 2 B ) C e ll wall s .
,
.

th e n app e a r dividi ng the e gg int o as m a ny y olk pyr a mid s a s the r e

,

a r e nu cl e i ( F i g 1 1 2 C ) T h e o ut e r e nd s o f th e se yo lk pyr a mi d s
.
,
.

co nt a ining th e nu cl e i a r e no w cut o ff by cr oss w a ll s th e r e sul t b e ing ,

a singl e l a y e r o f ce ll s t h e bla s tod e r m s u r r o unding t h e e nti r e e gg

, , .

T h e Side wa ll s of t h e inn e r p o r ti o n o f th e pyr a mids fus e p a r ti ally ,

r e st or ing th e o ri gin a l c o ndi ti o n .

FI G . 1 1 2. T hr e e s t a ge s i n th e
v e lo p m e nt of th e e
gg of th e
cra y fi sh . A, a fe w c l e a v a ge
nu cl e i i n th e e gg ; B nu cl e i
,

a r r a nge d ne a r the s u r f a ce ;
C, gg d
e ivi d e d i nt o yo k l
p yr a m d s i .
( Fro m Ko r sch e lt
a nd He i d e r . A a nd t B, af e r

M o ri n C ,
a f t e r R e i ch e nb a ch ) .

The mb r yo l o gy o f th e cra y
e

fi sh i s s o c o mpl e x th a t a d e
t a il e d a c co u nt will no t b e
a tt e mpt e d h e re It wi ll suffi c e to m ak e a sup e r fi ci a l e xa min a
.

ti o n o f th e de v e l oping e mb ryo with th e a i d o f Figu re s 1 1 2— 1 1 5 .

A thick e ni ng o f th e bl a s to d e r m o n o ne Side o f th e e gg i s kn o wn
21 6 AN INTRODU CTION TO Z OOLOGY

as the pri m i ti ve s tr e ak . F iv e
the r thi ck e ning s al so a ri se in this
o

r e gi o n a p a ir o f opti c lobe s ( Fi g 1 1 3 K ) a p a ir o f thor aco-abd om i na l

, .
,

p la te s a n d a m e di a n e ntod e r mal
p la te ( E S T h e sid e

o n whi ch th e se thick e nings o ccur will b e co m e th e v e ntr a l sur f a c e

o f th e a dult .

A l a t e r st ag e ( Fi g 1 1 4 ) sho ws a m e di a n o utgr o wt h j u st b a ck
.

o f th e o ptic l o b e s which will b e c o m e th e upp e r lip o r labr u m l

() , ,

a nd p a i rs o f r u di m e nt a r y a nte nnu le s a nte nna ( a ) a nd ma n 2

di ble s (m) B e twe e n th e a nt e nn a is a d e p re ssio n whi ch will

.
,

b e co m e the m ou th An op e ning in the thora co—abdo min al pl ate

.

TA ) r e p r e se nt s th e futu re a nus ( A)
. In a m or e a dvance d .

e mb r yo ( Fi g 1 1 5) t h e st r uctu r e s a l r e a dy m e nti on e d a r e se e n in a
.

l a t e r s t age o f de ve l opm e nt T h e th ora c o-abdo minal rudim e nt

.

h as b e co m e divid e d int o a numb e r o f s e gm e nt s fr o m e a ch of ,

which a p a ir of app e ndage s a ri se and de ve l op into those o f th e

a dul t cra yfi sh T h e l e ngth o f tim e r e qui r e d f or th e v a ri o u s
stage s in d e ve l opme nt wa s fo und by And r e w s to be a s foll ows :
'

C l e a v a ge t oo k up th e fi r st w e e k th e b e ginning o f a n e mb ry o th e
,

s e cond w e e k to p r ogr e ss a s f ar a s th e N a uplius the thi rd we e k

,

a nd m o r e t o e nl ar g e th e e mb r y o o v e r o ne h alf o f th e e gg a fo u r th
,

w e e k and m ore and to p e r fe ct th e e mb ryo f o r h a t ching a fifth a nd

,

Sixth w e e k or m o r e T he wh ol e e gg d e ve l opm e nt r e qui r e d fr o m

.

fi v e to e ight we e ks in di ff e r e nt s e ts o f e gg s und e r diff e re nt t e mp e ra

ture ( 1 59 pp 1 8 9 ,
.

HATC HI N G ( 1 5 9 In h a tching th e e gg cap s ul e spli ts and

, ,

th e l a rv a e m e r g e s h e a d f o r e m os t T h e h e lpl e ss yo ung c r a yfi sh
.

w o uld dr op awa y fr o m th e m o the r a t o n ce but f or a thre a d e xt e nd

ing fro m its t e lson to the inn e r surf a ce of the e gg caps ul e (Fi g .

So o n th e l arva p o s se ss e s st re ngth e no ugh to gr asp the e g s t ring g

with its cl a w s ( Fi g T h e t e l so n th r e a d the n b r e a ks

. Aft e r .

a b o ut f o r ty e ight h o u r s t h e l ar v a p asse s int o a se cond s ta ge

- Thi s .

i s in a ugu r a t e d by th e sh e dding o f th e fi r s t l a r va l cuticul ar cove r

ing a p r oce s s know n as m olti ng o r e cdysi s Thi s cas ting o ff o f
,
.

th e c o v e r ing of th e b o dy i s no t p e culi ar to th e yo ung but o ccurs ,

 3 Su rf a ce o f
1 1 . afi sh e gg with e m b ryo
cr a y

i i
g nn ng to f o rm E S e nt o d e r m a l p l a t e ; K o p ti c
.
, ,

lo be s ; TA th o r a co a bd o m i na l p l a t e s
,
-
.
( Fr o m
Ko r s ch e lt a nd H e i d e r a f t e r R e i ch e nba ch ).

FI G . 1 1 4 . E m b ryo of h i n th e N a u pli u s s ta ge
t h e crayfi s .

A r u d i m e nt o f e ye ; a 1 a nt e nnu l e ; 0 2 a nt e nna ;
, , ,

G ce r e b r a l ga ngli o n ; m m a nd i b l e ; l u pp e r li p ;
, , ,

TA th o r a co a bd o m i na l pl a t e ; A a nu s
,
-
,
.
( F ro m
Ko r s ch e lt a nd H e i d e r a f t e r R e i ch e nba ch ).
FI G 1 1 5 Ol d e r e m b r yo of the cr a y fi sh e ye s ;
. .
.

o r, an et nnu le s ; o z, t
an e nna ; a b, a b d o m e n; l, up

p e r li p ; m , m a nd i bl e ; m x1 , mxg , fi r s t a nd s e co nd
m a xill a ; T , t e l so n ; t7 , ts , th o r a ci c li m b s t1 , l s ,
;
m a xilli pe d e s ; ti , tg, w a l k i ng l e s
g ( Fr o m Ko r s ch e lt
.

a nd He i d e r t
af e r Re i ch e nba ch ).
21 8 AN INTR ODU C TION TO ZOOLOGY

T AB LE I X

THE D UR ATI ON OF THE LAR V AL STAG E S , AN D TH E SI Z E AND H A B TS

I OF Y OU N G
C R AYF I S HE S DU R I N G TH E F I R ST S U M M E R

S TA G E D U R AT I ON H AB I T

Att a ch e d to m o th e r .

Att a ch e d to m o th e r .

As so ci a t e d with
m o th e r f or o ne

w e e k th e n f re e
,
.

Fre e .

Fr e e .

F re e .

F re e .

F re e .

p e nd e nt o f the m o th e r altho ugh th e y a t fi rs t a lwa ys r e tu rn t o h e r

,

whe n s e p ar a te d F r o m thi s time o n the l a r va sh ift f o r the mse lve s

.
,

T AB LE X
V
TH E A E R AG E R ATE OF GR OWTH OF A C R AYF I SH ( Ca mba r us afi i ni s )

TI M E IN T R VA L
E SI ZE

Jus t h a t ch e d
End of I st su m m e r

End of 2d su m m e r

E nd of 3d su m m e r

End of 4 th s u mm e r

g ro wi ng f api dly and m olting a t l e as t fo ur m ore tim e s duri ng

,

th e fi r s t s umm e r T h e fi r s t wint e r no g r o wth nor m olt s o ccur

. .

The re ar e fo u r o r fi ve m olts the s e co nd sum me r thr e e or fo ur in ,

the thi r d s u m m e r and pe rh aps o ne or two in the fo ur th summ e r

,
.
 T HE C RA Y FI SH AN D AR T HR OP OD S I N GE N ER AL 21 9

The life of a singl e indi vidu al e xt e nds o ve r a p e ri o d o f a b o ut th re e

o r mo re ye ars .Ta bl e IX p r e se nts th e m ain fa ct s o f l arva l d e
v e lopm e nt du r ing th e fi r st summ e r T abl e X give s the
a ge a nd siz e o f a singl e sp e cim e n du ring i ts e nti r e lif e .

R e ge n e r a ti o n — T h e c r a yfi s h a nd m a ny o th e r C r ust a c e a ns
.

h ave the p owe r o f r e g e n e r ating l os t p a r t s but to a much m ore ,

limite d e xte nt th a n s uch a ni m als as Hydr a a nd the e a r thwo rm .

E xp e rim e nts h av e b e e n p e r fo r m e d up o n a lm os t e v e r y o ne o f th e
a pp e nda g e s a s w e ll a s th e e ye T h e se co nd a nd thi r d m a xillip e ds
.
,

th e w a lking l e gs th e s wimm e r e t s a nd th e e ye h a v e a ll b e e n
, ,

inju re d o r e xti rp a t e d a t v a rio us time s a nd s ub se que ntly re n e we d

t h e l o s t p a r ts. M a ny sp e ci e s o f c r a yfi sh o f v a ri o u s a ge s h av e b e e n
u se d f or the se e xp e r im e nts Th e g ro wth o f re ge n e ra t e d ti ss u e
.

i s m ore f r e qu e nt a nd r a pid in yo ung sp e cim e ns th a n in a dults

( 1 8 4)
Th e ne w t uctu r e is no t a lwa ys like th a t o f the o ne re m o ve d
s r .

F or e x a mpl e wh e n th e a nnulu s co nt a ining t h e sp e r m re c e pt acl e

,

o f a n a dult i s e xti rp a t e d a n o th e r i s r e g e n e r a t e d but a lth o ugh thi s

, , ,

i s a s l a rg e a s th a t o f th e a dul t it is c o mp ara bl e in c o mpl e xity t o

,

th a t of a n e arly l a rval
st ag e A m or e
re m a r k a bl e ph e no m e

no n is th e r e g e n e ra ti o n
o f a n app a r e ntly func

ti o nal ( t actil e ) an
t e nn a -lik e org an in
pl a ce of a de ge n e ra te
e ye which w a s re m o v e d

f ro m the b lind cra yfi sh ,

C a m ba r u s p e llu cidu s

te s ti i In thi s case a
.

no n-functi o n a l orga n F G 7 D i
I g
. m1 1h wi ng .
nt nn li ka g
ran s o a e a- e or a

w as r e pl ace d by a fun c g n t d i n
re p l e f n y
e ra ef P l a ce o a e e o a a

m n ( F m M g n ft H b t) ro or a a er er s
ti o nal o ne Of a di ff e r
o .
.

e nt Char act e r T h e r e ge n e r a ti o n o f a ne w pa r t whi ch di ff e r s

2 20 AN IN TR ODU CTION TO Z OOLOGY

fro m th e p ar t r e m ov e d i s t e rm e d he te r omorphosi s Figu re 1 1 7 .

sh o w s a n a nt e nn a which r e g e n e r a t e d in pl a c e o f a n e ye in a m a rin e

c ru st a c e a n P a la mon In st a nce s o f he te ro mo rphosi s have a lso

,
.

b e e n re co r de d in e xp e rim e nt s on o the r a nim al s

Au toto my . P e r h ap s th e m os t int e r e s ting m o rph ol ogic a l
s t r ucture c o nn e ct e d with th e r e g e n e ra tiv e p ro c e ss in C a mba r u s

i s th e d e fini t e b r e a king p oint n e a r th e b a se s o f th e w a l k ing l e g s .

If the che l a a re injur e d the y a r e b ro ke n off by th e cr a yfi sh a t the

,

b re a king p oint T h e o the r w a lki ng l e gs if injur e d m ay be th ro wn

.
, ,

o ff a t th e f re e j o int b e tw e e n th e se co nd a nd thi r d se gm e nt s A .

ne w le g a s l arg e a s th e o ne l os t d e v e l op s fr o m the e nd o f th e
, ,

s tump re m a ini ng Th is b r e a king o ff o f th e l e gs a t a de fi ni t e p oint

.

i s kn o wn as a ut o t o my a ph e n o m e n o n th a t a l so o c cu r s in a numb e r
,

o f o th e r a n i m al s T h e le g i s s e p a r a t e d a l o ng th e b re a king p o int
.

by s e v e ra l s ucce ssive muscular co ntra cti ons It has b e e n shown .

tha t a uto to my i s no t d u e to a w e a kn e ss a t th e b re a king p oint ,

but to a r e fle x a ctio n a nd tha t it m a y be b ro ught ab o ut by a

,

stimul a ti o n o f the tho ra ci c g angli o n as w e ll a s by a stimul a ti o n

o f th e n e r v e o f th e le g it s e lf (1 82 p Imme di a te ly a ft e r
,
.

th e le g h a s b e e n thr o wn o ff a m e mb r a n e o f e ct o d e r m c e ll s co v e r s
,

th e c a n a l th ro ugh which th e n e r v e a nd bl o o d p a s se d ; fi v e d ays

l a te r re ge n e ra tio n b e gins by a n o utwa r d g ro wth o f the e cto de rm
ce lls whi ch lin e d the e x oske l e to n An i nte r e sti ng p oint i n thi s
.

ne w g ro wth i s th a t th e mu scl e s o f t h e r e g e n e ra t e d p a r t a r e p ro b
a bly p ro duc e d by e ct o d e r m c e ll s wh e r e as in th e e mb ry o ni c d e
,

v e lopm e nt o f th e c r a yfi s h th e mus cl e s a r e suppo s e d to a r i se f r o m

the e nt o d e r m
Th e p o w e r o f a ut o t o my i s o f a dv a nt a ge to th e c ra yfi sh Sin ce ,

th e w o und cl ose s m ore qui ckly if th e le g i s l os t a t th e b r e a king

p o int N 0 one h a s ye t o ffe r e d a n a de qua te the o ry to a cco unt
.

f or a ut o t o my . It i s p rob ably a p ro ce s s th a t th e anim al h as

a cqui r e d in c o nn e cti o n w ith th e c o n di ti o n unde r which it liv e s ,

o r in o th e r w or ds a n a d aptiv e re sp o n se o f th e o r g a ni s m t o its
,

c o nditi o n o f life ( 1 7 6 p ,
.

As in the e arthw or m th e r ate of r e ge ne r ati on d e pe nds upon th e

,
222 AN INT RODU C TION TO Z OOLOGY

to turn th e b o dy o ve r Fr o m a l a rg e numb e r o f e xp e rime nts i t

.

h as b e e n p r o v e n th a t th e s t a t ocy s t s a r e th e o r g a n s o f e qui lib r a

tio n Th e str uctur e o f the se o rg an s i s de s cr ib e d o n p ag e 208
. .

T h e c o nt a ct o f th e s t a t olith s with th e st a t o cys t h a i r s fu r ni sh e s

t h e stimulus which c a us e s th e a ni m a l t o m aint ain an up right p o si
tio n .

Wh e n pl a ce d on its b a ck th e crayfish has so m e diffi culty in

,

r i ghting it se lf . T w o m e th o d s o f r e g aining i t s n o r m al p o siti o n

a r e e mpl o y e d . T h e u su a l m e th o d is th a t o f r a i sing i t se lf o n one
side a nd all o wing th e b o dy t o tip o ve r by th e for ce of g ra vity .

T h e s e co nd m e th o d is th a t o f co nt r a cting th e fle xo r abd omin al

mu scl e s which c a u se s a qui ck b ackw a r d fl0p b r inging th e b ody ,

r ight s id e up In g e ne ral th e ani m al s right the m se lv e s

, ,

wh e n pl a ce d on the i r b a cks by th e e a sie s t m e tho d ; and thi s i s

,

f ound to de p e nd u sua lly up on th e r e l a tive w e ight of the two side s

o f t h e b o dy . Wh e n pl ace d up o n a su r f ace whi ch i s no t l e v e l th e y ,

t a ke a dva nt ag e a ft e r a f e w e xp e rie nc e s o f th e in clin a ti on by tu rn

, ,

ing t o war d the l o we r Side ( 1 8 8 p ,

.

SE N SE S AN D TH E I R L OC ATI ON . T h e s e n s e o f touch in cr ayfi sh e s

is p e rh aps th e m os t v alu abl e since it a ids the m in fin d ing fo od
, ,

a v o iding o b s t a cl e s ,
a nd in m a ny o th e r w a ys It i s l o ca t e d in
.

sp e ci a liz e d h a i r s o n v a r i o u s p a r t s o f t h e b o dy Vi s i on in
cra yfi sh e s i s p r ob ably o f mi n o r imp o r t a nc e since th e co mp o und
,

e y e s a r e a lm o st u se l e s s in re co gnizing f or m a nd a re o f r e a l v a lu e
,

o nly in d e t e cting m o ving o bj e ct s N o r e a cti o n s t o s ou nd h av e

.

e v e r b e e n o bs e rv e d in cr a yfi sh e s a nd a pp a r e ntly th e y d o no t

he ar .
“ ,

Th e r e a cti o n s for m e r ly a tt r ibut e d to a u di ti o n a r e p rob

”
a bly d u e t o t a ctil e re fl e x e s ( 1 8 1 p ,
. In a qu a ti c a nim a l s

it is so di ffi cult t o di stinguish b e tw e e n r e acti on s o f taste and s me ll

th a t the se se n se s ar e b o th include d in th e t e r m che mi cal se nse .

T h e e nd o rg a n s o f thi s se ns e a r e di s t r ibut e d a ll o v e r th e b o dy
“
.

R E AC TI O N S To STI M U L I —
.
—
TH I GM OTR OP I S M Th e cr a yfi sh i s .

s e n siti v e t o t o uch o v e r th e who l e su r fa ce o f th e b o dy but e sp e ,

ci ally o n t h e ch e l a a nd ch e li pe d e s t h e m o uth p a r t s t h e v e nt ra l
, ,

surface o f th e abdom e n a nd the e dge o f th e t e l so n ( 1 64 p

, ,
.
 Y I
TH E C R A F SH AN D ART HR OP ODS I N G EN E R AL 2 23

Th e nt e nn a a re us ually co n sid e re d th e sp e ci a l o rg a ns o f to uch

a ,

but e xp e r ime nt s se e m to p ro ve tha t th e y a re no t so se n sitive a s

o th e r p a r t s o f t h e b o dy T h e t a ctil e h a i r s a re plum e d a nd su p
.
,

plie d by a singl e n e rve P osi ti ve thi gmotr opi s m i s e xhibit e d by

.

cr a yfi sh e s to a m a r k e d d e g r e e th e a nim a ls se e king to pl a ce th e i r
,

b o di e s in co nt act with a solid o bj e ct if p o ssibl e T he n o r m al ,

.

p o siti on o f th e crayfi s h wh e n a t r e s t unde r a s to n e i s s uch a s t o

b r ing i ts side s or dorsa l s u r fa ce in co nta ct with t he w all s o f i t s
hi ding pl ace Th igmo tropi s m no d o ubt is o f di s tinct a dv a n
.
, ,

t ag e Sinc e it fo r c e s th e a nim a l int o a pl a c e o f s a fe ty

,
.

P H OT OTR OP I SM Light o f v a ri o u s int e n s it ie s in th e m aj or ity

.

o f c a se s c a us e s th e c r a yfi sh to r e t r e a t i e to Sh o w ne ga ti ve
,
. .

p ho tot ro ism
p . Individu a l s p re f e r co l o r e d light s t o whit e h a ving ,

a Sp e c ia l liki ng f o r r e d N e g a tive r e acti o n s to light pl a y a n

.

imp o r t a nt r Ole in th e a n i m al s life sin ce th e y in flue nce it to se e k a

’

da rk pl a ce whe re it is co n ce al e d fro m its e n e m ie s .

C HE M OTR OP I S M T h e re acti o n s o f th e c r a yfi s h t o fo o d a r e d ue
.

in p ar t to a che mi cal s e n se Sm oo th h ai rs with n e rv e bundl e s

“
.
,

w ithi n th e m a r e p ro b a bly ch e mi ca l a nd sin ce

,
Th e a nim al s
, ,

r e a ct to ch e mic a l s timul a ti o n o n a n p a r t o f t h e b o dy w e
y
mu s t assum e th a t the re a r e ch e mic al s e n se o rga n s all o v e r th e
”
b ody ( 1 6 5 p ,
. T h e a nt e ri or app e nd a g e s h o w e v e r a re , ,

the m o s t se n sitiv e e sp e c i a lly t h e o ut e r ra mus o f th e a nt e nn ul e

,
.

P os i ti ve re acti ons re s ult fr o m th e applic a ti o n o f f o o d s ub st a nc e s .

F or e x a mpl e if m e a t j uic e i s pl a c e d in t h e w a t e r n e a r a n a ni m al
, ,

th e a nt e nn a m o ve Slightly a nd th e m o uth p a r t s p e r f or m vig oro u s

che wing move m e nt s T h e m e a t c a u se s g e n e ra l r e stl e s sn e ss
.

a nd v a gu e m o v e m e nt s t o w a r d th e so u rce o f t h e s timul a ti o n but

,

th e a nim a l s se e m t o d e p e nd chi e fly o n t o uch f o r th e a ccu r a t e

l o ca liza tio n o f foo d ( 1 6 5 p Acid s sa lt s s ug ar a nd o th e r
,
.
, , ,

ch e mic a l s p r o duc e a so r t o f ne a ti ve r e acti on indic a t e d by s cr a t ch

g
ing th e car ap a ce r ubbing th e che l a or pulling a t the p ar t s ti m u
, ,

l a te d .

HAB I T F OR M ATI ON — I t h as b e e n sh o wn by c e r t ain Simpl e

.

e xp e r im e nt s th a t cra fi sh e s a r e a bl e t o l e a r n h abit s a nd t o m o dify

y
2 24 AN IN T R ODU CTION TO Z OOLOGY

the m Th e y l e arn by e xp e ri e nce and m o dify th e i r b e h avi o r

.
,

sl o wly o r qui ckly d e p e nding up o n th e i r f a mili a rity with t h e

,

s itu a ti o n T h e foll o wing e xp e rim e nt s p ro v e th e a b o v e s t a t e

.

m e nts C ra yfi she s whe n pl a c e d in co mp ar tm e nt T o f a l aby rinth

.
,

lik e tha t shown in Figu re 1 1 8 will o n a tt e mpting to e scape , , ,

p ass o n o ne sid e o r th e o th e r o f th e p ar titi o n P A t ran sp a re nt .

gl a ss sc re e n G p re v e nt s th e i r e xit fr o m th e right side o f the p a r ti

ti o n ; th e o th e r side h o we v e r b e ing l e ft op e n all o ws a fre e pa s
, , ,

sa g e . I t w a s fo und by Y e r k e s a nd H uggins ( 1 88 ) th a t a ft e r
s ixty t ri a l s cr ayfi sh e s th a t a t t h e b e ginning ch os e t h e cl ose d
,

p a ssa ge 50 pe r c e nt o f th e tim e l e a rne d to avoid tha t side a nd in , ,

9 0 pe r c e nt o f the
t ri a l s cho se the
o p e n e xit Whe n .

t he gl ass pl a te
w as th e n r e m ov e d
fro m th e right o f
th e p ar titi o n and
p l a ce d o n the
o th e r sid e th e ,

c r a yfi s h e s w e r e

FI G 8 L by i nth
1 1 d i n p i m nt
a r n th
u se co nfu se d a t fi r St
ex er e s o e cr a
. .
y )

fi h s T .m p t m nt f m whi h ni m l w
,
co ar but l e arn e d the
e ro c a a as

t t d ; P p titi n t it ; G gl
s ar e ,
ar pl t ne w h abit o f e s
o a ex ,
ass a e

l i ng n it ( F m W hb n f t Y k
c os o e ex .

ca
ro
In at
as
the
ur a er er es

a nd Huggi ns ) p g
ight op e n side
.

r ,

m ore quick ly th an b e fore T he chie f fa ctor s in th e for m a ti o n .

o f s u ch h a bit s are t h e ch e mic a l se n s e ( p ro b a bly b o th s m e ll a nd

t aste ) to uch sight a nd the muscular se n sa ti o n s r e s ulting f ro m

, , ,

th e di re cti o n o f tu r ni ng T h e a ni m al s a r e a bl e to l e ar n a p a th
.

whe n th e p ossibility of foll owi ng a sc e nt i s e x clude d ( 1 88 p ,

.

P ro fe ssor S J Holm e s w rit e s th a t h e h as t r ai n e d cray

. .

fishe s to come to him f or foo d .

226 AN IN T R ODU CTION TO Z OOLOGY

FI G . 1 20 . P e ri patus ca pe nsi s .
( Fr o m Shi pl e y a nd M a cB r i d e t
af e r Se d gwi ck .
)

with th e e xce pti o n of P e r i pa tu s , are gm e nt ally a r rang e d ;

no t se

Ann e lid s h a v e a we ll d e v e l op e d coe l o m ;

in a r thr op o ds the coe l o m i s r e st rict e d
t o th e ca viti e s o f th e e xcr e t ory a nd
r e p ro ductiv e o r g a n s .

T h e C r u s t a c e a a r e divide d int o t w o
sub cl a s se s th e E nto m o s t r a c a a nd th e
,

M al acos t r a c a . T h e E nt o m ost r a c a i n
e lud e s m os t o f th e s m a ll simpl e sp e ci e s

( Fi g . Th e se h a v e a va ri abl e
numb e r o f segm e nts ; no ga stric mill
in th e st om ach ; and in m a ny sp e ci e s
h a t ch a s a l a r v a l fo r m call e d a N au pli u s
( Fi g 1 2 6 A) T h e M al a co st r a ca a re
.
,
.

u su ally l a rg e . Th e y h av e a de finit e
numb e r o f se gm e nt s fi v e in th e h e ad
—
,

e ight in th e tho r a x a nd six in th e ,

a bd o m e n ; th e s t o m a ch c o nt a in s a ga s

t ri c mi ll ; a nd th e N a upliu s st age is
u su ally p asse d thr o ugh within the e gg
b e fo r e h a tching T h e cr ayfi sh b e l ong s
.

in thi s sub cl ass .

C l a ss Ony ch oph ora c o nt ain s o nly a

f e w a nn e lid lik e a nim a l s th e b e s t kn o wn
-
,

1 21 . Do r sa l
vi e w o f a b e ing P e ri pa tu s ca pe nsi s ( Fi g .

ce nti p e d e 1 a nt e nna ;
.
,
Se gm e nt all y a rr a ng e d n e ph ridi a s tump ,

2 p0 i s o n cl a w
, (Fr o m
.
like l e g s simpl e e ye s a nd t r ache a a r e
, ,

Shi pl e y a nd M ac s o m e o f t h e o r g a n s p o sse s s e d by m e m
Br i d e )
b e r s o f this cl ass
.

.
 Y I
T HE C R A F SH AN D ART HR OP ODS I N G EN ER A L

FI G . 1 22 . A m illi p e d e .
( F ro m Shi pl e y a nd M a cB r i d e t K o ch )
af e r .

T h e m yr i op o d l
s a r e a so a nn e lid lik e in app e a r a nce Th e t wo
-
.

p rin cip a l o r de r s a r e th e C hi l o p o d a a nd Dipl o p o d a T h e C hil o .

.

p o da o r c e ntip e de s ( Fi g . 1 2 1 ) a r e fl a tt e n e d d o r s o v e nt r a lly h a v e
-
,

o ne p ai r o f l e gs a t lV INS EC
. TA
rach e d to e v e r y s e g

m e nt b a ck o f th e
h e a d and p o sse ss a
,
A RA A
C HN ID
p ai r o f p oi so n cl aws
~

a tt ach e d t o th e fi rs t Ill . M YR A I PO DA

se gm e nt T h e Dipl o .

p o d a o r M illip e d e s c nus m si

( Fi g . 1 22 ) a re cyli n

d r i ca l, h av e tw o p ai r s
of l e g s a tt ache d to
R ATU
PE IP S
e ve ry s e gm e nt b a ck
of th e fo u r th and ,

l ack p o i so n cl aws .

T h e in s e ct s C OIl S tl R M T V A R T HR
P I I I E O PODS

tut e ab o ut fo u r fifth s
of a ll t h e sp e ci e s o f

a nim a l s T h e num
.
A L A
NNE ID

b e r o f individu a l s i s
e ve n m o re r e m a rk a
ble . T h e b o dy o f a n
in se ct i s divide d int o
h e a d th or a x and ab
; ,
‘

FI G . 1 22 . D i a gr a m to s ho w th e a ffi n iti e s
do m e n T h e tho ra x
.
ar th r op o d s .
2 28 AN INTRODU CTION To ZOOLOGY

b e ar s thr e e p air s o f l e g s a nd with a f e w e xc e pti on s one o r

, , ,

t wo p ai r s o f wing s T h e a bd o m e n h as se v e r a l o r no a pp e nd
.

a ge s . Th e h o n e yb e e d e scri b e d i n C h apt e r XII h a s b e e n s e l e ct e d

a s a typ e o f t hi s cl a s
T h e C l a ss Ar a chnid a includ e s a nim al s whi ch h a v e th e h e a d
a nd th o r a x fu se d int o a c e ph a l o th or a x The y p o ss e ss fo u r p air s
-
.

o f l e g s a n abd o m e n u s u a lly with o ut a pp e nd a g e s a nd a h e a r t in

, ,

th e d or sa l p a r t o f th e a bd o m e n Such di v e rse a ni m al s a s th e
.

spid e r sc o rpi o n mi t e
, ,
a nd King c r a b a r e pl ace d t o g e th e r in thi s
,

cl ass T h e p r ob abl e affi niti e s of th e v ario u s cl a sse s o f a r th ro

.

p o ds a re shown in F igur e 1 23 .

Th e B i oge ne ti c Law In c on cluding thi s ch a pt e r we wish

.

t o r e fe r t o a law which h a s c o mm a nd e d th e a tt e nti o n o f

z ool ogi s t s f or almo st a c e ntury ; n a m e ly the law of bi o ,

g e n e sis a l so kno wn a s th e r e ca pi tu la ti on the ory Org a ni c e vo

,
.

lu ti on th a t is th e e v o luti o n o f o ne o r g ani s m fr o m a n o the r

, , ,

is a cce pte d as a n e stabli she d fa ct by pr acti ally all z oo l o gi s t s

at th e p r e se nt tim e This f act of org anic e vo lutio n i s
.

e xp r e sse d in the di ag ra m o f th e p r incip al phyl a of th e

a nim a l kingd o m o n p a g e 7 whe r e th e v a ri o us g ro ups app e a r
,

to be de riv e d fr o m a c e nt r a l st e m wh ch r e p r e se nts a i

se ri e s o f a nce s tr al fo r m s F r o m a n e x a min a tio n o f th e phyl a

.

r e p re s e nt e d in ou r di a g ra m w e g a in th e f a ct th a t th e m e mb e r s of

e a ch g r o up a r e m o r e c o mpl e x th a n th ose o f th e g ro up ju s t b e n e a th

the m on th e p age Evoluti o ni sts do no t cl aim th a t th e m o r e

.

compl e x fo rm s h ave e vo lve d di r e ctly fro m th e Simpl e r ani m al s ,

but th a t the i r a nc e s t o r s w e r e r e l a te d B e ginning with the Si m .

p l e s t a nim a l s w e find th a t a s ingl e c e ll p e r f o r m s a ll t h e n e ce ss a r y

p ro ce sse s of life e g Ame ba Within the l o we s t phylum the

,
. . .
,

P r o t o zo a th e r e a r e a nim al s c o n si s ting o f a numb e r o f c e ll s m o r e

,

o r l e ss intim a t e ly b o und t o g e th e r int o a h oll o w Sph e ric a l c o l ony ,

e g Volvox
. . P a ssing to th e n e xt high e r g ro up o f o r g a ni sm s w e ar e
.

intr o duce d to anim als th a t p osse ss t w o l aye r s of ce ll s s urro unding ,

a singl e c a v ity e g Hyd r a

,
. . All a nim a l s a b o ve th e C oe l e nt e r a t e s
.

h av e thr e e l aye r s of c e ll s fo r mi ng th e i r b ody wall s i e a re t riplo ,

. .
 23 0 AN I N T R ODU C T I ON T O Z OOLOGY

FI G . 1 25 . M ys i s oc u la ta . Se d gwi ck

n e a rly lik e th e a dult a s do e s th e

c ra yfish . Th e l o b st e r f o r,
ex

FI G . 1 26 . T wo t
s a ge s i n th e d e v e l o p
m e nt of th e s h ri m p ,
R e na ns . A,
N o npli ns s t a ge ; B P r otozoa a s t a ge .

( Fr o m Se d gwi ck t
af e r Fr it z
M ull e r ) .
 T HE Y I
C R A F SH AN D ART H R OP OD S I N G EN E RAL 23 1

ampl e up o n h a t ching ( Fi g 1 24) re s e mbl e s a l e ss sp e ci a liz e d

,
.

p r awn like cr us t ace an call e d M ysi s ( Fi g 1 2 a nd i s said to be

-
,
.

in th e M ysi s st age T h e Sh rimp P e na u s p asse s th ro ugh a

.
, ,

num b e r o f int e r e sting st a ge s b e for e th e a dult co nditio n is

FI G . 1 27 . T wo l a t e r s t a ge s i n th e d e v e l o pm e nt o f P e na u s A Z oa a s t age
.
, ;
B, M ys i s s t a ge
.
( Fro m Ko r s ch e lt a nd He i d e r a f t e r C l a u s )
.

a tt ain e d It h a tche s a s a l arv a t e r m e d a N a u pli us ( Fi g 1 2 6 A)

.
,
.
, ,

po sse ssing a fro nt a l e ye and thr e e p a i rs o f app e ndage s (A A "

,

thi s N au plius m oltS a nd g ro w s int o a P r otozoa a s t ag e

( Fi g 1 2 6 B ) which b e ar s th re e m o r e p ai r s o f app e nd a ge s a nd
.
,

th e r udim e nt s o f se gm e nt s III VIII

— T h e P r o to z o a a st ag e
.
23 2 AN INT RODU CT ION TO ZOOLOGY

g ro ws into th e Z e a e st age ( Fig 1 2 7 A) T h e c e ph a l o tho r a x

.
,
.

a nd a bd o m e n are di s tinct a t t hi s tim e ; e ight p a i r s o f a pp e nd a g e s a r e

p re se nt ( I VIII) and si x mo re ar e de ve l oping (a a ) T h e Z o a a

— —
l 6 .

g ro ws and m olt s and b e co me s a M ysi s ( Fi g 1 2 7 B ) with e ight .

,

p ai rs o f app e nda g e s ( I VIII ) o n th e c e ph a l o thor a x Fin a lly th e

— .

M ysi s p a sse s int o th e a dult sh rimp whi ch p osse sse s th e ,

ch ara ct e ris tic numb e r o f app e ndage s (I XIX ) e ach m o difi e d-

t o p e r for m i t s p a r ti cul a r fun cti o n . T h e N a u pli u s o f P e na u s

r e s e mbl e s th e l a r v a o f m a ny simpl e cr u s t a c e a n s th e Z oa a i s
s ome what sim il ar to th e co nditi o n o f a n a dult Cyclops ( Fi g .

th e M ys i s is lik e th e a dul t M ys i s ( F i g 1 2 . a nd fin a lly th e a dult

P e na u s is m o r e Sp e ci a liz e d th a n a ny o f its l ar v a l s t a g e s a nd b e ,

l ongs a mong th e highe r C r ust a c e a T h e a b ove fa ct s h ave co n

.

vi nce d s o m e z oo l ogist s th a t P e na u s r e c a pit ul a t e s in i t s l arv a l

de ve l opme nt the p r og re ss o f th e ra ce ; tha t th e l ob st e r h a s l o st

m any of the se s t age s r e taining o nly th e M ysi s ; a nd tha t the c ray
,

fi sh h a tch e s i n p r a ctic a lly th e a dult c o nditi on T h e N a u pli us

.

st ag e o f th e l a tt e r i s s upp o s e d t o b e r e p r e s e nt e d by a c e r t a in

e mb ry o nic ph a s e ( Fi g .

T h e la w o f bi oge n e si s Sh o uld no t b e t a k e n t oo se ri o u sly sinc e ,

it h as b e e n cr iticiz e d se v e re ly by m any p ro min e nt zo ol ogis t s '

,

but it h as fur ni she d a n hyp o th e si s which h a s c o nc e ntr a t e d th e

, .

a tt e nti o n o f s ci e nti s t s up o n fund a m e nt a l e mb ry o l o gic a l p r oce sse s ,

a nd h a s th e r e fo r e h ad a g r e a t influ e nc e up o n z o ol o gic a l p r og r e ss

( 1 75 . 1 77 .
 23 4 AN INT RODU CTION TO Z OOLOGY

Th e Dr o n e dr o n e ( Fi g 1 2 8) is the m al e b e e ; h e live s
.
— Th e .

in idle ne ss up o n th e foo d ga the r e d by th e w o r ke r s Hi s functi on .

,

ho we ve r i s a n imp o r t a nt one ; n a m e ly t o m a t e with th e qu e e n

, , .

Ho w thi s is acco mp lishe d wi ll be d e s crib e d l a t e r T h e b o dy o f .

th e d ro n e i s h e a vy a nd b r oa d a nd th e hi nd l e g s p oss e s s no p o ll e n
,

b aske t Hi s e ye s a r e co n sid e r ably l a rg e r th a n th o s e o f e ith e r th e

.

que e n o r wo rk e r .

Th e Wor k e r T h e wo r k e r ( Fi g 1 2 8) i s a s e x u a lly und e v e l o p e d

. .

fe m al e Unde r n o rm al co nditio n s it l a ys no e ggs but i s k e pt

.
,

Wo r k e r Q ue e n D r o ne

FI G . 1 28 . Ho ne ybe e , Api s me llifi ca .

( Fr o m Shi pl e y a nd M a cB r i d e )
.

bu sy g a the ring ho n e y p o ll e n a nd p ro p o li s or be e glue ca rry

, , ,
-
,

ing wa te r se cr e ting w a x b u i ldi ng co mb p r e paring foo d nu rsi ng

, , , ,

th e y o ung a nd cl e a ning a nd d e fe nding th e hi v e

,
Th e w o rk e r s .

a r e s m a ll e r th a n e ith e r qu e e n o r d r o n e ; th e y a r e th e b e e s u s u a lly

se e n h o v e r ing a b o ut fl o w e r s T h e a n a t o mi ca l d e s c r iptio n th a t
.

foll ow s co nce r n s th e w or ke r unl e ss o th e r wi se s ta te d .

An a tomy a nd P hys i ol ogy EX O SK E LE T ON . T h e b o dy o f the .

be e i s co mpl e t e ly co ve r e d with a s kin o r cuti cl e co n si s ting o f a ,

ve ry thin l aye r o f chitin wh ich i s se cre te d by th e hyp o de rm a l ce ll s

lying jus t b e n e a th it Thi s chitin p r o te cts th e in se ct fro m inju ry
.

a nd giv e s th e b o dy s t r e ngth D u ring th e y o ung s t age s th e e xo

.

sk e l e to n of in e l astic chitin i s c as t off a t inte r v als all owing the ,

b o dy to e xp and .

R E GI ON S OF TH E B ODY — Thre e di s tin ct re gio n s m ay b e r e cog

.

ni z e d in th e b ody o f th e b e e th e h e a d th o r a x a nd a bdo m e n , ,
.

E a ch divi si o n co n s i st s o f a d e finit e numb e r o f s e gm e nts m or e or

l e ss intim a te ly fuse d t oge the r .
FI G . 1 29 . He a d w o rk e r ho ne yb e e
of . a a nt e nna ; b bo u t o n ; g
, , ,
e
pi ph ar<

ynx ; m, m a nd i b l e ; m x m a xill a ; m xp m a xill a r y pa l p u ;

.
, .
, s l, hypo
p h a rynx ; lp l a b i a l p a lp u s
.
, ( Fr o m P a ck ar d a f t e r C h e s hi re )
. .
23 0 AN I N T R ODU C T ON T O I Z OOLOGY

int o th e oe sopha gu s T h e n e ct ar i s fi r s t c oll e ct e d by th e h ai rs on

.

th e ligul a ; it i s th e n f orce d upw a r d by t h e p r e ssing t o g e th e r o f

th e m a xill a a nd l a bi a l p a lpi .

T HE T H OR Ax AN D I T S AP P E N DA GE S T h e th o r a x c o n si s t s of .

thre e se gm e nts e a ch o f which b e a rs a p a ir o f l e g s T h e a nte ri or

, .

s e gm e nt i s kno wn a s the pr othor ax ,

t h e middl e se gm e nt a s th e m e sotho
r ax a nd th e p ost e ri o r s e gm e nt a s
, ,

th e m e ta thor a x T h e m e s o th or a x .

a nd m e t a th o r a x e a ch supp o r t a p a i r

o f wi ngs T h e se gm e nt s o f th e .

thor a x a r e co mp ar a tive ly l arg e ,

s ince th e y co nt a in t h e l a r g e s t a nd

m os t imp o r ta nt mu scl e s o f the

b o dy E xt e r n ally th e thor a x i s .

co v e r e d with fl e xibl e b r an ch e d
h ai rs whi ch a r e o f u se in g a th e ring ,

p oll e n ( Fi g .

LE G S P e r h a p s th e m os t i n .

t e r e s ti ng s t r u ctu r e s o f th e h o n e y
b e e a r e t h e l e g s o f t h e w o r k e r ( Fi g .

Th e y a re won de rfully
a d a pt e d f o r th e w o r k whi ch th e y
FIG 1
3 A w k
0 ° h n yb p e r fo r m T h e p a r t s o f a typical
, or er o e ee
.

with p ll n b k t f ll ; B
o e as e u
I n se ct le g na m I ng th e m I n o r d e r
,

p t f m th i l g with ,
ar o e so or ac c e
b e g mn mg a t th e p r o xi m a l e nd a r e
b n h d h i
ra c e nd p ll n a rs a o e ;

gr a i ns ; C o ne h a i r b e a r i ng th e coxa (c)
, ,
tr ocha nte r fe mu r
p o ll e n gr a i ns ( F r o m t
~ h e
(f ) ti bi a ,
a nd fi v e -j oint e d
C a m b r i dge N a t ura l Hi s
ta r su s ( t)
t o ry ) T h e pr othor aci c l e g s C)
( g
F i
'

. 1 3 1 ,

p oss e ss fo ll o wi ng u se ful
th e a nd tibi
s t r uctu r e s
(f ) T h e fe mu r
a .

( )
ti
. a r e Cl o th e d with br a nche d ha i r s f o r g a th e r ing p o ll e n E xt e nd .

i ng o n o ne sid e fro m th e di s t a l e nd o f th e tibi a a r e a numb e r o f

cur v e d b ri s tl e s th e polle n br u sh (b in C a nd E) whi ch a re use d to
, ,

b r ush up the p oll e n l oo se n e d by th e coar se r spin e s ; o n th e o the r side

FI G . 1 3 1 . Le gs wo rk e r b e e (Af t e r C h e shi r e ) A o u t e r s i d e o f m e ta
of . .
,

tho r a ci c le g : p m e t a t a r s u s ; t t a r su s ; ti tib i a B i nne r s i d e o f

, ,
.
,
.
,

m e t a th or a ci c le g : 6 co xa ; p m e t a t a r su s ; t t a r s u s ; ti ti b i a ; i r t ro
, , ,
.
,
.
,

ch a nt e r ; wp w a x p i nch e r s
.
, C pr o th o r a ci c le g : b p o ll e n b r u s h ; e b
.
, ,
.
,

e ye b r u s h ; p m e t a t a r su s ; t t a r s u s ; ti
,
tib i a ; v ve l u m D m e so
,
.
, ,
.
,

thor a ci c le g : l e tt e r i ng a s i n C ; s po ll e n sp u r E j o i nt o f p ro th o ra ci c
,
.
,

le g ; l e tt e r i ng a s i n C F t e e th o f a nt e nna co m b
.
,
G cr oss se cti o n o f .
,

tib i a th ro u gh po ll e n ba sk e t ; f a po ll e n ; 11 ho l di ng h a i rs ; n ne r v e
.
, , ,
.

H a nt e nna i n p ro ce ss o f cl e a ni ng ; a a nt e nna ; 6 a nt e nna co m b ; l

, , ,
.

s e cti o n o f le g s scr a p i ng e d ge o f v v e l u m
, ( Fro m R o o t )
,
. .
23 8 AN INTRODU CTION TO Z OOLOGY

in this b aske tlike st r uctur e it i s p ossibl e f or th e b e e to sp e nd m ore

,

tim e in th e fi e ld and to c a r ry a l arg e r l oa d a t e ach t rip T h e

, .

p o ll e n b ask e t i n cro ss s e cti o n i s sho wn in F igu r e 1 3 1 G Th e ,

.

p olle n c o m bs ( a t p i n B ) s e r v e t o fill t h e b a s k e t s by c o mbing out

th e p oll e n whi ch h a s b e co m e e nt angl e d in t h e h ai r s o n th e th o r a x

, ,

a nd t r a nsf e r r ing it t o t h e c o nc a vity in th e tibi a o f th e o pp o s it e

le g . At th e di s t a l e nd o f th e tibi a i s a r o w o f wide spin e s ; th e se

a r e o pp o se d by a s m oo th pl a t e on th e p r o xim a l e nd o f th e m e t a

t a rsu s T he t e r m wa x pi nche rs (wp in B ) h as b e e n appli e d to

. .

the se str uctu re s sin ce the y a re u se d t o r e m ov e th e w ax pl at e s

,

fro m th e abdom e n of the w o r k e r .

WALK I N G In w alking thr e e l e g s a re e xt e nde d a t o ne tim e

.
,

th e o th e r thr e e supp o r ting t h e b o dy lik e a t r ip o d T h e fo r e l e g .

pulls th e b o dy fo r wa r d th e middle le g m aint a in s e quilib rium

, ,

a nd t h e hind le g pu s h e s t h e in se ct f o r w a r d T h e d e t a ils o f walk .

ing in in se cts a r e ve ry c o mplica t e d .

WI N GS . E a ch o f th e fi r s t tw o th o r a cic se gm e nt s b e a rs a p ai r
o f m e mb r a n o u s wing s Wh e n a t r e s t th e wi ngs li e h o riz o nt ally
.

o v e r t h e a bd o m e n but wh e n flying a r e wid e ly sp r e a d

,
T h e wings .

m a y b e d e sc r ib e d a s t r a n sp a r e nt m e mb r a n e s s upp o r t e d by h oll o w
r ib s c a ll e d ne rve s o r ve i ns T h e p ai r o f wing s o n o ne sid e o f th e
.

b ody m ay be j o in e d to ge the r by a r o w o f hookle ts on th e ante rior

m argin o f th e hind wing whi ch a r e inse r te d int o a tr oughli lae fold
,

in th e p ost e r i or m argin of th e fo re wing .

F LI GH T . Wh e n flying th e wing s act a s inclin e d pl an e s and ,

l ocom o tion fo rw ar d i s a tt a in e d by b o th up and do wn st rok e s ,

t h e tip s o f t h e wing s m o ving in a cu r v e sh a p e d lik e a figu re 8 .

M o tio n b a ckw a r d o r a s ud d e n s t op m ay b e a c co mpli sh e d by

, ,

ch anging th e inclin a tio n o f th e pl an e o f oscill a ti on .

T H E AB DOM E N AN D I T S AP P E N DA GE —
T h e abd o m e n i s m ad e u p
.

o f a s e r i e s o f six vi s ibl e se gm e nt s o r r ing s o f chi tin incl os ing a l a rg e

p ar t o f th e alim e nt ary c an al n e rv o u s syste m r e p ro ductiv e organ s

‘

, , ,

e tc . E a ch r ing h as a d or s a l pl a t e th e te r gu m a nd a v e nt ra l pl a t e
, , ,

th e ste r nu m Thin chitino u s m e mb r an e s co nn e ct th e r ing s and

.
,

m ak e th e m ove m e nt and e xp an sio n of th e abdome n p o ssible .

 k , l, to m o e
Sti ng o f th e
w b a rb s o n t
d ar s l v v
b e rs
rk e r b e e
e
FI G . 1 33 . o .
,

t v ti l i l d i on sa c ;
d a r s ; n ne r e s ; p s ng f e e e r pg p
, ,
- . o
,
s o n g a n ; ps .
p o,
s

P k d f
S h e a th ; fi th i a l ga gli t
sh th f a b d o m n n o n ( F ro m a c a r a er
.
, 5 ,
.

C h e shir e ) .
2 40 .
AN IN T R ODU CTION TO ZOOLOGY

by m e ans o f h ar d t e nd o ns M u sc u l a r a cti o n i s e ith e r volunt ary

.

o r invo lunt a ry ; f or e x a mpl e th e j a w s a nd wing s a re m o v e d by

,

v ol unta ry m a ny inte r n a l o rg a n s by invo lunt a r y muscl e s T h e

, ,
.

st r e ng th o f the mu scl e s o f th e be e i s much g r e a t e r tha n tha t of

th e mu scl e s o f m a n c o mp a r e d with th e i r w e ight
,
T h e e xpl a n a .

tio n o f thi s i s quite simpl e sinc e th e we ight o f a muscl e in cr e a se s

,

a s th e cub e o f its di a m e t e r whil e i ts st r e ngth in cre a se s only

,

a s t h e s qu a re A l a rg e a nim a l ca n pu ll th e r e f or e co mp a r a tiv e ly
.
, ,

l e ss tha n a sm a ll e r one .

T H E DI GE S TI VE SY S TE M ( Fi g — T h e dig e stiv e ca n a l i s
.

m a de up of the foll o wing s t r ucture s n a m e d in o r de r b e ginning a t ,

th e a nt e r i or e nd : th e m o uth oe s oph a gu s o r gull e t h o n e y sa c o r

, ,

ho n e y s to m a ch t r u e s to m ach sm a ll int e s tin e o r il e um and l arg e

, , ,

inte stin e o r col on It op e n s a nte r i o r ly by th e m o uth and pos

.

te ri o rly by th e a nu s T h e oes ophagu s (oe s ) is a n a rr o w tub e

. .

whi ch p a s se s th ro ugh th e tho r a x ; i t s p o st e r i o r e nd i s e nl a rg e d

into th e hone y s a c (hs ) Situa t e d n e a r th e a nte r io r e nd o f th e abdo
.

m e n At th e p os t e r i o r e nd o f th e h o n e y sa c i s th e s tom a ch
.

m ou th (p) ; thi s s t r uctu re e xt e nd s Slightly f o r w a r d int o th e

h o n e y s a c It h a s fo u r t r i angula r lips whi ch m a y b e Op e n e d or
.
,

cl o se d by t w o se t s o f mu s cl e s l o ngitudin a l a nd cir cu l a r N e a r .
,

th e t op o f th e l i p s a r e a numb e r o f b r i s tl e s whi ch p r oj e ct b ack

wa r d If th e ali m e nt a ry can a l o f a fr e shly kill e d be e i s pl a ce d in
.

a to 3} pe r ce nt s a lt s o luti o n th e lip s will op e n a nd cl o se

,

r a pidly f or a o h a lf a n h o u r
b ut .
.

Th e tr ue s tom ach i s a cylind r i ca l sa c ; its w a lls cont a in a

nu mb e r o f ci r cul a r mu s cl e s a nd a l a ye r o f l o ngitudin al m u scl e s
,
.

T h e dige s tive jui ce s s e cr e t e d by i ts w a ll s ch a ng e th e fo o d int o

chym e P a r t o f thi s chym e i s a b so r b e d th e r e s t i s fo r ce d by
.
,

muscul ar c o nt r a cti ons i nt o th e sm all int e s tin e Un di ge ste d foo d .

i s dis so lve d in th e s m a ll i nte s ti ne or il e um a nd th e dig e s t e d

,

f oo d o r chyl e i s ab sorb e d by i ts w all s At i t s p o s t e ri or e nd th e .

sm all int e stin e g r a du a ll y m e rg e s int o an e nl arge m e nt th e colon (Z) ,

.

Thi s p a r t o f th e a lim e nt ary ca n al r e ce ive s all th e un di ge ste d

matt e r a nd disch arge s it to th e o ut sid e th r o ugh th e a n us T he
, .
1 34 . I nt e r na l h o ne ybe e bl M a l p ighi a n t ub ul e s ; o s t rue
o r ga ns o f . .
, . .
,

s om ac
t h ; d v d o r sa l ve sse l ; e e ye ; g ga ngli a o f ne r ve cha i n ; bs
. .
, , ,
- .
,

ho ne y sa c li r e ct u m [ p l a b i a l pa l p u s m e s a t m e so tho ra x m e ta t
.
, .
, . .
, . .
,

m e t a th o r a x ; m ac m a xill a ; n ne r v e s N o 1 N o 2 N o
; 3 sa liva ry
.
, , .
,
.
,
.
,

gl a nd s ; oe s .oe so p h a gu s ;
, p s t o m a ch m o u th ; p o t p ro th o r a x ; s i
, r . .
,
.
,

s m a ll i nt e s ti ne
( il e u m ) v v e nt r i cl e s o f d o rsa l ve ss e l
, ( F ro m P a ck a r d .

a f t e r C h e s hir e ) .
24 2
‘

AN IN TRODU C T ION TO ZOOLOGY

f a ce s a r e n e ve r vo ide d within th e hive if th e b e e s a r e k e pt unde r

p rop e r co ndi tion s .

T HE SA LI VAR Y Th e r e are tw o p a ir s o f s alivary

GL AN D S — .

gl a nds o ne withi n th e h e a d th e o th e r withi n th e th o r ax Those

, ,
.

in th e h e a d li e a g a in s t th e p o s te ri o r w a ll s o f th e cr a ni um T h e .

o th e r p a i r li e in t h e v e nt r a l p a r t o f th e a nt e r i o r h a lf o f th e th or a x .

B o th p ai r s o f gl a nd s p ro du ce w e a kly a lk a lin e s e cr e ti on s whi ch a re

p o u r e d o u t up o n th e l abium wh e r e th e y a ct up on th e fo od tak e n
th ro ugh th e p r o b o s ci s ( Sn o dg r a ss ,

T HE E X C R E TOR Y OR GA N S — . T he M a lpi ghi a n o r u r i na ry tu bu le s

( Fi g . I 3 4 ,
bt ) . a re a numb e r
l ong fi ne h ai rli ke str uctu r e s whi ch
of , ,

o p e n int o th e a nt e r i or e nd o f th e int e s tin e Th e y w e r e di scove re d .

by a nd n a me d a fte t e g e a t It a l a n an a t omi st M alpighi Ex

r h r i .

cr e ti o ns a r e t a k e n f r o m t h e bl o o d a nd th e f a t b o dy in th e f or m

o f u r a t e s ; th e y p a ss int o t h e int e s tin e a nd th e n ce ou t o f the ,

b ody .

T H E VA S C U LAR SY S TE M —
T h e blood o f th e h o n e yb e e i s simil ar
.

t o th a t o f t h e cr a yfi sh It i s a co l o rl e ss pl a sm a co nt aining a m e
.

b o id co rpu scl e s It diffe r s in one imp o r t ant p o int fr o m th e bl ood

.

o f m o s t o th e r a nim a l s i t d oe s no t ca r r y v e r y mu ch if a ny
—
, ,

o xyg e n T h e di s t r ibuti o n o f o xyg e n i s a cc o mpli sh e d by th e

.

r e s pi r a t o ry s y s t e m a s w e sh a ll s e e l a t e r
,
.

An e ve n l e s s co mpl e t e s y s t e m o f blood ve s s e ls i s p r e se nt th an
in th e cra yfi sh T h e p r in cip a l o r ga n o f ci r cul a tio n i s th e d orsal
.

ve sse l o r he ar t ( F i g 1 3 4
,
Thi s i s a tub e lyi ng in th e m e dian
.
,

l ine ju st b e n e a th th e do r sa l s u r fa c e ; it i s cl o se d ne a r th e p ost e ri or
e nd o f t h e a bd o m e n but o p e n s in t h e h e a d ,
T h e w all s o f the .

h e a r t a re muscul a r a nd co nt r a ct a t i nt e r v a l s B l oo d e nte rs
,
.

thr o ugh fi ve p ai r s o f os ti a o ne p a i r t o e a ch of th e fi v e c omp a rt

,

m e nt s o r ve ntr i cle s (v) int o whi ch th e h e a r t i s div id e d Va lve s .

p r e ve nt the fl o wing b a ck o f th e bl oo d s o th a t e a ch co ntra cti on ,

se nd s a st r e a m t o w a r d t h e h e a d F ro m h e r e it p a sse s th r o ugh th e
.

sp ace s a mo ng th e ti ss u e s fin a lly r e a ching th e v e nt r al p ar ts o f th e

,

b o dy B e n e a th th e h e a r t i s a h or iz ont a l di aphr agm o f muscl e

.
,

which fo r m s a pe r i car di a l s i nu s B y cont r a cting thi s dia phragm .

,
 T H E HON E Y BEE AN D B EE S I N GEN E R AL 243

forc e s the olood fro m th e v e ntral p a r t o f th e b o dy into the s inus ,

a nd th e nc e th r o ugh t h e o s ti a int o th e h e a r t .

T HE R E SP I R AT OR Y SY S TE M ( F i g 1 3 — T h e h o n e yb e e b r e a th e s .

through op e ning s c all e d spi r acle s ,

situa t e d in the side s o f ce r
ta in thor a ci c a nd a bdo min a l se gm e nts N o air e nte rs thro ugh .

a p e r tu re s in t h e h e a d a s in ,

ve r te b ra te s Th e re a re two .

p ai rs of sp i ra cl e s in th e thor a x ,

o ne in th e Side s o f the pr o
th ora x the o the r i n th e m e ta
,

thor a x F iv e p a i rs a re p r e se nt
.

in th e a bd om e n T h e spi r a cl e s .

o p e n int o tub e s c a ll e d tr ache a ,

.

Th e se unit e with l ongitudin a l

t ra ch e ae which e xte nd al ong th e
sid e s o f th e b o dy Oth e r tr a .

ch e aa ar i se f r o m th e s e longi t u

di n a l t r unk s a nd di s t ribut e th e i r
b r a n che s to all p ar ts of th e
b o dy T he t r a ch e ae ( Fi g 1 3 6)
. .

a re tub e s c o mp ose d o f a s ingl e

l a ye r of c e ll s ( a ) a nd lin e d with
a thin chi tin o us w a ll This w all .

i s thi ck e n e d a t r e gul a r int e r

v al s for mi ng a spi r al thr e ad
, ,

whi ch se rve s to ke e p th e t r a FI G 3 6 P ti n f t ch . 1 . or o o a ra ea . a,

ch e ae op e n C e r t a in t r a ch e ae
.
ll
ft
l
w
L yd g)
ll ( F m P k d ce

a
u ar

er e l
a . ro ac ar

a r e di l a t e d int o a i r s acs
( Fi g .

t h e l a r g e s t o f which a r e situ a t e d in th e a nt e ri or p ar t o f th e

FI G . 1 35 . R e spi ra t o ry s ys t e m o f w o r k e r b e e a s se e n f ro m abo ve ,
o ne an e t
i
r or pa i r o f a bd o m i na l s acs r e m o v e d a nd
tra ns v e rs e v e ntr al com m i ss ur e s
o f a b d o m e n no t s h o w n I S p I I I S p VI I S p sp i r a cl e s ; Ht Tr a S e . . . .
,
,
. . .
, ,

Tr a S e 1 2 4 7 8 I O t r a ch e a l s a cs ; Tr a t ra ch e ae
. .
, , , , , , , ( F ro m Snod .
,
.

grass T e ch ni ca l Se r i e s 1 8 B u r e a u o f E nt om o l o gy U nit e d St a t e s De
, , ,

par tm e nt o f Agr i cu lt ur e ) .
244 AN INT R ODU C TION TO Z OOLOGY

a bd o m e n The se a r e supp ose d t o b e o f se r vi ce to th e b e e during

.

flight sinc e the i r Size can b e in cr e ase d a t will a nd th e sp e cific

, ,

g r avity o f th e in se ct the r e fo r e d e cre a se d Ai r i s dr a wn int o

, , .

a nd e xp e ll e d f r o m t h e t r a ch e w by a lt e r n a t e e xp a n s i o n s a nd co n

t ra ctio n s o f th e a bdo m e n E ach spi ra cl e co nt a in s a v a lve which

.
,

m ay b e cl o se d o r Op e n e d D u s t i s p r e v e nt e d f ro m e nt e r ing by
.

h a i r s wh ich surro und th e op e ning .

Th e r a te o f r e s pi r a ti on d e p e nd s up o n th e a ctivity o f th e indi
vidu al N or m a lly th e r e a r e ab o ut fo r ty in spi r a ti on s pe r minute
.
,

but in fa tigue d b e e s the numb e r re a ch e s a s high a s o ne hundre d

a nd Sixty pe r mi nut e T h e a i r b r o u ght int o th e b e e s b o dy i s
’
.

c arri e d by th e tr a ch e ae dir e ctly to th e ti ssue s no cir cula tor y sys ,

t e m b e ing n e c e ss a ry f o r th e d is t r ibuti o n o f o xyg e n .

T H N R V OU S SY S M ( g 1 3 7 Op F i L — — T h e n e rvo us
E E TE 7 .
,
. .

s y s t e m o f th e b e e co n si s t s o f a l a r g e ch a in o f p a i re d g a ngli a a nd

t w o g r o up s o f s m a ll e r g a ngli a ca ll e d t h e s t o m a t o g a s t r ic a nd th e
, ,

s ymp a th e ti c r e sp e ctiv e ly ,
T h e l a r g e g a ngli o nic ch a in i s by f a r
.

t h e m os t imp o r t a nt It i s fo r m e d o f se v e n m asse s o f n e r vo u s
.

tiss ue tw o in th e h e a d tw o in th e thor a x a nd fi v e in t he abdo

, , ,

m e n E ach m a s s i s co mp o se d o f t w o g a ngli a lying s id e by Sid e

.
,

a nd c o nn e ct e d with th e m a s s in f r o nt a nd b e hind by tw o n e r ve

co r d s All e x ce pt th e for e m o s t g a ngli o ni c m a ss a re Situa t e d

.
, ,

n e a r the c e nt e r o f th e v e nt r a l b o dy wa ll T h e g angli a a t the .

e xt r e m e a nt e r i o r e nd o c cupy a ca vity in th e d o r s a l p a r t o f th e

h e a d ; the y a r e kn own a s th e br a i n o r s up r a oe soph age al g a nglia ,

.

N e rv e s co nne ct th e b r a i n with th e co mp o und e ye s ( Fi g 1 3 7 0p

’

. .
,

t h e o ce lli th e a nt e nn ae a nd th e l a b r um
, ,
B e n e a th th e oe soph a .

g u s in t h e h e a d li e s t h e s u boe s o h
p g
a e a l a
g g n li on whi ch in n e rv a t e s

th e m a ndi bl e s l abium a nd o th e r m o uth p a r t s

, ,
T h e a nte r i or .

FIG . 1 37 . Lo ngit u d i nal , m e d i a n, v e r ti ca l se c ti o n of e ti r e

n bo dy of work e r
be e , ho wi ng ne r vo us sys t e m (Op
s 7 s
.
y s t e m ( Tr a S c
L — . t r a ch e a l . .

1 d o r sal a nd v e nt r a l d i ap h r agm s o f a b d o m e n ( D Dph a nd V . . .

a nd d o r s al v e ss e l co ns i s ti ng o f h e a r t ( Ht ) a nd ao r t a ( F rom
Sno d gr a ss T e ch ni ca l Se r i e s 1 8 B u r e a u o f E nt o m o l o gy U nit e d St a t e s
, , ,

De pa r tm e nt o f Agr i cu lt u r e ) .
 T HE HON E Y B EE AN D B EE S I N G E N E RAL 24 5

ga ngli on in nd
th e thora xs n e r v e s int o th e f r o nt p a i r o f l e gs
se .

T h e poste r i or thor aci c ga ngli on i s c o mp a r a tiv e ly l a r g e c o nsisting ,

re a lly o f s e v e r a l g a ngli a wh i ch h a ve g r o wn t o g e th e r Th e a n .

t e ri o r p a r t of thi s g a ngli o n suppli e s th e f o r e wing s a nd th e middl e

p air of l e gs ; the p os te ri or p ar t inn e r va te s the hind wing s a nd
l e gs Va rio us p ar ts o f the abdo m e n a re suppli e d with n e rv e s
.

fro m the a bdomi na l ga ngli a ; th e l as t o f the se i s la rge r th a n the

o th e r s ,
b e c a use o f th e imp o r t a nt o r g a n s th e ge ni tal app ara tus
,

a nd th e sting wh ich a r e inn e r v a t e d by th e m

, .

Th e stomato—gas tr i c p a r t o f th e n e r v o u s s y s t e m is m a d e up o f
m any sm all ga ngli a co nn e cte d with the o rg a ns o f dige stion ci r ,

c u l a ti o n a nd r e spi r a tio n
,
.

E a ch s e gm e nt o f th e b o dy c o nt a ins a t ri a ngul ar g a ngli o n f ro m

which fi ne n e rv e fib e r s p ass to a ll p a r ts o f the b o dy Thi s is the .

s o c a ll e d s ympathe ti c ne r vou s s ys te m
- .

T HE SE N S OR Y OR G AN S — T HE E YE S AN D VI SI ON — Each
compou nd e ye i s m a d e up o f a g re a t numb e r o f l o ng Sl e nd e r ,

s t r uctu r e s c a ll e d omma tidi a The re m ay b e a s m a ny a s fi ve

.

tho usand o f th e se in a singl e e ye Th e o mm a tidi a a re all alike

.

in str uctu re Th e y m ay b e r e co gni z e d e xte r n ally a s minute

.

he x ag on al are a s o r face ts a m o ng whi ch a ri se l o ng p ro te ctive h a i rs

,

which a r e unb r a n ch e d P a ssing fro m without in th e o mm a

.
,

ti di u m i s f o und t o c o nt a in th e f o ll ow ing p a r t s : th e cor ne a ; th e

cr ys talli ne cone c o mp o s e d o f f o u r m o difi e d c e ll s s u rr o und e d by

tw o c e lls c o nt a ining c o l or ing m a tt e r a nd th e r ha bd om e a d e li ,

ca t e t ran sp a re nt ro d s u r ro unde d by e ight sl e nde r re ti nu lar

ce lls a nd ab o ut tw e lv e i m nt ll w hi ch e xt e nd to a bas a l
, pg e ce s ,

me mbr a ne . In a ll th e re a re tw e nty e ight p a r t s to e ach -

o mm a ti dium T h e pigm e nt c e ll s p r e v e nt th e r e fle ctio n

o f li ght within th e o mm a tidium a nd b e tw e e n n e ighb o ring
om m a tidi a .

Th e oce lli th ough c o mm onl y kn o wn a s simpl e e ye s a r e alm o st

, ,

as c o mpl e x as th e co mp o und s t r uctu r e s ju s t d e sc r ib e d Ea ch .

o c e llu s c o n s i s ts Of a n e xt r e m e ly co nve x c o r n e a a nd a l a rg e b i
,

c onve x cryst allin e c o ne b e hind whi ch a re a g re a t numb e r of rods

,
2 46 AN INTR ODU CT ION TO Z OOLOGY

r e se mbling som e wh a t th e omm a tidia An o ptic n e r v e fro m the .

b ra in p asse s to e ach oce llu s .

VI SI ON — A numb e r o f int e r e s ting bi o l o gic a l p r obl e ms a r e

.

di r e ctly co n ce r n e d with t h e vi s i o n o f th e h o n e yb e e a nd o th e r
in se cts ; a mo ng the se m ay be m e ntio n e d th e o rigin o f fl o we rs a nd
cro ss — p ollin a ti on th e m e tho d o f finding th e w ay b ack to th e
,

hi ve a nd th e finding o f th e qu e e n by th e d ron e dur ing th e nupti al

,

FI G . 1 38 . Lo ngit u d i na l
ti o n th ro ugh p a r t
se c of an an e t nna o f th e h o ne y
t a ctil e h a i r s ; a u d it o r y p it s ; n ne r v e s ;
'

be e c co no i d h a i r s ; f
.
, , ho .
, ,

3 s m e ll h o ll o w s
, ( Fr o m C h e shi r e )
. .

fli ght The se que s tio n s ca nn o t b e d e finit e ly a n swe r e d b e ca us e

.

th e e x a ct ch a r a ct e r Of th e im ag e p ro duce d by th e e y e s i s no t
.
“
kn o wn A m o difi ca tio n o f th e mo sa ic the o r y p r op o se d by
”
,

M ii lle r in 1 8 2 6 a nd d e s c rib e d f o r th e co mp o und e ye o f th e c r ay

fi sh o n p a g e 2 0 7 i s s till h e ld by m a ny inv e s tig a t o r s
,
M o ve m e nts .

a r e m a d e kn o wn t o th e b e e v e r y qui ckly acco r di ng t o thi s th e o ry

, ,

a nd th e for m o f o bj e ct s whil e th e in s e ct i s m o ving a r e lik e wi se

in sta ntly p e r ce iv e d s in ce th e v a r io u s f a ce t s w o uld b e affe ct e d

,

in s ucce ssion .

Th e r e i s e xp e r im e nta l e vid e n ce th a t th e oce lli e n abl e th e

in se ct t o di s tinguish light fro m d a rkn e ss If im age s a r e for m e d .

by the m the y must b e of o bj e ct s a t a de finite r a ng e since th e

, ,

l e n s i s in cap abl e o f a c co mm o d a ti on F u r th e r m or e th e gr e a t
.
,

co nv e xity o f t h e l e ns m a k e s it p ro b a bl e th a t f o r m ca n b e p e r c e iv e d
24 8 AN INT RODU C TION TO Z OOLOGY

1 3 8 , ha .
) s upp ose d to b e th e g a n s o f h e aring E ach pit
e nd or .

h a s a c o n e a t i t s b a s e c onn e ct e d with a se n so r y c e ll It i s ve ry .

d o ubtful h o w e v e r w h e the r b e e s h a v e a ny se n se o f h e a ring

, , ,

s inc e th e e x a ct fun c ti o n s o f a nt e nna l o r g a n s is no t kn o wn in

a ny c a s e .

Th e s o unds m a d e by b e e s m a y b e e nti r e ly incid e nta l to

o th e r a ctiviti e s So und s r e sul t f r o m th e vib r a ti o n o f th e wings
.

th e vib r a ti o n o f th e a bdo min a l s e gm e nt s a nd t h e a ctivity o f ,

t h e spir a cul a r v o c a l a pp a r a tu s T h e s iz e o f th e wing s a nd

.

physiol ogica l co nditio n o f th e b e e d e t e r min e th e r a te of vib ra

tio n and co n se que ntly th e pitch Whe n in full flight the 440
,
.
,

vi b ra ti o n s p e r s e c o nd giv e a in t h e t r e bl e cl e f but if f a tigu e d , , ,

on l y 3 3 0 vib ra tio n s pe r s e c o nd m a y b e p r o du ce d giving e Th e , .

SO c all e d vocal a ppa r a tus

- li e s with in th e spir acul ar op e ni ngs
o f t h e r e spi ra t or y s y s t e m It co n si s t s o f a vocal m e mb rane
.
,

a s o unding b o x a nd a m e ch a ni s m f o r r e gul a ting th e s iz e o f t he

,

Op e ning Ai r p a ssing t o th e o ut sid e vib r a t e s th e m e mb r an e

.

p ro ducing a humming s o und .

T OU C H B e e s p osse ss a t a ctil e se n se th e e nd o r g an s o f which

.
,

a r e h a i rlik e st r uctu r e s o n v a r i o u s p a r t s o f t h e b o dy but e sp e ci ally ,

o n t h e a nt e nn a At l e a s t t w o kind s o f t actil e o r g a n s o ccu r o n

.

t h e a nt e nn a One o f th e se c o nsi st s o f a s m all h a i r ( Fi g 1 3 8 f)

. .
,

which p r oj e cts th ro ugh a minut e o p e ning in th e ch itin and i s ,

c o nn e ct e d with a n e r ve c e ll (n) within T h e o the r t ouch .

o r g a n s a r e t e r m e d conoi d ha i r s ( F i g 13 8 the y a re l arge r

.
, ,

a nd h a v e a ce nt r a l ca vity co nt a ining a n e r ve fib e r M ore .

t a ctil e h ai r s ar e p r e se nt n e ar th e e nds o f the a nt e nn a than

e ls e wh e r e .

Th e R e pro d u cti ve Sys te m M ALE R E P R ODU C TI V E OR GAN S

.

— bd m n d n o te s te s ( T
( g
F i . I n th e a o e o f t h e r o e a r e t w
e a ch c o n si s ting o f a b o ut th r e e hund r e d sp e r m a ti c tub e s in whi ch

th e Sp e r m a t o zo a ar e for m e d ; th e y a r e co nn e ct e d by a p ai r o f fi ne
tub e s th e vasa def e r e nti a ( V Def ) with th e s e mi nal ve si cle s
,
.
,

T h e s e m in al v e sicl e s Op e n int o a p ai r o f l a rge mu cou s

2 50 AN IN T RODU CTION TO ZOOLOGY

unde v e l op e d fe m al e s T h e l a tt e r h o we ve r co nta in v e stigi al

.
, ,

re p ro du ctiv e or g a n s whi ch m ay e v e n und e r ce r t a in c o ndi ti o n s

, ,

b e co m e cap abl e o f p ro du cing e ggs T h e abdo me n o f the .

que e n i s a lm os t c o mpl e t e ly fill e d by th e tw o ova r i e s ( Fi g 1 4 2 .

,

E ach o v a r y i s m a d e up o f a g re a t numb e r o f ova r i a n tu bu le s ( ov ) .

c o nt aining e ggs o f va ri o us siz e s th e l a rg e s t a t the p os t e ri or e nd

,
.

E gg s p a ss fro m th e s e tubul e s int o th e ovi d u cts (0c the nce .

into the vagi na a nd ou t o f th e b o dy by w a y o f th e g e ni t a l

a p e r tu r e
. Op e ning into th e v a gin a i s a sph e r i ca l s a c th e spe r ,

ma the ca S fi ll e d with sp e r m a to z o a r e c e iv e d f r o m th e m al e
du ring copula tio n .

SP E R M ATOGE N E S I S T h e m a tu ra ti o n o f th e m a l e c e ll s o f the
.

ho n e yb e e di ffe r s m a r k e dly fro m the u su a l typ e (p

-
a nd .

fro m th a t o f th e a n i m al s thu s f ar de sc rib e d T he p rimordi a l .

g e r m ce ll s g ro w into sp e r m a to go ni a a s u s ual T h e fi rs t sp e r .

m a to cyte divi sio n h o we v e r , which o r din a r ily r e s ul t s in two se c

,

o nd a ry sp e r m a t o cyt e s o f e qu a l Siz e i s a so r t Of budding p ro ce ss

,
.

A sm all p o r tio n o f th e cyto pl a s m i s pin ch e d o ff a nd di sint e g ra te s .

T h e c e ll r e m a ining th e se co nd a r y sp e r m a t o cyt e r e t a in s a ll o f
, ,

th e ch ro m a tin o r igin a lly co nt a in e d in th e sp e r m a t o g o nium Th e .

se c o nd ar y sp e r m a t o cyt e no w di vid e s p ro ducing o ne sm a ll ce ll

,

with ha lf o f th e chr o m a tin but v e ry littl e cytopl asm a nd one

, ,

l arge c e ll T h e s m all ce ll b e gin s to d e ve l op into a sp e r ma to zoon

.
,

but p rob abl y d e ge n e ra t e s T h e l a rg e r ce ll whi ch m ay now be

.
,

ca ll e d a sp e r m a tid m e ta morphose s into a singl e functi on al

,

sp e r m a to z oo n ( 20 9 ,

OOGE N E SI S . It is now p r e tty w e ll e s tablishe d th a t the e ggs

which p r o duce d ro n e s are no t fe r tilize d whil e tho se tha t p ro duce ,

th e wo r k e r s a nd qu e e n s a r e T h e r ip e ning o f th e l a tt e r i s si m il ar
.

t o t hi s p r oce s s in o th e r a ni m a l s but th e m a tu r a ti o n o f th e u nf e r
,

tili z e d dr o ne e gg i s unique A full a cco unt of thi s p r o ce ss

.

h a s b e e n publi sh e d by P e t r u nk e wi t ch
C OP U L ATI ON .
-
T h e Sp e r m a t o z oa a r e t ra n s fe rr e d f r o m the
dron e to the que e n whil e the l a tt e r i s t a king h e r nupti a l fli ght .

U sually fro m fi v e t o e ight d a y s a fte r the qu e e n e me rge s fro m he r

 T H E HON E Y B EE AN D B EE S I N G E N ER AL 251

ce ll sh e v e ntur e s ou t o f th e hive fi r st cr a wling ab o ut n e a r th e

, ,

e nt r a nc e the n t aking Sho r t fli ght s a nd fin ally h e r w e dding t r ip

, ,

o f f r o m thr e e t o thi r ty mi nut e s Sh e i s fo ll o w e d by th e d r o n e s

.
,

o ne o f whi ch c op u l a t e s with h e r T h e r e s ult o f copul a tio n i s th e

.

filling of the sp e r m a the ca o f th e que e n with sp e r m a to z oa ( Fi g .

Sh e u s u ally copul a t e s o nly o nc e th e sp e r m o bt a in e d a t ,

tha t tim e b e ing s uffi ci e nt t o fe r tiliz e th o u sa nd s o f e gg s Afte r .

h e r nupti a l flight th e qu e e n n e v e r
l e a ve s the hive e xc e pt w ith a sw ar m .

F E R TI LIZ A TI ON — T h e e gg s a r e f e r
.

ti li z e d ju s t b e fo r e d e p ositi o n Ho w .

thi s i s a cco mpli sh e d i s no t d e finite ly

kn o wn T h e qu e e n s e e m s t o b e abl e
.

to la y f e r tiliz e d o r u nf e r tiliz e d e gg s
a cc o r ding t o th e s iz e o f th e c e ll in
which th e individu a l i s to d e ve l op ,

but it h a s b e e n p r o v e n th a t th e Si z e *

o f th e c e ll d o e s no t a ut o ma tic ally
F G 43 5 p m th f I 1 er a e ca 0
d e t e r mi n e th i s F e r ti l i z e d e gg s d e
.

q n h n yb ue e o e ee . a,
v e lOp in to qu e e n s a nd wo rk e r s p fill d with fl i d ; , S a ce e u

whe r e a s th e u nf e r tili ze d e gg s whi ch b m f p m t ; ,

as s o s er a oz o a

d e v e l o p b e co m e d r o n e s T h e me th o d
, .
d t; 1 tiv p 6, uc ( ,
ac e S er

o i co nt r o l o f f e r tiliz a ti o n is still a
m t ( F m P k a o z oa ' ro aC “

d f t C h hi ) ar a er es re
m ys t e r y
.

E GG LAYI N G — T he e gg s a r e blui sh whit e a nd Ob lo ng in sh ap e ;

.
,

the y are d e p o site d by the qu e e n a t th e b a se o f th e c e ll s a nd f a s

te ne d in a p ositio n p a ra ll e l to th e sid e s o f t h e ce ll s by a glutin o us
s e cre ti on T he fe r tiliz e d e ggs a r e l aid e ithe r in s m all wo rk e r
.

ce ll s ,
o r in l a r g e i rr e gul a r qu e e n ce ll s Unfe r tiliz e d e ggs a re .

u s ua lly l a id in d ron e ce ll s .

E m b ry olo gy — T h e fe r tiliz e d e gg i s m a d e up o f a l a r g e ce nt r a l
.

m ass o f yolk s phe r e s a m o ng which a re t r ace s o f cy toplas m and

, ,

a p e r iph e r a l l a y e r o f cyt o pl a s m A s ingl e nucl e u s 1 5 p r e se nt ;

.

this i s th e nu cl e u s o f th e e gg a nd tha t of th e sp e r m a to z oo n co m
bi ne d . A chitin o u s s h e ll th e chor i on s u r ro und s t h e e gg ; thi s is
, .
2 52 AN I NT R ODU CT ON TO I ZOOLOGY

li n e d by a d e lic a t e vi te lli ne m e mbr a ne As in t he cr a yfi sh cle av .

,

a ge p r o c e e d s with o ut t h e int e r v e nti o n o i c e ll w a lls M ost of the .

cl e a v a g e nu cl e i mig ra t e t o th e p e r iph e r y wh e r e th e y f o r m a s ingl e ,

l ay e r o f ce ll s th e blas tode r m ; th e r e s t r e m a in in th e yolk whi ch it

, ,

i s th e i r duty t o ch a ng e int o p r o t opl a sm a s d e ve l opm e nt co ntinu e s .

T h e bl a s t o d e r m s o o n b e c o m e s t hi ck e r o n one Sid e o f th e e gg ,

fo rming th e ge rm band or p rimi tive stre ak This l a te r b e come s .

FI G . 1 44 . hone yb e e within e ggs h e ll a b hi nd i nt e s ti ne ; c

E m b r yo of . .
,
-
,

oe sop h a ge a l co nne ctiv e s ; ch ch o r i o n or e ggs h e ll ; b r e i nt e s ti ne ;

.
, f f o .
,
-

ga g.a n, gli a ; m b m i d i nt e s ti
. ne,;
-
s ga b r a in ( F r o m P a ck
. a r d a f
. t e
,
r .

C h e s hi r e ) .

th e v e nt r al Si d e o f the y oung b e e N e xt a m e di an gr oove appe ars .

in th e ge rm b an d and two l a ye r s of c e ll s a ri se in thi s r e gion ; the

o ut e r l a y e r i s th e e ctode r m th e inn e r th e e ntom e s ode r m
,
The ,
.

l att e r as its n am e impli e s giv e s r i se to b o th the e ntode rm and

, ,

m e sode r m T h e g e r m b and no w g r o ws a r o und th e e gg until it

.

cove r s th e e nti re s u r fa ce T h e a nte nn a a nd fo u r p ai r s o f app e nd

.

a g e s a pp e a r n e a r th e a nt e r i o r e nd o f th e e mb ry o One p ai r of .

th e l a tt e r di s app e a r ; th e o th e r s b e c o m e th e m o uth p ar t s Thre e .

p ai rs o f app e nda ge s al so d e v e l op on th e tho r a x but di sapp e ar

‘

b e fore th e e mb r yo ha tch e s So m e o f th e o rg an s o f the

e mb r y o a r e Sh o wn in F igu r e 1 44 T h e g a ngli a o f th e b rain .

( g )
s. o a n.d v e nt ra l n v
er e c o r d (g )
a a r e quit e d i s tinct a t thi s tim
.
e ,

a nd th e th r e e p rincip a l p a r t s o f th e a lim e nt a r y ca n a l the for e ,

i nte sti ne mi d i nte sti ne -

a nd hi nd i nte s ti ne ( a b ) o ccupy - .

th e l o ngitudi n a l a xis o f th e b o dy .

M e tam o rph o s i s — T he lif e hi t

s or y o f a n individu a l m a y b e

divide d int o fo ur p e ri o ds e gg la rva pu pa a nd adu lt or i mago

, , , ,
.
FI G . 1 45 . La r v a a nd p u p a o f o ne yb e e i nh e r th i ce ll s . Sl . l a r va sp i nni ng
,

co co o n N , pupa FL , yo u ng a r a l v
an , an . t e nna ; cc e ye co co coo n;
.
, .
,

m , m a nd i b l e ; sp .
, i
sp r a c e s l ; t t o ngue
, ; w, wi ng ( Fro m P a ck a r d a f t e r
.

C h e s hi re .
)

FI G . 1 46 . Int e r na ll ar va l ho ne ybe e an anu s ; bm ne r ve ch a i n ;

o r ga ns o f . .
, .
,

cd . t r u e s to m a ch ; cd hi nd —i nt e s ti ne ; g br a i n ; s t sp ir a cl e s ; oz
, .
, ,
.
, ,

ce so ph a gu s ; s d s p i nni ng gl a nd
.
, vm M a lpi ghi a n t u b u l e ( Fr o m P a ck
.
, .

a r d a f t e r Le u ck a r t ) .
 T HE HON E Y B EE AN D B E E S I N GE N E R AL 2 53

Th e l e ngth of tim e of ea ch s t a te is shown in Ta bl e X I ( 1 9 1 ,

p .

TAB LE X I

THE TI ME OC C UP I E D I N T H E DE VE L OP M E N T OF Q U E E N , WOR K E R , AND DR ON E

AR V A UP A TO TA L FR OM D T
E P OSI I ON
L P
OF E GG To A U LT
D

Aft e rthr e e da ys the l a rva e me rg e s fro m th e e gg a nd li e s a t ,

th e b ase o f th e ce ll ( F i g 1 4 5 F L ) fl oa ting in th e fo o d p r e p are d by

.
,
.

”
th e w or k e r s a nd kn o wn as chyle o r b e e milk C hyl e is co m .

p o se d of di ge st e d ho n e y and p o ll e n p rob ably mi x e d with a ,

g l a ndul a r s e c r e ti o n a nd i,
s giv e n t o a ll o f th e l arva by th e nu rse

b e e s during the fi rs t th re e d a ys The n th e l a rva tha t will be .

c o m e wo rk e rs ar e giv e n h on e y a nd dig e s te d p oll e n in g ra dually

in cre a sing a m o unts ; th e d ro n e l a r va a ft e r th e fo u r th d ay al so , ,

r e c e iv e h o n e y a nd undig e s t e d p o ll e n ; but t h e qu e e n l ar va are f e d

l a vi shly on th e r ich a lbumin o us b e e m ilk th e ro yal j e lly , ,

un t il th e y cha nge to pupa .

GR OWTH du r ing th e l a r va l p e ri o d i s a cco mp a ni e d by s e v e r a l

mou lts o f th e chitin o u s l a r v a l e nv e l op e At th e e nd o f th e l a rv al .

p e ri o d th e c e ll s co ntaining th e yo ung b ro o d a r e cov e r e d o ve r

with wa x fe e di ng c e ase s a nd th e l a rva p r o c e e d to spin a cocoon
, ,
.

Of Silk f r o m th e i r spi nni ng gl a nd s ( F i S T h pi nning

g 451 e s e S .
,

gl a nds ( Fi g 1 4 6 sd ) b e co m e th e saliv a ry gl a nds ( s ys te m s I a nd

.
,
.

II) of th e a dult T h e simpl e str u ctu r e o f th e l arv a i s sho wn in

.

Figur e 1 4 6 . T h e a lim e nt a r y ca n a l co nsi s t s o f a n ( e s opha gus

( ) a chyle s tomach
a , a hi nd i nte s ti ne a nd t w o se t s o f
-

a pp e nd a g e s th e s i nni n l nd n d h M l i h i n tubul s
, p gg a s a t e a pg a e

Alm o st e v e ry se gm e nt c ont ain s a p a i r o f spi r acle s

and a gangli on o f the c e nt r al n e r v o us syst e m
2 54 AN IN T R ODU CTION TO Z OOLOGY

It t ak es w k thi ty i h ou r s to spi n its co coo n the n it

th e or er r -s x
,

sl o wly ch a ng e s int o a pup a or C h ry s a lis ( F i g 1 4 5 N)

,
P r a cti .
,
.

ca lly t h e e nti r e b o dy i s m a d e o v e r a t thi s tim e ; th e thr e e re gi o n s ,

h e a d tho ra x a nd abdo m e n b e co me di s tin ct ; e xt e rn ally th e wi ngs

, , ,

( )
w ,
l e g s m o
,
uth p a r t s ( t m ) s ting a nt e nn a
,
a nd ,e y e s a r e ,

vi sibl e ; and th e inte rn a l cha ng e s a r e e v e n m o r e s tr iki ng the ,

l a r v al o r gan s de ve l oping into those o f th e a dul t a nd ne w o rg a n s . ,

a pp e a ring Aft e r a p e r i o d o f re s t th e pup a c a s t s Off its e xosk e le

.

t o n a nd e m e rg e s a s a n a d u lt
,
.

Th e Acti vi ti e s o f th e Wor k e r s — T h e functi o n s o f th e qu e e n .

a nd d r o n e a re f e w as c o mp a r e d with th o s e o f th e w or k e r The .

q u e e n l a ys e ggs a n d t h e dr one f e r tiliz e s th e qu e e n

,
B u t t h e .

wor ke r s h a v e a la rg e numb e r o f v a r i e d a ctiviti e s su ch a s th e build ,

”
ing o f h o n e y co mb th e coll e cti o n o f p rop olis o r b e e glue
,

w i t h whi ch th e in sid e o f th e hi v e i s v a r ni s h e d th e g a th e r ing o f ,

”
p oll e n or b e e b r e a d a nd i ts p r e p a r a tio n a s f oo d th e fe e di ng ,

o f th e qu e e n a nd y o ung b e e s t h e ca r r ying o f w a t e r th e co lle c , ,

ti on o f fl ow e r n e cta r and its m a nuf acture int o ho n e y and the ,

cl e a ning war mi ng v e ntil a ting and gu a r ding of th e hi ve Work

, , ,
.

e r s a ls o a cc o mpli sh t h e p o llina ti o n o f fl o w e r s r ais e ne w qu e e ns ,

wh e n ne c e ss a r y a nd i n cre as e th e numb e r o f col oni e s by s warming

,
.

THE B U I LDI N G OF H ON E Y C OMB — Wax i s p r o duce d by p ai r s .

o f w a x gl a nd s o n th e s t e r n a o f th e l a st fo u r visibl e a bd omi n al

s e gm e nts of the w or ke r Hon e y and p oll e n a re consum e d in th e

.

p ro c e ss and a te mp e r a tur e of fr o m 9 7 to 98 F i s ma int ain e d

,
° °
. .

Whe n co mb i s to b e built th e b e e s g or g e the m se lve s with hon e y

, ,

a nd h a ng in d e n se clu s t e r s f ro m t h e t op o f t he hiv e f or s e v e r a l

ho ur s Thin s cal e s o f w a x a re the n se cr e t e d by the w ax gl ands

.
,

re m o v e d by th e w a x pinch e r s o n th e m e t a th o r a ci c l e gs tran s ‘

fe rre d to th e p ro tho r a ci c l e g s a nd the n to th e m o uth whe r e th e y, ,

a r e mix e d with sa liv a a nd kn e a d e d by th e j a w s If a ne w comb .

i s to b e b uil t th e w a x i s pl a s t e re d t o th e roo f a nd in s o m e
, ,

m y s t e r i o us w a y e a ch b e e puts i t s co nt r ibutio n a lmo st e x a ctly

wh e r e it i s t o r e ma in T h e c e ll s whi ch a r e gra d u ally built up
.
.
, ,

a re h e x a go na l in Sh a pe .
256 AN IN TR ODU CTION TO Z OOLOGY

hone y ce lls , but al so in d ron e worke r a nd tra n sitio n ce lls C are

, ,
.

f ul m e a s ur e m e nt s h a v e s h o wn th a t t he c e ll s a r e s e ldo m p e r fe ctly
symm e tri ca l a ltho ugh in m any ca se s the y app e ar so t o our e y e s
, .

T h e h o n e y ce ll s a r e built with e nt r a n ce s sli ghtly a b ove th e i r b a s e s ,

s o th a t t h e h o n e y s t ore d in th e m will no t flo w ou t b e f or e it b e
”
c om e s r ip e
T HE C OLLE C TI ON OF P RO P OLI S B e e glu e a s p r op oli s i s
.
,

s om e tim e s c all e d i s a r e sin o u s m a t e ri a l co ll e ct e d fr o m bud s

,

a nd c r e vic e s o f t r e e s It i s t r a n sp or t e d in th e p oll e n b ask e ts and

.
,

is u se d a s soo n a s coll e ct e d to p a int the in side o f the hi ve to

, , ,

fill up cr a ck s a nd to s t re ngth e n a ny l o ose p a r t s

, .

GATH E RI N G P OLLE N — P o ll e n g r a in s a r e of in e stim abl e v alu e

.

in th e b e e h o us e h old Th e y a r e v e r y sm all o f v a ri o u s Sh ap e s and

.
,

c ol or s a nd a r e fo r me d within a p ar t o f th e fl ow e r kn own a s th e
, ,

a nth e r . T h e p o ll e n g r a ins co nt a in th e m a l e e l e m e nt in th e fe r tili

z a ti o n o f fl o w e r s a nd co n s e qu e ntly a r e n e ce ssa r y f o r th e p r o duc
,

ti on o f fe r til e s e e d T o th e b e e p oll e n i s inva lu abl e a s a food

'

,
.
,

a nd i s a l so u se d in p re p a r i ng t h e ce ll s c o nt a ining pup a We .

h a ve al r e a dy d e scr ib e d th e p e culi ar st r u ctu re s on th e l e g s and

o th e r p a r ts o f t h e b e e s b o dy u se d in c o ll e cting p o ll e n ( p
’
.

Up o n r e aching th e hiv e th e p e ll e t s o f p oll e n a r e p ri e d ou t o f th e

p oll e n b as ke t by th e spu r a t th e t e r min a ti o n o f th e tibi a o f th e
m iddl e le g ( Fi g 1 3 1 D s) a nd d e p osit e d o ut s id e th e b r oo d
.
, , ,

clu s te r s in wh a te ve r ce lls a r e a va il abl e u su ally in w o rk e r ce lls .

P o ll e n i s the p r in cip a l foo d o f the l a r va It i s v e ry r ich in .

nitr oge no u s m a te ri al a foo d e l e m e nt no t fo und in hon e y and

, ,

with out whi ch th e yo ung w o uld s t a rv e T h e g a the ring o f p oll e n .

by b e e s h a s a g r e a t influ e nce up o n th e fl o we r s vi sit e d a s i s e x ,

pl ain e d in a n o the r pl a c e (p .

C AR R YI N G WATE R D u r ing w ar m w e a th e r w a t e r i s suck e d up

.

into th e h on e y sac fro m d e w o r b ro ok s a nd p o o l s a nd c arri e d

, ,

t o th e l ar va in th e hiv e In coo l w e a th e r m oi s tu re co nde n se s

.

in th e hiv e in suffi ci e nt qu antiti e s a nd und e r so m e co nditio n s , , ,

t o s uch a n e xt e nt a s t o inju r e t h e inh abit a nt s .

T HE M AN U F A C T UR E OF HON E Y B e e s d o no t c oll e ct h o n e y fr om
.
 T HE HON E Y B EE AN D B E ES I N G E N ER AL 257

fl o we rs but g a the r n e cta r which i s l a t e r t ra n s fo r m e d int o h o n e y

, , .

T he n e ct a r i s l a pp e d up by th e t o ngu e ( F i g 1 2 9 l) a nd t r a n s .
, ,

fe rre d to the ho ne y sa c ( Fi g 1 3 4 wh e r e it i s s tore d whil e th e

.
,

be e is in th e fi e ld P a r t o f th e w a t e r co nt ain e d in t h e n e ct a r
.

m a y b e e xc r e t e d b e fo r e th e hive i s re a ch e d N e ct a r i s pl ace d .

in Op e n ce ll s in the we ll ve ntil a t e d hi ve until all but 1 8 to 20 pe r

-

c e nt o f the wa te r co nt a in e d in it h a s e va p o ra t e d Whe n a ce ll .

is fin ally fille d with r ip e hon e y it is se a l e d with a ca p o f w a x , .

T h e fl a vo r o f h o n e y d e p e nd s up o n th e kind o f fl o we r s f ro m
which th e n e cta r is coll e cte d Am o ng th e m o s t imp o r t a nt
.

p ro duc e r s o f the b e s t h o ne y in th e E a s t a nd N o r th a re white cl ove r ,

b a sswoo d bu ckwh e a t a nd th e f r uit t re e s a nd s m a ll f ruits ; in

,

the m iddl e s t a t e s a r e th e tulip t r e e s o r r e l t re e s w e e t cl o v e r a nd , , ,

a lfa lfa ; in the So uth a re th e m ang rove c abb age a nd sa w ,

p a lm e tt os a nd s o r r e l t r e e ; whil e in th e We s t a re a lfa lf a a nd whi t e

,

s age T he b e st and m o s t o f the C aliforni a h o n e y is f r o m th e

.

wild whit e s a g e ( 2 0 7 p ,
T h e a m o unt o f h o n e y p ro du ce d
.

in o ne hive in a fa i r s e a so n ra nge s fro m a n a ve r ag e o f a b o ut

thi r ty p o unds o f c o mb ho ne y t o p o s s ibly fifty p o unds o f e x
t ra ct e d h o n e y Thi s will ne t th e b e e k e e p e r fro m te n to fift e e n
.

c e nts pe r p o und
C LE AN I N G THE HI V E T h e h e a lth o f th e swa r m d e p e nds up o n
.
-

the cl e a nlin e s s o f th e i r d o micil e s in ce p e r f e ct sa nit a r y co nditi o ns

,

a re n e c e ss a r y wh e re s o m a ny in di vi du a l s live in s uch cl o se qu ar

t e rs De a d b e e s pie c e s o f Old c o mb th e e xcre t a of the qu e e n

.
, , ,

d ro ne s a nd o th e r s tha t r e m a in in th e hi ve a nd any o the r wa ste

, ,

m a te ri a l i s imm e dia t e ly r e m o ve d .

VE N TI LATI N G THE HI V E F re sh a i r f or th e hive is o bt a in e d

.

by the e xe r tio ns o f ce r t ain o f th e wo r ke rs M a ny b e e s n e a r the .

e nt r a nc e a nd a t o th e r pl a c e s in th e hiv e
, a re bu s ily e ng a g e d in ,

vib r a ting th e ir wing s a nd c re a ting a cu rre nt o f a i r which k e e ps

, ,

the hiv e fr e s h a nd a id s in r ip e ning th e n e ct a r

, T h e l o ud buzzing .

which a cc omp a nie s this a cti vi ty i s o fte n h e a r d a t night a fte r a

l a rge a m o unt of n e ct a r h a s b e e n co ll e ct e d
GU AR DI N G TH E HI V E T h e hive i s gu a r d e d a ga inst th e int r a
.

s
258 AN INT ROD U CTION TO Z OOLOGY

sio ns o f ye ll ow j a ck e ts b e e m o th s a nd o the r b e e s by worke rs

, , ,

wh o w a nd e r b a ck a nd f or th n e a r th e e nt r a n ce a nd e x a min e e v e ry ,

c r e a tur e tha t vi sits th e c ol o ny If th e s wa r m i s s t rong th e gua rds

.
,

s u cc e e d with th e a i d o f t h e b e e k e e p e r in w a r di ng o ff a ll h o n e y
, ,

l o ving e n e mi e s .

SWAR M I N G — T h e numb e r o f b e e s in a hiv e inc re a s e s v e ry

.

r a pidly s inc e th e qu e e n u s u a lly l a y s f r o m 9 5 0 t o 1 2 0 0 e gg s

, pe r
d ay . Whe n th e colo ny i s in a p r o spe r o us conditio n a nd th e re ,

is da ng e r of o ve r c r o wdi ng que e n c e ll s a re built by th e wo r ke r s

, ,

usually aro und th e f e r tiliz e d e ggs and ne w que e ns a re r e a r e d ,

.

I wo qu e e n s d o no t live a mi cably in o ne hi v e a nd if s uch a co n

‘ ‘

, ,

mli ti on a ri se s e ith e r th e re i s a b a ttl e b e tw e e n th e tw o r e s ulting in

, ,

t he d e a th o f o ne o f th e m o r th e w o r k e r s kill o ne o r e l se th e o ld
, ,

q ue e n co ll e cts f ro m tw o to tw e nty tho usa nd wo rke rs ab o ut he r

a nd fli e s a w a y with th e m t o f o und a ne w co l o ny T hi s is kn o wn .

a s sw a r m ing T h e Old hi ve is no t b r o k e n up b ut c o ntinue s its

.
,

e xi s t e nc e as b e fo r e .

Swa r ming o ccu r s in M ay Jun e o r July a cc o rding to l a titude

, , , ,

a nd a se co nd s w a r ming p e r i o d m a y b e in a ugu r a t e d if w e a th e r

c o nditio n s r e sult in a mids umm e r flo w o f h o n e y B e fo re issuing .

f ro m th e hi ve th e h o ne y sacs a r e fille d with ho ne y t o s e rv e until

,

a ne w h o m e i s f o und T h e s w a r m a ft e r fl ying a sh o r t di s t a nce

.
, ,

c o m e s to r e s t up o n the limb o f a t r e e o r o the r obj e ct whe re it

r e m a ins so m e tim e s f o r s e v e r a l h o u r s A sit e f o r th e ne w col o ny
.

is s o m e time s ch os e n by s co uting b e e s se ve ra l d ays b e fore the

swar m l e ave s the p are nt hive The se sco uts m a y a l so p a r tially
.

p re p are th e pl ace by cl e a ni ng o u t l oose dir t b a r k e tc T he , ,

.

u s u al choi ce i s a holl ow t re e such a s th e wild a nce st ors o f th e

,
”
h on e yb e e inh abit e d a nd he n ce for th i s call e d a b e e t re e
,
One .

o f t h e duti e s o f th e b e e k e e p e r i s t o hiv e th e s w a r ms b e fo r e th e y

succ e e d in e scaping to th e woo ds Swa r ms m ay a l so be forme d .

a r tifici a lly .

Th e E n e m i e s o f th e H o n e yb e e — We ak o r n e gl e ct e d hive s
.

m a y b e a tt a ck e d by th e B e e M oth Ga lle r i a me llone lla which , ,

t ak e s adva nt ag e o f e ve r y o pp o rtunity to e nte r a nd lay i ts e ggs .

2 60 AN I NT R ODU C T ION TO Z OOLOGY

2 . BEES IN G E N E R AL

a . C lassi fica ti on f
o H one ybe e s

T h e b e e s b e l o ng with th e a nt s w asps e t c t o th e or de r Hy
, , ,
.
,
o

m e nopte r a In this or d e r a r e i nclude d all in se ct s with fo u r

.

m e mb r a n o u s wing s th e hind wing s b e ing th e s m all e r ; with

,

biting a nd s u ck ing m o uth p a r ts ; with a s ting pi e rce r or saw a t , ,

t h e e nd o f t h e a bd o m e n o f th e f e m al e ; a nd with a c o mpl e t e
m e tam orpho s is i e with l arva l a nd pup al s t age s dur ing d e ve l op
,
. .

m e nt T h e hone yb e e b e l o ng s in th e f a mily Apida and i s th e

.
,

m ost sp e ci aliz e d with re g ar d to its co mmuni s tic life o f a ny of th e

g ro up Th e sp e cie s o f h o ne yb e e fou nd in thi s c o unt ry i s Api
. s

m e lli fica A numb e r o f o th e r sp e ci e s Of h o n e yb e e s inh abiting

.

Asi a a nd Afr ica a re pl a c e d with m e llifi ca in th e ge nu s Api s .

T he in di vidu a l s Of th e Sp e ci e s Api s m e llifi ca a r e no t a ll a lik e in

str uctur e c ol o r o r a ctiviti e s Se ve n or m o r e r a ce s a r e r e c ogniz e d
, ,
. .

T h e ch aract e r i s tic s o f th e m o re imp o r t a nt ra c e s a r e c ont ra st e d

in Tabl e XII T h e r e l a ti o ns o f the h o n e yb e e s t o o th e r ins e cts
.

a nd t o e a ch o th e r a r e sh o wn in o utlin e in T a bl e XIII .

b Gyna nd r om or phs
.

A n orm al col o ny o f h o n e yb e e s c ont ains as st a te d b e fore a , ,

fe r tiliz e d e gg l a ying qu e e n a numb e r o f m al e s or d ro n e s a nd

,
-
, ,

tho us a nds o f st e ril e fe mal e s o r w o r ke r s A numb e r o f b e e s h a v e .

b e e n di s cov e re d whi ch show e d m al e char a ct e rs in c e r t a in p ar ts

o f th e b o dy a nd f e m a l e ch a r a ct e r s in o th e r p a r t s Abn or m a l .

inse cts of this kind a r e kn o wn a s gyn a ndro m o rphs B utt e r flie s .

,

a nts ,
a nd b e e s a pp e ar t o b e m o r e o ft e n a ff li ct e d th a n o th e r

ins e cts T he b e st w e ll kn o wn inst a nce o f gyn a ndr om o rphi sm

.
-

o ccu r r e d i n a hive o f b e e s a t E u gs t e r a nd w a s r e p o r t e d by v o n

Si e b olt This hi v e c o nt a in e d a n Ita li a n que e n a nd Ge rm an

.

d ron e s T h e w o r ke r s p ro duce d by this que e n w e re the r e fo r e

.

hyb rid s So m e o f th e gynand ro m or phs in thi s c ol o ny h ad th e

.

a nt e r i o r e nd o f t h e b o dy m al e th e p o st e rio r fe m al e ; o the rs,

e xhibit e d m a l e ch a r a ct e rs o n th e r ight a nd f e m a l e o n th e l e ft ,
 T H E HON E YBEE AN D B EE S I N G E N ER AL 26 1

T AB LE X I I

S OME OF T H E C H AR AC T E R I S T I C S OF T H E M OR E I M P ORT AN T R AC E S OF
l
H ON E Y B E E S

L R
C O O OF DI SP OS I Q U AL T
I Y C A
P P I N GS
R A CE
AB DOME N T ON
I
'

As AP O
R OF C OM B RE MA RK S
DU C E R HON E Y

Fi rst r a ce i nt r o d uce d
Ge rm an B l a ck C r os s Whi t e
In o t Am e ri ca

I t li
a an G e ntl e M os t po p u l ar ra ce

So m e v o ca t e s
ad i n the
C ar m o la n Gr ay G e ntl e GOOd Wh i t e
U nit e d St a t e s
R e ce ntl y i nt r o d u ce d
re
C aucas I a n Goo d Wh l te
'

Go o d f or am a te urs

B a na t Go o d Whit e R e ce nt

b
'

H t 11

1 t
F ro m i nf o r m a io n f u r nis h e d by Dr E F P h i i ps
. . . ll .

T ABLE XIII
TH E R E LATI ON S OF T HE H ON E YB E E S T O OT HE R I NSE CT S AN D T o E AC H
OT H E R

Ins e cta
ORDE R S Hym e no pt e r a
( Be e s Was ps Ant s
.
, ,
e tc .
)

F A M IL I E S

G E N ERA

SP E C I E S Ap i s i nd i ca Api s m e lli fi ca
( E as t I nd i an Ho ne ybe e ) ( C o m m o n H o n e yb e e )

R AC E S C a r ni o l a ns I t a li a ns G e r m a ns
262 AN INT RODU CTION TO Z OOLOGY

or vi ce ve r s a ; s till
the r s h ad m al e and fe m al e chara ct e rs in
o

di f fe r e nt p ar ts o f th e sa m e o rg an T h e r e p r o ductive or g a ns .

w e r e o ft e n p ar tly m a l e a nd p ar tly fe m al e a nd the i r ch ara ct e r ,

coul d no t b e d e te rmi ne d by the e xt e r n al app e ar ance of the

gyn andr o m orphs Vari o u s e xpl an a tio n s h ave b e e n o ff e re d to
.

a c co unt f or thi s p e culia r c o nditi o n but as ye t th e d a t a n e c e ssa ry ,

t o d e cid e th e qu e s ti o n h a v e no t b e e n fu r ni s h e d .

c . The Re la ti ons o
f B e e s to Othe r Or ga ni sms

C h ar l e s Darwin in T he Origin use d the bum of Sp e ci e s h as

ble b e e to illu s tr a t e h ow pl ants a nd a ni m a l s r e mo te in the sca l e ,
”
o f n a tu r e a r e b o und t oge th e r by a w e b of c o mpl e x re l a tio n s
,
.

He fo und th a t th e vi sits o f b e e s a r e n e c e ss ary f or th e fe r tiliz a

tion o f s o me ki nds o f clo ve r ; fo r in st a nce twe nty he a ds of Dutch ,

cl o ve r ( Tr ifoli u m r e pe ns) yi e ld e d 2 2 90 s e e ds but tw e nty o the r ,

h e a ds p r o t e ct e d fro m b e e s p ro duc e d no t o ne
, ,
Humbl e .

b e e s al on e vi sit r e d cl ove r a s o the r b e e s c a nno t re a ch the n e ctar

, ,

he nce w e m ay infe r a s hi ghly p r ob abl e tha t if th e who le

-
, ,

ge nus o f humbl e b e e s b e ca me e xtinct o r v e ry r a re in E ngl a nd the ,

h e ar t se a se a nd r e d cl o ve r wo uld b e com e v e r y r are o r wholly

’
,

di sapp e a r The numb e r of humbl e b e e s in a ny dis t ri ct de pe nds

.

in a g re a t m e asu r e up o n th e numb e r o f fi e ld mi ce which de stro y ,

the i r com b s and n e st s N o w th e numb e r o f mic e is l a rg e ly

.

de p e nde nt as e v e ry o ne kn o ws o n th e numb e r o f ca ts
.
,
.

He n ce it i s quit e c r e dibl e th a t t h e p r e se nc e o f a fe lin e a nim a l in

l arg e numb e r s in a di st r ict might de te r min e thro ugh the inte r ,

v e nti o n fi r s t o f mic e a nd th e n o f b e e s th e f r e qu e ncy o f c e r t a in ,

”
fl o we r s in tha t d is t rict " ( 2 3 6 p T h e influe n ce o f o ld
,
.

m a ids up o n the numb e r o f c a t s w as sugge st e d by Huxl e y a s an

a d di ti o n to Da r win s illu s t r a ti o n
’
.

B e e s a nd Flowe r s C R OS S-P OLLI N ATI ON

. B e e s in flying from .

fl owe r to flo we r ga the r ing n e cta r a nd p o ll e n a cco mpli sh wha t is

kn own as c ro ss p ollin a ti o n i a the p o ll e n fro m o ne fl owe r is
-
,
. .

c arr i e d by th e b e e to a n o the r fl ow e r C r oss p ollin a ti o n se e ms to .

-

be o f a dva nt a ge t o th e s e e d sinc e m a ny fl o w e r s a r e st r uctur ally

,
FI G . 1 48 . lli na ti on o f a n o r chi d ( Cypr i pe di u m) by a bu m bl e be e A be e forcing i ts
Po .
,

w a y i nto th e fl o w e r ; B o b t a i ni ng ne ct a r withi n th e fl o w e r ; C be e e sca p i ng br u sh e s

, ,

p o ll e n u p o n t h e s ti gm a o f t h e fl o w e r ; b e f o re fi na ll y e sca i
p gn th e b e e r e ce iv e s a no the r

lo a d o f po ll e n f r o m th e a nth e r ( Fr o m C o u lt e r a f t e r Gi bson )
. .
 T HE HON E Y B EE AN D B E ES I N GEN ERAL 2 63

constitute d so a s t o p r e v e nt s e lf p ollin a tio n and the visit atio ns

-

of un s u it abl e i n se cts . On th e o th e r h a nd th e y a r e so for m e d a s

,

to s e cur e vi sits f r o m in se ct s th a t fly r a pi dly a nd e nt e r m a ny ,

flo we rs thus insu r ing the wid e di s t r ibution o f p oll e n T h e b e e s

,
.

are a m o ng th e m o s t imp or t a nt o f th e p ollinizing in se cts The y .

a r e e sp e ci ally v a lu a bl e n e a r fr uit t r e e s sinc e it h a s b e e n d e m o n

,

s t r a t e d th a t o r ch a r d s co nt ai ni ng c o l o ni e s o f b e e s ar e m o r e p r o

d u cti v e tha n n e ighb o r ing o r ch a r d s w ith o ut b e e s .

So m e of th e m o s t int e r e s ting a rr a ng e m e nt s f or th e s e cu ring o f

c ross p ollin a tio n ar e fo und a m ong th e fl o w e r s o f the or chid s
-
.

Fo r e x a mpl e th e co mm o n l a dy sli pp e r (C ypr i pe di u m) is a d apt e d

,
-

t o the visit a ti o ns o f bumbl e b e e s T h e fl o w e r h a s a c on spi cu o u s

.

p o uch with an op e ning o n th e upp e r Sid e of th e inn e r e nd o ve r ,

whi ch ha ngs a flap p o s se s sing tw o a nthe r s and a s tigm a Th e .

b e e fo r c e s i ts w ay int o th e p ouch ( Fi g 1 4 8 A) a nd suck s up th e

.
,

n e ct ar co nta in e d within ( B ) In e s caping fr o m the p o uch i t

.

b r ushe s its b a ck fi r s t a g ain s t th e stigm a ( C ) and the n ag a in s t the

a nth e r ( D) Any p oll e n p re s e nt up o n th e b e e s b a ck is b r u sh e d
’
.

o ff up o n t h e s tigm a du r ing i t s e s c ap e a nd a ne w supply i s th e n

,

ga th e r e d fro m the a nthe r Th e n e xt o r chi d visite d r e ce ive s this

.

p oll e n up o n i ts s tigm a a nd a dd s a ne w bu r de n to th e b e e s l oa d
,
’
.

T h e bumbl e b e e thu s i s o blig e d to t r a n s fe r p oll e n f ro m o ne pl a nt

t o a n o the r whil e g a th e r ing n e ct a r
T HE C OLOR S OF F L OWE R S . M a ny s ci e nti s t s b e li e ve tha t the
b rilli a nt col or s o f m a ny fl o we r s a tt r a ct b e e s and o the r in se ct s ,

a nd a r e th e r e f o r e in s t r um e nt a l in ca u s i ng cr o ss p o llin a ti o n
- It i s .

fu r the r cl aime d tha t the b r ight co l o r s the m se lve s a r e th e r e sult

o f t h e vi sit s o f in s e ct s s in ce th ose fl o w e r s th a t h a pp e n e d t o b e
,

m o re b r ightly c ol o re d wo uld b e m o re ce r ta in to a tt r a ct in se ct s ,

a nd th e r e fo r e m o re li abl e to b e p o llin a t e d a nd p r o duc e se e d T he .

s e l e ctio n o f th e m o re b r ightly co l o r e d fl o we r s f or a s uffi cie nt

numb e r o f ye a rs w o uld r e s ult in the sur viv al o f pl ant s whi ch
t e nd to p ro duce m or e highly co l o r e d fl o we r s Thi s e nti re the o ry .

o f th e o r igin o f t h e co l o r s o f fl o w e rs b e c a u se o f th e vi sit s o f i n

s e ct s s e e m s to d e p e nd up o n the fa ct o r tha t a tt ra cts the ins e ct to

2 64 AN INTR ODU CTION TO ZOOLOGY

th e fl ow e r . C e r t ain
b se r v a tio ns app are ntly p rove th a t sm e ll
o

a nd no t col or i s th e d o min a nt f a ct or whe r e as o the r ob

s e rv a ti o ns h a v e r e s ult e d in th e c o nclu si o n th a t ins e cts such as ,

b e e s a nd butt e r flie s tha t Sho w a high de g re e o f a dapt a ti o n to

,

flowe r s p re fe r r e d ; purpl e and blue a nd tha t ins e ct s p oorly

, , ,

a d apt e d t o fl o w e r s f a v o r y e ll o w a nd whi t e P e r h ap s t h e
s a fe s t vi e w to a dopt a t p re s e nt i s tha t co l or o d or a nd st r uctu ral , ,

cha ra cte r s a re a ll imp or t ant fa ct or s influ e ncing the vi sits o f b e e s

a nd o th e r in se cts to fl o w e r s ( 2 04 ,

d . The S oci al Li fe o f Bees

C e r t ain sp e cie s o f ant s b e e s and w a sps e xhi bit a s in the ca se

, , ,

o f the hon e yb e e r e ma r k abl e so ci al o r g aniz a ti o n s

,
Ho w this .

h a s c o m e a b o ut i s a p r obl e m no t ye t s o lv e d but p r a ctic a lly a ll ,

st age s fro m a s o lit ar y h abit to a c ompl e x co mmunity a re illu s

, ,

tra t e d by v ar i o u s m e mb e r s o f th e f a mily Apida ( 1 99 ,

— T h e l e a f cutt e r M e gachi le acu ta is

( 1 ) A SOLI TAR Y B E E .
-
, ,

a s olit a r y l o ng t o ngu e d b e e
,
- In bui ldi ng h e r n e s t Sh e digs a
.

tunn e l u su ally in d e ca ye d woo d a nd e x ca va te s thimbl e shap e d

, ,
-

c avitie s in th e b o tt o m o f it The se cavi tie s a re line d with pi e ce s

.

o f l e a v e s g e n e r a lly cu t f r o m a r ose bu s h
,
In the b o tto m the .
,

b e e pl a c e s a qu a ntity o f p o ll e n a nd n e ct a r up o n which Sh e l ays ,

a n e gg . She th e n plug s th e e nt r a nc e with pi e c e s o f l e a ve s a nd ,

fli e s a way The young th a t h a tch f ro m h e r e gg s live up on the

.

sto r e d foo d .

( )
2 A S OLI T A R Y B E E T H AT W AT C H E S I T S Y OU N G — T h e car .

e n t b e e C r a ti na du pla m a k e s h e r n e st by digging th e pith

p e r e , ,

fr o m th e ce nt e r of a de a d twig o f suma ch or o the r pl a nt Afte r .

a l o ng tunn e l i s e xc a v a t e d sh e b e gin s a t t h e b o tt o m a nd c o n s t r ucts

a s e ri e s o f ch a mb e r s with p a r titi o ns c o mp o se d o f pith At th e .

b o tto m of e a ch ch a mb e r Sh e pl a ce s a m ass o f p oll e n and l ays

a n e gg . Sh e th e n w a its f or h e r o ff sp ring t o e m e r ge
“ The .

l o we r o ne h atch e s fi r s t ; and a ft e r it h as a tt a in e d i ts gr owth it

, ,

t e a r s do wn th e p ar titio n a b ove it a nd the n w a it s p a tie ntly f or ,

th e one ab o v e t o d o t h e s a m e F in ally a ft e r th e l a st o ne in the

.
,
2 66 AN INTR ODU C TION TO ZOOLOGY

( 4) SOLI TAR Y B E E S WI TH T AE N DE N C Y T OW A R D C OM M UN I TY
LI F E .
— T h e mining b e e s o f th e g e nus Hali ctu s m a k e bu rr o ws in
s a nd b a nk s a nd t h e s id e s o f cl iff s ( Fi g 1 4 9 A) . A r e m a r k abl e
,
.

f e a tur e in th e h abit s o f th e b e e s o f thi s g e nu s i s tha t se ve r al

f e m al e s unite in m a king a bu rro w into a b ank a fte r which e a ch .

fe m a l e m a k e s p assag e s e xt e nding Side wi se f ro m this m ain b urro w

o r public co rr id o r t o h e r o w n ce ll s Wh il e A ndr e na buil ds vil
.

l ag e s c o mp ose d o f individua l ho m e s H a li ctu s m ak e s citie s co m

'

p ose d o f ap a r tm e nt h o u se s ( 1 99 p ,
.

— umbl b s
5 ) S O C I A L B E E S WI T H A N N U A L C o L ON I E s T h e h . e e e

o f t h e g e nu s B ombu s f o r m co mmuniti e s co mp o se d o f qu e e n s m a l e s , ,

a nd w o r k e r s. Only y o ung f e r ti liz e d qu e e n s s u r viv e th e wint e r .

In the sp ring e ach se l e cts a n o ld de se r te d fi e ld m o u se n e s t a nd -

st a r t s a co l o ny F i rs t a numb e r o f w or k e r s a re re a re d b e ing
.
,

c a r e d f o r by the i r m o the r ; th e se w o r ke r s th e n t a ke o v e r the

ho use hold dutie s a nd the qu e e n de vo te s h e rse lf to l a ying e gg s
, .

As a utumn app r o a ch e s m a l e s a nd qu e e n s a pp e a r a nd fin ally a ll ,

pe ri sh e xce pt th e y o ung que e n s .

—
( ) SO
6 C I A L B E E S WI TH P E R MAN E N T C O LON I AL L I F E T he .

h on e yb e e diff e r s fr o m the bumbl e b e e in m a ny wa ys I ts life ac .

t i vi ti e s a re m o r e c o mpl e x a nd i t s c ol o ni e s a r e a bl e t o p a s s th e
,

wint e r witho ut p e r i shing Tha t the ir compl e x co mmunity life

.

h ad e v olv e d f ro m a so lit a r y co nditi o n th ro ugh t h e st a ge s m e n

ti one d a b o v e s h o uld no t b e und e r s t ood but th e life hi s t o r i e s of
,

t h e s o lit ar y b e e s a nd th e bumbl e b e e Sh o w m a ny gr a d a ti ons be

twe e n the s t rictly s olit ary li fe and the c ompl e x s o ci a l live s of
th e se re m a r k abl e ins e cts .
PI G . 1 50 . Ar i s to tl e , 3 84-3 22 (F rom Locy )
.
2 68 AN IN TRODU CTION TO ZOOLOGY

a nim al s Th e y r e ve al a r e m ark abl e fa mili arity with th e fact s o f

.

co mp a r a tiv e a n a t o my phy si o l ogy a nd e mb r y o l o gy Ar is t otl e s

’
.
, ,

ide as l a t e r fur ni she d th e st a r ting p oint f or th e fo unding of ou r

m ode r n syste m s of cl a ssific a tio n a nd th e o r i e s of e volution but ,

his gre ate st cont r ibuti o ns to z ool ogy w e r e th e m e tho ds of w or k

which h e int r oduc e d He wa s a cr itical c o mpil e r a nd fr o m the
.
, ,

f ab ric o f scatt e r e d fa ct s and fa nci e s which e xi st e d a t hi s tim e ,

p ro duce d a c o mp act a nd fa i rly a ccur a t e a c co unt of a ni m al s He .

w a s no t co nt e nt h o w e v e r t o a cc e pt o ld s t a t e m e nt s but v e r ifi e d
, , ,

e v e r y thi ng by c a r e ful e x a min a ti o n s o f th e a nim a ls th e ms e lve s ,

and a dd e d m a ny ne w f a cts .

2 . T HE R OM AN P E RI OD

P liny ( 2379
-
A D . ctiv
.
) e public
le d lif e und
an a e r t h e R o m a n
Empi r e a s a n av a l c o mm a nd e r Hi s w r itings co n sist o f thir ty
.

s e v e n v o lum e s whi ch h a d a g r e a t influ e nc e o n t h e id e a s o f n a tu

,

r a li s t s du r ing suc ce e ding c e ntu r i e s Unfor tun a t e ly the y ar e not .

,

c r itical c o mbining fa ct f abl e and f ancy in a cco unt s of d r ago n s

, , , ,

gorgo n s a nd o the r im agin ar y m o n st e r s As a wh ol e P liny s

, .
,
’

influe n ce w a s d e t rim e nt al to z o ol ogical p r ogr e ss and he lp e d ,

in a ugu ra te a n e ra o f s upe rs titi o n .

C l a udi u s Gal e n ( 1 3 0 2 00 A D ) w a s a Gre e k physici a n wh o

—

p r a ctic e d f o r a tim e in R o m e He w as th e g re a t e st a n a to mist o f

.

a ntiquity a nd hi s w r iting s re m ain e d t h e b e s t o n t h e s ubj e ct unt i l

,

th e S ixt e e nth c e ntu ry Th e se w o rk s we r e t h e re sult s o f hi s o wn

.

c are fu l studi e s a nd disse cti o n s o f the high e r a nim als and hi s ,

de scripti o ns w e re r e m a rk ably cl e ar and fo rce ful .

3 . M I DDLE AGE S

T he M iddl e Age s bl ank so f ar as z ool o gical p r ogr e ss is

are a ,

co n ce r n e d Sup e r s titi o n w a s r a mp a nt a nd th e b e li e f in v ari o u s

.
,

fabl e d a nim als w a s p re val e nt All z o o l ogica l qu e stio n s we re .

re fe rr e d t o th e a nci e nt a uth o r iti e s a nd o r igin a l inv e stig a ti o n w a s ,

a t a st a nd s till In one cont rove r sy a se ri e s o f p ap e r s w a s pub

. f
FI G Li nna us Si x t y , —1
. 1 51 . at 1 70 7 7 78 .
( Fro m Lo cy )
.
2 70 AN INT R ODU C T ION TO ZOOLOGY

with whi ch h e w a s f a mili ar t o ge th e r with a syste m of cla ssi fi ca

,

tio n Thus th e w ay wa s cl e a re d f o r th e g re a te s t worke r in this

.

fi e ld C a r l Li nna u s ( 1 70 7 1 77 8 Fi g
,
— wh o a tt e mpt e d t o
,
.

de s c rib e a ll th e e xi s ting sp e ci e s of a nim a l s a nd pl a nts He .

succ e e de d in listing 4 3 7 8 in the te nth e di tio n o f hi s gr e a t e s t wo r k ,

S ys te ma N a tu r a H i s g r e a t influ e nce a nd th e wide re co gniti on

.
,

which w a s a ccorde d his wo r k m a de th e sys te m a tic s ide o f z oo l ogy

,

the m o st a ctiv e fi e ld o f inv e s tig a ti o n f or a l o ng tim e a ft e r hi s

de a th T he a i m o f th e sys te m a tic z oolo gis t h a s b e e n to d e scrib e
.

a ll th e sp e ci e s of a nim a l s a nd t o a rra ng e th e m a cc or ding to a

,

n a tu ra l syst e m i e a sys te m th a t will Show the i r t ru e r e l a tion

,
. .

shi ps to one a n o the r .

b C ompa r ati ve Ana tomy

.

An a to my up to the sixte e nth c e ntu ry consiste d in de sc riptions

o f t he s t r uctu re o f s ingl e a nim a l s T h e p o ints of re se mbl a nce of
.

diff e re nt a nim a l s fin ally le d z oo l ogis ts to co mp are th e a na tomy

o f o ne with a n o th e r T h e F re nch s ci e nti s t C uvi e r ( 1 7 6 9 1 83 2
.
—
,

Fi g I
. m a y h o w e v e r b e c o n sid e r e d th e fo und e r o f thi s b r a nch
, ,

o f z oo l o gy s inc e h e e xt e nd e d hi s s tu di e s o v e r th e e ntir e a nim a l

,
.

kingd o m a nd a dd e d a g re a t ma ss o f p e r son a l ob se r va ti ons to th e

,
‘

m a ny de scr ipti o n s publi she d by hi s p re d e c e ssor s B e side s a . .

numb e r o f t r e a tis e s o n co mp a ra tiv e a n a to my h e wro te a b oo k o n ,

th e fo ssil r e m a ins o f a nim a ls w hi ch fo und e d th e sci e nc e o f v e r t e

b ra te p al e o nto l o gy .

Am o ng C uvi e r s m o r e n o t e d s ucc e ss or s w e re th e Englishm e n

’
,

R ich ar d Owe n ( 1 8 0 4 1 8 9 2 ) a nd Th o m a s H H uxl e y ( 1 8 2 5 1 8 9 5)

— —
.

a nd th e Am e r i ca n E D C o p e ( 1 8 4 0
,
. . To R ich a r d Owe n
.

w e o w e t h e int r o ducti o n o f th e id e a s o f a n al o gy a nd h o m ol ogy .

Hi s w o r k o n th e c o mp ara tiv e a n a t o my o f v e r t e b ra t e s h as b e e n o f
gre a t se r vice e v e r Sinc e i ts publi ca tio n Altho ugh H uxle y .

m a de m any inv e s tiga ti o ns a nd publishe d a numb e r o f p ap e rs on

th e c o mp ara tiv e a n a to my o f a nim a l s h e i s b e s t kn o wn b e c a u se
,

o f h i s influ e nc e in p o pul a r izing z o o l o gy C o pe pl a ye d a n imp or

.
C uvi e r , 6 —1 8
1 7 9 3 2 .
1 53 . Willia m Ha r ve y, 1 57
-
8 1 667 .
( From Locy .
)
 HI ST ORI C AL ZOOLOGY 271

ta nt r Ole in th e a dva nce m e nt o f hi s cho s e n fi e ld o f wor k . He no t

o nly studi e d living f o r m s but w as o ne o f th e g r e a t e st
,
cont rib u

to rs to th e scie nce of p ale o ntol ogy .

c . Hi stology

B ich a t — f und hist l gy H w k

( 77 1 1 1 80 1 ) was th e
o e r o f o o i s o r .

w a s o f a hi gh or de r r a nking with th a t o f th e physi o l o gist J

, .

M ull e r and the e mb ryol ogis ts Va n B ae r and B alfo ur He .

di r e cte d the a tte ntion of biol o gists to th e study o“f tiss ue s a nd , ,

tho ugh h e did no t u se a mic ros cop e and f ail e d to m ak e o u t th e

c e llul ar str uctur e s hi s inv e stig a tions le d to th e mo de r n sci e nce
,

o f hist o l o gy .

T he int r oductio n o f th e mic r o scope a dd e d ne w imp e tus t o th e

scie nce of th e a n a to my o f ti ssue s and r e sulte d in th e anno unce
,

m e nt of th e c e ll the ory by Schw a nn in 1 83 9 a s de sc rib e d in ,

C h a pte r III (pp 3 5 . M ax Schultz e ( 1 8 2 5 by i de nti

f yi ng v e g e t a bl e p r o topl a s m with a nim al s a r c o d e r e fo r m e d th e ,

ide as re gardi ng c e lls R udolph Vi r cho w ( 1 8 2 1 1 903 ) t oo k up

.
—

th e study o f di se a se d ti ss u e s a nd h e lp e d e st ablish th e ce ll the o ry

'

a s it is und e r st oo d t o— d ay .

d E mbr yology
.

T he i e of e mb ry ol o gy d a t e s fro m the time of Willi am Harv e y

r s

( 1 57 8 1 66 7 Fi g I 5 3 ) a nd M a r ce ll o M a lpighi ( 1 6 2 8
—
,
. T he
fo r me r publishe d in 1 6 5 1 a w or k d e sc r iptive o f th e e mb ryol ogy of
t h e chi ck a nd o f se v e ra l m a mm a l s
, M alpighi w a s fur the r r e
.

m o ve d fro m the infl ue nc e o f th e a nci e nt s tha n Harv e y a nd hi s ,

w or k s on e mb ryo l o gy a re o f g re at e r v alue Hi s co nt ributi o ns .

include a de t ail e d de scriptio n o f the de ve l opm e nt of th e chick ,

illust r a t e d by e xc e ll e nt d r a wings .

P re vi o us t o th e tim e o f F ri e d r ich K Wo l ff e mb r yol o gi s t s b e

.
,

li e v e d in wh a t i s kn o wn as th e p r e fo r m a ti o n the ory Acco r ding .

t o thi s th e o r y th e e mb r yo e xi s t s in mini a tu re within th e e gg ,

a nd du ring d e v e l o pm e nt simply unf o ld s a nd e xp a nd s Fr o m .

272 AN INT R ODU CTION TO Z OOLOGY

thi s th e or y gr e w th e ide a of e nca se m e nt i e th e p re form e d e m

,
. .

b r yo mu s t co nt a in a se co nd sm a ll e r mini a tu re a nd thi s a n o th e r ,

ad i n fi n i tu m ,
ju s t a s a se rIe s o f b o x e s m a
y b e m a d e t o fit wit hi n
o ne a n o th e r . Wol ff did m ore than a ny o th e r e mb ryol o gist t o
o v e r thr o w th e p r e f o r m a tio n th e o r y a nd t o int r o duc e in i t s st e a d

t h e id e a o f e pig e n e s i s B y e pige n e si s Wo l ff m e a nt th e gra du a l

.

fo r m a ti on o f o rga n s fro m an unor g a niz e d e gg Th e e st abli sh .

~

m e nt o f thi s the o ry cha ng e d a ll sub se qu e nt wo r k in e mb ryol ogy .

Ka r l E v on B ae r ( 1 7 9 2 1 8 7 6 Fi g 1 54) w as th e g r e a t e s t Of a ll
.
—
,
.

e mb r y o l ogi s ts Hi s m os t imp o r t a nt s e r vic e s w e r e ( 1 ) th e e xc e l

‘

l e nce of hi s wor k whi ch r a i se d th e s t a nd ar d o f e mb ryol ogical

,

inv e s tig a tio n s ( 2) th e e s tablishm e nt o f th e ge r m l aye r the ory

,
-
,

a nd ( 3 ) th e b r o a d e ni ng o f t h e fi e ld o f e mb r y o l o gy by m a ki ng it
“
c o mp a r a tiv e He i s now kn o wn a s th e fa the r of m o de rn e m
.

b r yol ogy .
”
.
— —
F ra n ci s M B a lfo u r ( 1 8 5 1 1 88 2) in 1 880 1 88 1 pub
li sh e d h i s C ompa r a ti ve E m bryology T hi s w or k i s a summ ary o f
.

t h e e n o r m o u s lit e r a tu r e o n th e s ubj e ct th a t h a d a ccumul a t e d

during th e p e ri o d in a ugu ra t e d by v o n B ae r It al so cont ain s th e .

r e sul t o f much o r igin a l r e se a r ch a nd m a ny b r oa d g e n e r aliz a ti on s .

e . P hysi ology

T he m e di c al m e n o f a n cie nt tim e s d e p e nde d l arge ly up o n the

inve s tiga tio n s o f Gal e n Di se a se s w e re s upp o se d t o be d ue to
.

t h e p r e se n ce o f spi r it s a nd hum o r s in th e b o dy Thi s ide a ca ll e d.

,

th e pn e um a th e o r y w a s no t o v e r th r o wn until th e R e viv a l of
-
,

Le ar ning . T h e n a m e s o f Willi a m H ar v e y ( 1 57 8 Alb r e cht

Hall e r ( 1 7 0 8 a nd J o h a nn e s M illle r ( 1 8 0 1 1 8 5 8 ) a r e a m o ng
—

th e m o st fa m o us in th e fi e ld o f phy si o l ogy Harv e y d e m on .

s t ra t e d th e co nn e cti o n s b e tw e e n a r t e r i e s a nd v e in s a nd di s cov ,

e r e d th e ci r cul a ti o n o f th e bl oo d Alth o ugh th e s e co ntr ibuti o n s

.

t o kn owl e dg e h a v e m a d e hi s n am e f a m o u s th e int r o ducti o n by ,

him o f e xp e r im e nt a l m e th od s in physi ol ogic a l inv e s tig a tions h ave

h a d a m o r e p r o fo und influ e nc e up o n th e p rog r e ss o f phy si o l ogy .

Hall e r se p ar a t e d th e s tudy o f phy si o l o gy fro m m e di cin e a nd

 Jo ha nne s M ull e r —
FI G . 1 55 .
,
1 80 1 1 8 5 8 .
( Fro m Locy .
)
 AN INTRODU CTION TO ZOOLOGY

the i r o rigi n a l form though the y co ntinu e t o e xe r t an imp or tant

,

influe nce up on m a ny lin e s o f re se ar ch .

So m e of th e m or e r e ce nt w or k e r s in thi s fi e ld a r e M e nde l ,

We i sm ann and de Vr i e s Gre gor M e nde l ( 1 8 2 2 1 884) w a s a n

,
.
—

Au st ri a n m o nk wh o s e r e s ult s f ro m a study o f th e c ro s sing of

diff e re nt ki nd s o f p e a s and o the r pl a nt s h ave give n us one o f the
f e w l a w s o f h e r e dity An a cco unt o f hi s w o r k will b e fo und in
.

C h apte r X I V ( p . Augu s t We ism a nn (b o r n 1 8 3 4) i s a n

a r d e nt s upp or t e r o f Da rwini s m He is th e f o r e mo s t living e vo
.

lu ti onary z ool ogi s t .Hug o d e Vr i e s h a s r e c e ntly b ro ught fo r th

a w o r k e ntitl e d Di e M u ta ti ons the or i e publi she d in 1 90 1 which
, ,

co mb a ts Darwin s the orie s o f th e o rigin o f sp e ci e s and off e rs

’

“ ”
in its st e a d th e mut atio n th e or y a discussio n o f which is
,
,

r e s e rv e d f or th e n e xt ch a pt e r .

g
. Z oology f
o To-day

As th e fa cts o f zool o gical s cie nce s h ave incr e ase d in numb e r ,

th e fi e ld s o f w o r k h a v e b e c o m e m o re num e ro u s a nd n a rrow e r a nd ,

inve stig a tio n s ar e no w c a rr i e d on by m ore imp r o ve d m e tho d s a nd

in gr e a t e r de tail th a n e ve r b e for e M o rphol o gi cal s tudie s a r e
.

b e ing suppl e me nt e d by e xp e r im e nt al inve s tig a tio n s in e mb ryol

o gy r e g e n e r a ti o n h e r e dity e v o luti o n a nd o th e r s ci e nc e s
, , , ,
Ani.

m al b e h a vi or i s o ne o f th e m os t f a v o r e d s ubj e ct s o f r e se a rch a nd ,

i s r apidly l e a di ng p sych o l ogists t o a b e tt e r und e r s t andi ng of th e

a nim a l m ind .

Z ool o gy a t th e p r e se nt tim e p r e s e nts a r i ch fi e ld f o r o rigin al

re s e a rch. M a ny o f th e a pp are ntly sim pl e “ l aws h ave b e e n
found o n cl o se e x a min a ti on to b e re ally ve ry c o mpl e x All lin e s .

o f e xp e r im e nt a l w o r k o ff e r l a r g e r e w a r d s f o r t h e s tud e nt a nd o p e n

up f o r him a co untl e ss numb e r of f ascin a ting p robl e m s .

FI G 1 5 6. C har l e s Da r wi n, —1 8 8 2
. 1 8 09 .
( Fr om Lo cy )
.
2 76 AN INT RODU C TION TO Z OOLOGY

c ontin e nt o f N o r th Am e r i ca h a s b e e n divide d by s ci e ntis t s into

de finite r e gi o n s a cco r ding to th e s u m to t a l o f th e t e mp e ra tu re
,

du ring th e se a so n o f g rowth ; a nd r e gio ns o f a ce r t a in t e mp e r a tu r e ,

tho ugh wide ly se p ara t e d a r e liabl e to s upp o r t simil ar f a un as

,

Wint e r is m e t by n o r the r n a ni m al s in o ne of fo ur w ays

( I ) by dying e g a dult butt e rfli e s ( 2 ) mig r a ting e g bi r d s
,
. .
, ,
. .
,

( 3 ) hi b e r na ting e g b e a r s (4 ) r e m aining a ctiv e e g rabbit s

,
. .
, ,
. . .

Anim a l s living in t r o pi ca l r e gi o n s p ass th e s umm e r in m a ny c a s e s

i n a t o rpid co nditi o n a nd a r e s a id t o b e a sti v a ti ng
,
.

A ce r t a in a m o unt o f w a t e r i s n e ce s sa r y f o r life a s th e b odi e s ,

of an i m al s a r e m a d e up o f f ro m 55 to 9 5 pe r ce nt w a t e r In d ry .

clim a te s a nim al s h a ve thi ck Ski n s a nd thus e va p or a ti o n i s pr e ,

ve nt e d Aqu a ti c a nim al s obt a in ai r fro m w a t e r as d o a l so

.
,

s o m e t e rr e s t r i a l sp e ci e s e g th e e a r thw o r m
,
. T h e d e n s ity o f
. .

t h e w a t e r a nd i t s s a linity d e t e r min e th e di s t r ibuti on o f m a ny

a qu a ti c o r g a ni s m s .

Light a s w e h a v e se e n pl a y s a l e a ding r Ole in t h e liv e s of a ni

, ,

m al s ; m a ny sp e ci e s r e qui re it ( Eu gle na ) but o the r s shun it a s ,

much a s p ossibl e ( cr a yfi sh) p r in cip a lly in o r de r t o e scap e the ir

,

e n e mi e s .M ol a r a g e nt s s u ch a s curr e nt s l i mit th e numb e r o f

, ,

sp e ci e s a nd individu a l s e sp e ci a lly wh e r e th e y a ct with mu ch

,

fo r c e And fin a lly foo d co nditio n s a r e m o s t e ff e ctive Sin ce

.
, ,

ca r niv o r o u s a ni m a l s e g li on s mu s t liv e wh e r e th e y m ay obt a in

,
. .
,

fl e sh h e rbivor o us anim a l s e g d e e r mu s t liv e wh e r e s uit abl e

, ,
. .
,

v e ge ta ti o n ab o und s a nd o mnivo ro us a nim a l s e g cr ayfi sh e s

, ,
. .
,

Wh e r e b o th fl e sh a nd v e g e t a ti o n o f c e r t a in so r t s e xis t Tabl e .

X I V p re se nt s ro ugh ly th e f o u r p r in cip a l kind s o f f a un a a nd th e i r ,

m o de s o f e xi st e nc e .

Th e g e n e r a l s t a t e m e nt m a y b e m a d e th a t t h e m a j o r h abit a t s
a re b r o k e n up int o min or h a bit a t s by v a r i a ti o n s in t h e co n di ti o n s ,

a nd th e c o n s tituti o n o f th e o r g a ni s m Th e r e a r e g re a t diff e r .

e nce s in th e e x actn e ss with whi ch t h e di ff e r e nt f o r m s a r e c o nfin e d

‘

to di ff e r e nt se t s o f e nvi r o nm e nt a l co nd iti o n s Thi s will b e .

b e tt e r u nd e rs too d a fte r th e o the r l a w s o f di s t r ibution h av e b e e n

di s cusse d .
 G EN E R AL ZOOLOGI C AL FAC T S AN D T HEOR ES I 277

T AB LE X I V

THE F OU R KI N DS OF EAUN A AND THE I R M ODE S OF E X I S TE N C E

F AU N A H A B I T AT F AC T OR S OF H AB I TAT EX AM P LE S

Ha l ob i o s M a r i ne

Li m no b i o s F re sh wa te r De ns it y
Wa t e r Da rk ne ss
M o l a r a ge nt s

'

Ge o b 103
D ry , l i ght ,
lo w s pe C I fi c gra vi ty

Sub t e rre s t r i a l T hi gm o t a xi s , d a r k ne ss

B i o bio s P a ra siti c F l u i d f o o d , da r k ne ss

T HE LA W OP D I SP E R S I ON . Anim a l s
t e nd t o mig ra t e f ro m th e
r e gi o n o f th e i r o r igin T h e r e a so n f o r thi s i s in di sput e a t th e
.

p re se nt ti me It h as b e e n h e ld f o r m a ny y e a rs th a t e v e r y a nima l
.

p roduce s a g re a te r numb e r o f o ff sp r ing th a n ca n b e supp or te d

in i ts p ar ticula r ha bit a t a nd Sin ce p a re nt s a nd o ffsp ring c anno t
, ,

o ccupy t h e s a m e a r e a s o m e individu a l s mu s t e ith e r mig r a t e o r

,

d ie . The re a r e m a ny sci e nti s t s o n th e o the r h a nd wh o b e lie ve

, ,

th a t no o ve r c r o wding t a k e s pl ac e but tha t di sp e r sio n is th e ,

r e s ult o f a s e a r ch f or f oo d .

E a ch Sp e ci e s o f a nim a l i s s upp o s e d t o h a v e o r igin a t e d in a d e h

nite r e gio n o f th e e ar th s s u r face a nd t o h av e mig ra t e d in v ari ou s
’

di re ctio n s e nlarging its h abit a t y e a r by ye a r until a n appro xi

,
2 78 AN INTR ODU CTION TO ZOOLOGY

m ate ly p e rm an e nt ar e a is o ccupi e d T h e r e gi o n fro m which .

mi gra tio n took pl a ce i s te rm e d th e ce nte r of di sp e r sal T h e .

fo ll ow ing c rite ri a have b e e n giv e n to de te r min e the c e nte r of d i s

p e rsal o f a sp e cie s : th e abund anc e a nd siz e o f individu al s the ,

l o ca tio n of cl ose ly re l ate d forms a nd the mig ra ti o n ro ute s now

,

se l e cte d by th e sp e cie s
T H E LAW OF B ARR I E R S AN D HI GH WAY S — Anima ls a re c o nfin e d .

t o c e r t a in h abit a ts by b ar r i e r s The y a r e p r e ve nt e d fro m ga ini ng

.

a cc e ss t o a ne w r e gi o n by th e ch a ng e in t h e m e di a by d e a r th o f
,

foo d a nd th e inte r fe re nc e o f o th e r anim al s C omm on b a rr i e rs

, .

a r e m o unt a in s b o di e s o f w a t e r o p e n c o unt r y f o r f ore s t a nim a l s

, , ,

a nd f or e s ts f o r p r a i ri e inh a biting sp e ci e s
-
F or e x a mpl e th e c r a y .
,

fi sh C a mbar u s mig r a t e s up a nd d o wn st r e a m s but c a nn o t t r a ve l

, , ,

o v e r l a nd t o n e ighb o r ing s t r e a m s t h e h o n e yb e e c a nn o t fly a cr oss

,

t he o ce a n nor ca n Hydr a e nt e r th e s e a
, .

T h e r e ve rse o f a b a rri e r is a highw a y App a r e ntly th e r e a r e .

r o ut e s o f m ig ra ti o n whi ch a r e e sp e ci a lly f a v ore d Thi s m ay be .

illust rat e d by th e fli ght o f bi r d s so uthwar d in th e a utumn a nd ,

n orthward ag ain th e fo llo wing sp ring M a ny bi r d s mig r a t e .

up and do wn the M issi ssippi Vall e y a nd -al o ng the Atl a ntic ,

co as ta l pl ain .

C O SMOP OLI TAN GR OU P S OF AN I MAL S So me sp e ci e s o f a nim a l s

.

have wid e ra nge s e g so me are fo und inhabiti ng p ra ctic ally

,
. .

e ve ry l a r g e l a nd a r e a o n th e e a r th s su r fa c e Sixt e e n f a mili e s
’
.

o f bi r d s including dov e s o wl s a nd ducks a nd o ne f a mily o f

, , , ,

m amm al s the b a ts a r e co s mop o lit a n g ro up s Do ubtl e s s th e

, ,
.

wings o f the bi r ds have facilita t e d the i r di sp e rsal since the y g ive ,

the m r e m a rk abl e p o we rs o f l o co m otio n .

R E STR I C TE D GR OU P S OP AN I M AL S In a numb e r o f c ase s ce r ta in

.

sp e cie s a r e r e s tr ict e d to v e ry limit e d a r e a s T h e m o unt ai n goa t .

i s fo u nd only in t he high e r R o cky a nd C a s c a d e m o unt ain s to

Al a sk a . I sl a nds a re fa mo us f o r th e p re se n ce o f r e st r ict e d sp e cie s .

D a r win s d e sc riptio n s o f th e a nim al s h e fo und in th e Ga l ap a go s

’

Isl ands re a d like fairy ta l e s

DI SC ON TI N U OU S DI STRI B U TI ON —
Wh e n e ve r a sp e cie s o ccur s in
.
 280 AN IN T R ODU C TI ON T O Z OOLOG Y

orde r o f th e i r succe ssio n a nd th e tim e ,

of or igi n a nd re a l tive
numb e r o f th e diff e r e nt g r oup s o f a ni m al s
T AB LE X V

T HE G E OL OG I C AL P E RI ODS AN D TH E ORI G I N AN D RE L AT I VE NUM B ER OF

TH E DIF F E R E N T G R O U P S OF A N I M A LS

PE R IOD

CO
0) on
5
iii E S: 2
<
2.
2 m 1
In m S 2

R e ce nt
P l e i s t o ce ne J

C e no z o i c P li o ce ne l
|‘
M i o ce ne r Io

E o ce ne l J

IO 1

Me s o z o i c 5
I
5 J

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S il u r i a n 20

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A r ch a an Lau r e nti a n

(3 ) Var i a ti ons in Ani m als

We h ave al r e a dy n o te d (p 7 9) th a t no tw o o r g a ni sm s ar e e ve r
.

e x actly a lik e I n o th e r w o r d s individu a l s Of diff e r e nt sp e ci e s

.
,

a r e s o dis si mi l a r a s t o b e in m o s t ca se s e a sily r e cogni z e d Indi

‘

.
 GEN E R AL Z OOLOGI C AL FAC T S AN D TH EOR ES I 28 1

vi d u a ls o f s a m e sp e cie s if e x a mi n e d cl ose ly will a l so b e f o und

th e , ,

to v a r y in ce r t a in r e sp e ct s ; a nd e v e n t h e o f fsp r ing o f th e s a m e
p a r e nt s di ff e r fr o m o ne a n o th e r t o a g re a te r o r l e ss de gre e .

T h e di ffe r e n ce b e tw e e n chil d a nd p a re nt a nd b e tw e e n child r e n is

ca ll e d v a r i a ti o n Va r i a ti o n s a r e imp o r t a nt in a ny di scu ssio n Of
.

e voluti o n a nd h e r e dity sin ce th e i r or igin a nd influ e n ce up o n th e

,

t r a n smut a ti o n o f sp e ci e s co n s titute th e fund a m e nt a l p robl e m s

th a t a r e o ccupying th e a tte ntio n o f phi l osop hi ca l z o ol ogi st s a t
t h e p r e s e nt tim e .

C ON TI N U OU S OR F L U C T U ATI N G VAR I A TI ON S If a hundr e d m e n

.

a r e g a th e r e d a t ra nd o m a nd a r ra ng e d in a r o w a cc o r ding t o th e i r
,

he ights it will b e fo und th a t th e y d e c re a se g ra dua lly f ro m th e

,

t all e s t a t o ne e nd t o th e sho r t e s t a t th e o th e r This i s a n i llu S .

a

t r a t i o n o f co ntinu o u s v a r i a ti o n If no w w e e x amin e this lin e

.

m o r e cl o se ly w e will find th a t th e g re a t e s t numb e r m e a s u r e a b o ut

,

s ixty fi v e in ch e s in h e i ght
-
a nd th a t t h e numb e r o f a ny h e ight
,

d e cre a se s a s th e e nds o f th e lin e a re app ro a ch e d T h e m e as ure .

~

m e nt s d e cr e a se o r i ncre a se g r a du a lly fr o m th e ce nt r a l m e a n to
o ne e nd o r th e o th e r a nd a s m a ny v a r i a ti o n s will b e fo und a b o v e
,

th e m e a n a s b e l o w T h e Off sp r ing o f a ny o f th e in d ividu al s

.

s e l e ct e d will Sh o w a v a r i a bil ity th a t i s Simil a r ly flu c tu a ti ng a nd ,

foll o w s th e law o f p r ob a biliti e s This la w h old s th a t Slight .

m o di fica tio ns fro m th e m e a n a r e m os t abund a nt a nd th a t th e ,

gr e a t e r th e v a ri a tio n a nd th e n e a re r th e e xt re m e s a re a pp roach e d
, ,

t h e l e ss num e ro u s th e m o difi ca ti o n s b e co m e .

D I S C ON TI N U OU S VAR I A TI ON S Anim al s kn o wn a s
. sp or ts ,
”
s a lt a ti o n s or ,
mut a tio n s a re e x a mpl e s o f di sco ntinu o u s
v a ri ability The se di ff e r fr o m co ntinuo u s v a ri a tio ns in th a t th e y
.

are abl e t o t r a n s m it th e i r diff e r e n ce s fro m th e m e a n to the i r d e

s ce nd a nt s .So m e o f th e m o s t n o t e d s udd e n v a ri a tio n s a r e th e
ho rnl e ss c a ttl e o f P a r a gu a y th e ni a t a b r e e d o f o x e n o f So uth
,

Am e ri ca a nd t h e m a u ch a m p a nd a n co n Sh e e p
,
T h e l as t n am e d
.
-

o r igin a t e d f ro m a r a m with cr o o k e d l e g s a nd a l o ng b a ck th a t w a s ,

b o r n in M a ssa chu se tt s in 1 7 9 1 Wh e n cr o sse d with comm o n .

Sh e e p ,
hi s p e culi ar ch a ra ct e r i s ti cs r e app e a r e d in th e o ff sp r i ng ,
28 2 AN IN T R ODU C TION TO ZOOLOGY

a nd by inte rb r e e ding th e indi vidu al s th a t sho we d the se t r aits the

a nc o n ra ce o f sh e e p w a s e s t a bli s h e d T h e mo s t fa mo u s
di sco ntinu o us v a r i a tio n s kn o wn in th e pl a nt ki ngdo m a r e th e
di s tinct sp e ci e s o f e v e ning p rim rose s whi ch h a v e o r igin a t e d in
,

r e ce nt y e a r s f ro m th e co mm o n p r im ro se a nd h a v e b e e n so w e ll
,

de scrib e d by th e Dutch n a tu r ali s t d e Vrie s

(4) Ad apta ti o n s of Anim al s

The d apta ti o n s o f a nim a l s to the i r e nvi ro nme nt i s so u sua l a s

a

to p a s s unn o tice d E a ch o f th e typ e s w e h av e s tu di e d i s fitt e d

.

f o r life in i t s p a r ti cul a r h a bit a t a nd in m os t c a s e s in no o th e r a

, ,

fa ct whi ch a s w e h a v e s e e n a cco unts l a rge ly f or th e ge ogr a phi cal

, ,

di stributio n o f o rg a ni sm s Anim a l s no t o nly a re a dju ste d to the i r

.

s u r r o unding s but p oss e ss s t r uctu r e s which s e r v e th e m in obt ain

,

ing foo d in de fe ndi ng the m se lv e s a nd th e i r yo ung a nd in m ain

, ,

t a ining s up re m a cy a mo ng i ndividu a l s o f the ir ki nd T h e cr a y .

fi sh i s a d apt e d to i ts e nvi r o nm e nt by th e p osse ss io n o f g i ll s f o r

br e a thing e y e s f or d e t e cting m o vi ng o bj e ct s i n th e w a t e r a h a rd
, ,

e x os k e l e t o n t o p r o t e ct it f ro m v a ri o u s e n e mi e s pin ch e rs f o r o ff e n s e
,

a nd d e f e n se a nd a p ositiv e
,
thigm o t r opic r e a cti on which c a u se s
it to s e e k a d ar k sp o t ou t of ra ng e o f m a r a ude rs .

Inse cts a r e e sp e ci ally favo r abl e f or i llu st ra ting a dapt a tion s .

Th e foll o wing s t r uctu re s o f th e h o n e yb e e a re e xc e ll e nt e x a mpl e s

o f a d a pt a ti o ns : t h e h a i r s a nd l e g s f or g a th e r ing p o ll e n th e a li ,

m e nt ary c an al f o r o bt aining a nd s to r ing n e ct ar th e m a ndi bl e s ,

f o r m o l di ng w a x th e s ting f or se lf d e fe n se f o r th e d e fe n se o f the
,
-
,

hiv e a nd in th e case o f th e qu e e n f or u se in d e te r mi ning h e r

, , ,

s up r e m a cy T h e o rigin o f a d apt a ti o ns i s a m o o t qu e s ti o n a m ong

.

biol ogis ts Th e y a r e v a r i a tio n s which h av e b e e n inh e rit e d a nd

.
,

hav e b e co m e m o r e p r o n o unce d in th e co u r se o f co untl e ss ge n e ra

ti ons .
2 84 .
AN INT R ODU CTION TO Z OOLOGY

influ e nce th e i r life a ctiviti e s SO th a t w e a r e abl e to se e v e r y littl e

,

if a ny incr e ase but e v e n a p o s sibl e de cre a se in t he numb e r o f

, ,

bir ds tha t g r e e t u s with th e co mi ng o f e a ch sp ri ng .

Th e s e ani m al s a r e no t p e culi ar in a ny w ay sin ce any sp e cie s ,

tha t might b e se l e ct e d wo uld s oo n co v e r th e e a r th with i ts de

sce nd a nt s if a ll o w e d t o r e p r o duc e unch e ck e d In m os t ca se s th e .

numb e r o f individu a l s o f a sp e ci e s i s p r a ctic ally co nsta nt fr om

y e ar t o ye ar Th s i s the r e s ult o f th e st r uggl e f o r e xi st e nce
i
. .

An ani m al mu s t c o nt e nd with i ts s u rr o undi ng s ; E u gle na mu st

h av e light t o car r y o n i ts life p roce sse s ; a nd th e h o n e yb e e m ust
sto r e up p ro vi si o n s and m ake i t s hive se cu r e in or d e r to su rvive
t h e wint e r .

T h e h abit a t o f a sp e ci e s i s n e v e r l a r ge e n o ugh t o a cc o mm o d a t e
a ll th e individu al s o f a s p e ci e s a nd a n a nim a l i s f o r ce d t o b a ttl e
,

f o r life a g a in s t o th e rs o f i ts kind T h e s t r o ng e r o th e r thi ngs

.
,

b e ing e qu al alwa y s win s in th e s t r uggle in o th e r w o r ds th e fitte st

, , ,

s u r viv e s . F o r e x a mpl e in p oo r se a s o n s f o r g a th e r ing h o n e y

, ,

be e s f ro m o ne hi v e a tt a ck th o s e o f a n o th e r a nd in m a ny c a se s , , ,

ca r r y o ff th e p o ll e n a nd h o n e y l e a ving th e i r victim s t o s t a r v e
,
.

Oth e r b a ttl e s o ccu r betw e e n ind iv i du al s o f di ffe r e nt Sp e ci e s .

Wha t e ve r the ca use o f th e st r uggl e it i s alwa y s th e w e a ke s t tha t

,

i s v a nqui s he d T h e s t ro ng e s t individu a l s o f a Sp e ci e s liv e o n

.

a nd k e e p up th e vig o r o f th e r ace thi s b e ing o ne o f the a dvan

,

t age s o f th e e limin a tio n o f th e unfit .

( )
6 H e r e d y it
He re dity h a s b e e n de fin e d (p 7 9) a s th e r e se mbl an ce o f child
.

t o p a r e nt A ch a r a ct e r m a y b e sa id t o b e inh e rit e d whe n it

'

a lw a y s in o ne g e n e r a ti o n a ft e r a n o th e r i s o ne o f t h e ch aract e r s o f
, ,

t h e sp e ci e s , ( 2 69 p T h e ch a r a ct e r s o f a n o rg a ni sm
,
.

de p e nd o n th e n a tu re o f i t s s urr o undings e sp e ci a lly du ring d e ,

v e lopm e nt a nd o n t h e co n s tituti o n o f th e p a r t o f t h e p a r e nt f r om
,

which it g re w In M e t a zo o n s thi s m ay b e a b u d as in Hydr a

.
, ,

o r a n unfe r t i liz e d e gg ce ll a s in th e d r o n e b e e
,
but i s u su ally an ,

e gg fe r tiliz e d by a s p e r m .
 G EN ER AL ZOOLOGI C AL F AC T S AN D T H EOR E S I 28 5

We m ay study inhe r it a n ce by e x a mining th e g e r m c e lls by ,

co mpiling s t a tisti cs o r by th e e xp e r im e nt a l m e th o d o f cross

,

b re e ding a nd sub se qu e nt a n a ly si s Of th e ch a ra ct e rs o f th e off

spring I t i s no t p o ss ibl e in a ny c a se to p r e di ct wh a t cha r a ct e rs
.

will b e fo und in th e o ff sp ri ng o f kn own p ar e nts altho ugh in ,

ce r t a in in s t a n ce s to b e b r o ught ou t l a t e r thi s m a y be d o n e with

, ,

som e de g re e o f a ccur a cy

( 7) Th e Eff e ct s of Is o l a ti o n
M o st e v o luti o ni s t s b e li e ve th a t n e c e ssa ry co nditio n f o r the
a

form a tio n o f ne w spcci e s i s th e i sol a tio n se p a r a tio n or se g re, ,

g a t i o n o f c e r t a in individu a l s f r o m o th e r s o f th e i r kind T hi s m ay .

be d u e to e ith e r g e o g r a phi ca l o r phy si o l o gica l ca u se s .

GE OG R AP HI C AL I S OL AT I ON . In th e ca se o f g e og raphic Or top o
g raphic iso l a tio n th e i sol a t e d g ro up o r gro up s o f indivi du al s a re
actu a lly i n a n o th e r r e gi o n o r l o c a lity fr o m th e r e s t o f t h e sp e ci e s ,

this b e ing the r e s ult o f m ig r a tio n v o lunt a r y o r inv o lunt a ry

,

( 5
2 0,
p . W e h a v e S h o w n (p
'

2 8 4). th a t n o r m a lly th e r e is
a n o v e r p r o ducti o n o f a nim a l s a nd th a t di sp e rsa l i s n e ce ssa r y f o r

th e p re s e rv a ti o n Of lif e .
“ ,

In r e gi o n s b r o ke n by f e w b a rr ie r s ,

mig r a tio n a nd inte rb r e e ding b e ing a ll o w e d w e find wide ly ,

dist ribut e d sp e cie s h o m og e n e o u s i n th e i r ch a ra ct e r th e m e m

, ,

b e rs Sho wing individua l fluctu a ti o n a nd clim a ti c e ffe cts but ,

re m a ining unifo r m in m os t re g a r d s . In r e gi o ns b rok e n by

bar r ie r s whi ch i sol a t e g r o ups o f individu al s w e find a g r e a t
numb e r of r e l a t e d sp e ci e s th o ugh in m os t c a se s th e sa m e r e gi o n
,

conta in s a sm all e r numb e r of g e n e r a o r familie s .Giv e n

a ny sp e ci e s in a n r e gi o n t h e n e a r e s t r e l a t e d sp e ci e s i s no t lik e ly
y ,

t o be fo und in t h e s a m e r e gi o n no r in a r e m o t e r e gi o n but in a ,

n e ighb o ring dis t r i ct se p a r a t e d fro m th e fi rs t by a b a rr i e r o f s om e

so r t

T h e f a un a o f i sl a nd s sh o w in a s t r iking m a nn e r th e e f fe ct s o f
i sol a tio n Som e o f the se i sl a nd s if no t o f volc a nic o rigin a nd
.
,

th e i r a nim al s th e d e sce nd a nt s o f indivi d ual s th a t w e r e d r ive n

28 6 AN INT RODU CTION TO ZOOLOGY

th e re by winds or c arri e d o n flo a ting o bj e ct s h av e f or a v e ry ,

l o ng time b e e n i sol a te d a nd thi s i sol a ti o n h a s so cha ng e d the

,

cha ra ct e r o f the se ani m al s th a t the y a re now re cogni ze d as

di s tinct sp e ci e s The r e can be no do ubt tha t th e sp e cie s o rigi
.

na te d f ro m indi vidu a l s f r o m th e m a inl a nd a nd th a t th m os t ,

e

impor ta nt f a cto r in the i r t ra n smut a ti o n h a s b e e n g e ographi ca l

i sol a tio n
P H Y SI OLOGI C AL I S OL ATI ON — Inte r n al di ff e re n ce s m ay c a use
.

physiol ogica l i sol a tion a fa cto r in th e origin o f sp e ci e s a s imp o r

,

ta nt p rob ably a s g e ogr aphic al i sol a ti o n P hy si o l ogi ca l o r a s it i s

.
, ,

so m e tim e s c a lle d se xu a l i so l a ti o n i s ca u se d by
,
th e influ e n ce
o f s o m e v a r i a ti o n t e ndi ng to m a k e d ifli cul t o r imp ossibl e wh o lly

f re e a nd misce ll an e o us m a ting or b r e e ding insid e o f a sp e ci e s .

This v ari ati on m ay be o f pu r e ly physiol o gi cal ch ara cte r o r m ay

be a s t r uctu r al one : th a t i s th e hind r a nc e to m a ting m a y be o ne
,

o f ins tinctiv e f e e ling a r a ce f e e ling

,
de p e n di ng o n a n a ntip a thy
-

t o Od o ur t o a ge to app e a r a nc e e t c or m ay b e a sli ght m o difi ca

, , ,
.
,

ti o n of the copul a tory o r g an s m aking su ch m a ting difli cult o r ,

e v e n a m o difi ca ti o n o f th e e gg o r t h e sp e r m a t o z o id s m a ki ng f e r

tili z a ti on d ifli cult It i s a w e ll kn o wn fact th a t n u me r o us

.
-

v a ri e ti e s of do m e s tica t e d a nim al sp e i e s ra r e ly b r e e d tog e the r

c ,

a lth o ugh quit e a bl e t o a nd p r o vid e d wit h f ull o pp o r tun

,
i ty On .

th e o th e r h a nd a nim a l s o f diff e r e nt sp e ci e s w hi ch in N a tu r e
,

r are ly or n e v e r b r e e d t o g e th e r m a y if k e pt l ong in c onfin e m e nt

, ,

a s in z oo l o gic a l g a r d e n s m a t e a nd p ro duc e y o ung

,
In e a ch case .

th e re s e e m s to b e que stio n o f a r ace fe e ling in th e fi rst c ase -

a se xu a l a v e rsi o n k e e ping a p a r t individu a ls o f th e s a m e sp e ci e s ,

in th e s e cond th e b r e a ki ng do wn o f r a ce fe e ling th a t in N a tu re -

”
h a s s uff ic e d t o p r e ve nt hyb ridi zing ( 5
2 0 p In m a ny
,
.

c as e s if int e rb re e di ng doe s o ccur th e hyb rid o ff sp r ing ar e ste r il e

, , ,

e g th e m u l e p r o duc e d by a c r oss b e tw e e n a f e m a l e h or s e a nd a
. .

m al e a ss . B y physi ol ogic a l i so l a ti o n th e n g r o up s of i ndividu als

, ,

a r e f or m e d withi n th e sa m e re gi o n T h e r e s ult is in e v e ry way

.

similar t o se gre ga tio n by g e og r aphi c a l b arr ie r s .

2 88 AN INTRODU C TION TO Z OOLOGY

th e ge r m c e ll s a s w e ll a s the b ody c e ll s a r e a ffe ct e d by th e e xte r

nal s timuli a nd th e b o dy ce ll s h a v e no e ff e ct up o n th e g e r m c e ll s
,
.

Th e th e or y i s a l so a pp a r e ntly va lue l e ss i n e xpl a ining i nh e r it a nce

in th e P r o t o z oa (p .

T HE C ON TI N U I T Y OF TH E GE R M P L AS M T h e p r e s e nt wid e
.
-

sp re ad b e li e f in th e th e or y o f th e c o ntinui ty o f th e g e r m pl asm i s
l arge ly d ue to th e e ff or ts o f We i sm a nn We ha ve al r e a dy i llus
.

t r a t e d th e th e o r y in a g e n e r a l w a y by th e life hi s t o r y o f Volvox
,

p
( 9. 8,
F i g. a n d in a n o th e r pl a c e ( p 3 2 ) h a.v e p o int e d ou t
tha t th e chr o moso m e s a r e tho ught by m a ny to b e th e b e a re rs o f
he r e dit a ry cha r acte r s .

Acc o r di ng t o We i sm a nn th e r e a r e
,
tw o g re a t c a t e g o ri e s o f
living sub sta nce h e re d it a ry s ub s t ance o r idi opl asm a nd nu ,

t ri ti v e s ub s t a nc e or t r oph o pl as m ( 2 7 8 Vol I p
”
’
,
He .
,
.

r e co gniz e s th e ch r o m a tin a s t h e idi o pl a s m a nd c a ll s t h e idi o pl a s m

,

of th e g e r m c e ll s g e r m pl a s m
,
Thi s g e r m pl a sm is n e v e r
.

fo r m e d de novo but it g ro ws and inc r e a s e s ce a se l e ssly ; it i s h and e d

,

o n f ro m o ne ge ne r a ti o n to a no the r like a l o ng r oo t cre e ping

thr o ugh th e e ar th fr o m which a t r e gul ar d ist a nce s shoot s gro w u p
,

a nd b e co m e pl a nt s t h e individu al s o f t h e s u cc e s s iv e g e n e r a ti o n s
,

( 7
2 8,
V o l
. I p,
.

T he d e t a il s o f We i s m ann s th e o r y a nd th e ing e nio u s e xp l a n a

’
,

ti o n s o f s u ch phe n o m e n a a s c e llul a r diff e re nti a tio n a nd r e ge n e r a

ti on a re e xt re m e ly c o mplica t e d B ri e fly s t a t e d th e m e cha nis m
, .
,

i s co n ce iv e d by We i s m ann a s fo ll o w s : E a ch l a rg e ch r o m oso m e
o f a g e r m ce ll is co mp os e d o f ch r o m a tin g r a n ul e s c a ll e d
“ i ds
”
,
.

Each id i s e ith e r m al e o r f e m a l e a nd c o nt a ins all o f th e

,
co m

pl e xe s o f p rim a ry co n s titu e nt s n e ce ss ar y to the p ro ductio n o f a

”
co mpl e t e individu a l ( 2 78 p ,
. T h e p r o du cti o n o f a p a r
ti cu lar p ar t o f a n o r g a ni s m i s th e r Ole o f th e p r im a r y co n s titu e nt s
o f th e id a nd sinc e th e se d e t e r min e t h e n a tu r e o f t h e p a r t p r o

,
, ,

d u ce d th e y a r e n a m e d
“ d e t e r mi na nt s ”
F in ally t h e d e t e r m i
.

n ants a r e co mp ose d o f o ne o r m o re p a r ti cl e s ca ll e d bi opho rs ”

“ “
.
,

The se a re f a r b e l o w th e l imit s o f vi s ibility but a r e l a rg e r

”
,

than any che mica l m ol e cul e s b e c a use the y th e m se lve s co n si s t of

 G E N E R AL Z OOLOGI C AL F AC T S AN D T HEOR ES I 2 89

a g ro up o f mol e cul e s a m o ng which ar e some o f c o mpl e x c omp osi

,

ti on a nd th e re f o r e o f r e l a tiv e ly c o n s id e r abl e s ize

, ( 78 p
2 ,
.

—
GALTON S LAW OF AN C E S TR AL I N HE R I TAN C E F ra nci s Ga lt o n
’
,

f rom a study Of p e digr e e d d og s a nd th e ge n e al ogic a l re c or ds of the

B r iti sh P e e r a ge c a m e to th e c o nclu sio n th a t in a n o rg ani sm o f
,

bise xu al p ar e nta ge T h e tw o p ar e nts b e tw e e n the m cont ribut e

,

o n th e a v e r a g e o ne h a lf o f e a ch inh e r it e d f aculty e a ch o f th e m ,

cont ributing one qu ar te r o f it T h e fo u r g r a ndp a r e nt s co nt ribut e

.

b e twe e n the m o ne qu ar te r o r e a ch o f the m o ne s ixte e nth ; a nd so

,

—
o n th e sum of t h e s e r i e s é f i f 3 + T g +
- -
b e ing e qu al t o
l
, 5
”
1 ,
a s it s h o uld b e It h as b e e n d e m on s t r a te d tha t thi s la w i s
.

onl y a n a pp r o xi m a ti o n o f th e r e a l re sul t

M E N DE L S LAW OF I N HE RI TAN C E ( 2 2 7) T h e re sul t s o f M e nde l s

’ — ’

e xp e rim e nts in hyb r i di zing p e a s a nd o th e r pl a nt s w e r e publi sh e d

by him in 1 8 66 but di d no t re c e iv e a ny a tt e ntio n f ro m biol ogi s ts

,

until 1 900 wh e n the y w e r e dis co v e r e d in a n ob scu re p e rio dical

,

by se v e ral b o t a ni sts Since the latt e r d a t e M e nde l s law h as

.
,
’

b e co m e a favor it e obj e ct o f inve s tig a tio n with b o th b o tanists and

z ool ogis t s until now i ts c o mpl e xitie s a r e so g re a t as to m ak e th e
,

d e t a il s o f th e subj e ct unint e lligibl e e xc e pt to p e r so n s w o r king in

this p ar ticul ar fi e ld .

An o utlin e o f o ne o f M e nd e l s e xp e r im e nts will se rv e t o illu s

’

t ra t e hi s la w T h e v ar i e ti e s o f t h e e dibl e pe a w e r e se
le cte d by hi m f or hyb r idiz a tio n Wh e n a t all pe a w a s c r o sse d
.

with a dwa r f pe a all o f th e o ffsp r ing w e re t all T alln e ss

“ ,
.
,

the re fo re i s s a id to b e a d o min a nt ch a ra ct e r ( D) a nd dwa r f

,

n e ss a r e ce ssive ch ar a cte r ( R) .

Wh e n th e hyb rid s w e re all o we d to fe r til iz e th e m se lv e s th r e e ,

fo u r th s of th e i r o ffsp ring w e r e t all ( D) a nd o ne fo u r th d wa r f

( R) ; th e r e w e r e no int e r m e dia t e fo r m s Wh e n th e se dwa r fs w e re
.

a ll o we d t o fe r tiliz e the m se lv e s o nly dwa r f o ff sp ring re s ult e d .

,

Wh e n th e t a ll p e a s w e r e a ll o we d to fe r tiliz e the m se lve s o ne thi r d ,

o f th e m p r o du ce d t a ll o ff sp r ing whi ch wh e n inb re d p r o duc e d

,

o nly t a ll p e a s
; t h e o th e r t w o thi r d s wh e n inb r e d p r o du ce d th re e
fou r ths t all ( D) a nd o ne fo u r th dwar f ( R) Off sp r ing T h e r e s ult .

U
2 90 AN I NTRODU CTION TO ZOOLOGY

m ay b e xp re s se d di ag ra mm atically a s in Figur e 1 5 7
e

M a ny o th e r pl a nt s a nd num e ro u s anim al s h av e b e e n e xp e r im e nt e d
with ; in so m e ca se s M e nde l s law h a s b e e n c o nfi r m e d ; in o th e r
’

ca se s th e inhe rit a nce i s app a r e ntly d e te r min e d in a diffe r e nt way .

T o a cc o u nt f o r th e inh e r it a nc e o f ch a r a ct e r s in th e M e nd e li a n
r a ti o o f th r e e d o min a nt s ( D) to o ne r e c e s siv e ( R) th e th e o r y o f ,

th e se g r e g a ti o n o f g e r m c e ll s h a s b e e n p r op ose d Thi s i s th a t .

th e ge r m ce ll s o r g a m e te s p r o duce d by c r oss b re d o r g a ni sm s m ay -

in r e sp e ct o f giv e n ch a r act e r s b e o f th e pu re p a r e nt al typ e s a nd ,

co ns e qu e ntly inc ap a bl e o f t r a n s mitting t h e o pp o s it e ch ar a ct e r ;

th a t wh e n s uch pu re simil ar g a m e te s o f opp osit e s e x e s a r e unite d

in fe r tili z a ti on th e individu al s so fo r m e d a nd th e i r p ost e r ity a r e
,

fre e from all t a int o f th e cro ss ; th a t th e r e m ay b e i n shor t pe r ,

f e ct or alm o st p e r fe ct di s co ntinuity b e tw e e n th e se g e r ms in r e
sp e ct o f o ne o f e a ch p ai r o f Opp osite ch ara ct e rs ( B a te son) .

TALL 2 ARF 6 2 TALL

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FI G 1 5 7
. . D i a gr a m h o wi ng th e i nh e r it a nce o f t a ll ne ss a nd d wa r f ne ss i n pe a s
S

a cco r d i ng t o M e nd e l s La w ( M ifi d f t T h m on )
’
. o d e a e r o s .
292 AN INT R ODU CTION TO Z OOLOGY

(c) P a le ontology tudy o f fossil a nim al s h as b rought

— T he s

ou t som e r e m ar k ably c onvincing e vid e nc e of o rgani c e voluti o n .

FI G . 1 58 . Sk e l e t o ns of th e fo re li m bs of v a r io us ve r t e br a t e s a wi ng o f
.
,

b i r d ; b, f o r e le g of d og ; c, a r m o f m a n ; d wi ng o f b a t
,
.
( Fro m M e t
calf ) .

Tabl e X V o n p age 2 8 0 Sho ws tha t in the o lde s t s t rat a of the e a rth s ’

c r us t co nta ining o rga nic re m ain s th e C a mb ri a n a nd Silu ri a n , ,

o nly inv e r t e b r a t e s e xi s t e d To war d th e e nd o f th e l a tte r p e ri od

.

fi she s m a d e the i r app e a r ance the n a mphi bi a n s re ptile s m am, , ,

m al s a nd fin ally M a n E a ch o f the se g ro up s i s o f a high e r o r de r

,
.

th a n the l ast a nd th e sp e cie s in e ach g ro up b e come m ore

,

co m pl e x a s th e p re se nt e ra i s app roache d a fa ct no t shown in ,

th e t abl e .

E v e n m o r e co nvin cing a re th e fa ct s r e ve a l e d by a s tudy o f th e

a n ce s t or s o f a Singl e sp e ci e s s uch a s th e co mm o n h o r s e
,
E qu u s ,

( Fi g . T h e h o r se s o f th e E o c e n e p e ri o d ar e re p r e s e nt e d by
Orohi ppu s a n a nim al p o sse ssing f o u r t oe d fo r e fe e t a nd th r e e t oe d
,
- -

hi nd f e e t Thi s fo r m w a s r e pl ace d by M e s ohi ppu s in th e Low e r

.

M i o ce n e ; the n P r otohi ppu s a pp e a r s i n th e Lo we r P li o c e n e P li o ,

hi ppu s in th e P li o c e ne a nd fin ally th e E qu u s o f r e ce nt tim e s

,
In .

th e se forms the re h as b e e n a gra dual de ge n e ra ti o n of c e r t a i n

 GEN ER AL Z OOLOGI C AL F AC T S AN D T HEOR E S I 29 3
-

toe s and a corre sp onding inc r e ase in th e

,

Siz e o f o ne t oe until an a nim al E qu u s i s

, , ,

e v o lv e d with o ne to e o n e a ch f oo t a nd th e

splintlik e re m a in s o f wh a t we r e functio n a l
t oe s in its E o c e n e ance s tor .

D r i bu t i o n
— Th e
( )
d G e og p
r a hi ca l i s t .

g
f acts o f e o g r aphi c al di s t r ibutio n all se e m
to Sho w th a t sp e ci e s o rigin a te in so m e par
t icu la r pl a c e fr o m w hi ch th e y di sp e r s e a s
,

the ir numb e rs inc re a s e Wh e n g r o up s are .

p r e ve nt e d fro m m ingling w ith o the r s o f

th e ir k ind by e nvi r o nm e nt al b arr i e r s or
physi ol ogic a l c o nditi o n s th e fa ctor o f i sol a ,

tion c o mbin e d w ith o th e r fa ct or s r e sult in

, ,

the e v oluti o n o f ne w g e o g r a phi ca l r a c e s a nd

fin ally ne w sp e ci e s Acc o r dingly wh e n
.
,

sp e ci e s th a t r e s e mbl e e ach o the r cl o se ly a re

fo und o ccupying ne ighb o r ing r e gi o n s but ,

s e p ar at e d by b arri e r s w e c o nclud e th a t ,

th e y h av e e vo lve d fr o m a c o mmo n a nce s

t o r a nd h a v e div e r g e d until th e i r di ffe r
,

e me e s a r e c o n sid e r e d o f sp e cific r a nk .

T HE T H E OR Y OF N AT U R AL SE LE C TI ON .

Th e o r igin o f sp e ci e s a s c o nc e iv e d by ,

Dar win is l a r g e ly d u e t o n a tu r a l s e l e ctio n

,
.

We m ay g ain a cl e a r id e a o f w h a t i s m e ant
by n a tu ra l s e l e cti o n if w e e x a min e th e r e
sul ts o f a r tifi ci a l se l e cti o n F o r e x a mpl e .
,

the r e a r e a t th e p r e se nt tim e o ve r o ne hun

dre d a nd fifty diffe r e nt d om e s ti c va r ie ti e s o f
pige on s ; a f e w o f th e s e a r e sh own in F igur e
FI G .
5 9 D i agr a m s ill u s t r a ti ng th e gr a d u a l ch a nge s
1 .

i n f o o t s t r u ct u r e i n f o ss il a nd r e ce nt spe ci e s o f
th e ho r se f a m i ly . a, b o ne s Of f o r e f oo t ;
th e
b, b o ne s of th e hi nd f oo t .
( Fr o m M e t ca l f a f t e r
M arsh ) .
2 94 AN INTRODU CTION TO Z OOLOGY

1 60 . Th e y d iffe r wid e ly fro m o ne a n o th e r I n s t r uctu r e a nd habit s ,

but in m a ny c ase s m a y be t r a ce d b ack t o a singl e a nce stor th e ,

blue rock pige o n (C olu mba li vi a) s till in e xi s t e n ce in i ts wi ld st a t e

-
,

( g
F i 1.6 0 ,
T h e th e or y i s t h a t th e s e pig e o n s h a v e b e e n o r igi
na t e d by th e p re se r v a ti o n f o r b r e e di ng pu rp ose s o f bir d s th a t
sho we d f a vo r abl e v a r i a t io ns .

N ow a nim al s v a r y in a wild s t a t e as w e ll a s und e r th e c a r e o f

m a n a nd wh e n e ve r a n individu a l a pp e a r s with a v a ri a ti o n o f
,

v alue in th e st r uggl e f o r e xi s te n ce thi s p a r tic u l a r a nim al b e co m e s

,

o ne o f th e fitt e s t t o s u rviv e a nd th e r e f o r e
, ,
liv e s t o t r an smit i ts
,

fa vor abl e v ari atio n to i t s o ffsp r i ng In o the r wo r ds it i s se l e ct e d

.
,

by n a tur e ju s t a s a do m e sti ca te d a ni m al with a favo r abl e va r i a

tio n i s s e l e ct e d by M a n As a n e xa mpl e w e m ay t ake th e r obin
.
,
.

We h ave sh own (p 2 8 3 ) th a t thi s bi r d r e a r s a numb e r o f y o ung

.

e ach y e a r ye t th e numb e r o f r o bin s r e m a in s p ra ctic a lly c o n s t a nt

,

f r o m y e ar to ye a r T h e m aj o r ity a re k ill e d in so m e w a v Those

. .

tha t a r e str ong e s t a nd a r e a bl e to w e a the r th e s t or m s o r a r e ,

e nd o w e d with e xt r e m e sp e e d so a s t o e s c a p e th e i r e n e mi e s a r e ,

s e l e ct e d by N a tu re whil e w e ak e r o r sl o w e r individual s a r e
,

d oo m e d to de st r ucti o n .

Anim al s b e co m e a d apt e d to th e i r s u r r o unding s th r o ugh th e

o p e r a ti o n o f thi s p ro ce s s o f n a tu ra l se l e cti o n T h e w o nd e r fully
.

a d apt e d s t r uctu re s o n t h e l e g s o f t h e h o n e yb e e f o r g a the ring

p oll e n m ay h av e e v olve d in thi s w ay .

Of r e c e nt y e a r s m a ny bi ol ogi s t s h a v e qu e s ti o n e d th e a d e qu a cy
o f n a tu r a l s e l e cti o n t o e xpl a in a ll t h e m o difi ca ti o n s o f sp e ci e s .

Thi s w a s n e ve r cl aim e d f o r th e th e o r y e v e n by Da r win f o r h e

, ,

s ays in th e int ro ducti o n t o th e Or i gi n of S pe ci e s I a m co nvin ce d

, ,

th a t n a tu r al se l e ctio n h as b e e n t h e m ain but no t th e e x clu sive

, ,

m e a n s o f m o difi ca tio n
T H E T H E OR Y OF OR TH O GE N E SI S M a ny ph e n o m e n a o f th e e v
.

o lu ti o no f o r g a ni s m s ca nn o t b e e xpl a in e d by n a tu r a l se l e cti o n F or .

e x a mpl e ,
th e d e v e l opm e nt o f s t r u ctu re s o f a pp a re ntly no a d v a n
t age to th e a ni mal s u ch a s th e a rr ang e m e nt o f th e v e in s in th e
,

wi ng s o f in se ct s a nd th e d e ve l opm e nt o f m o difica tio n s ha r mf ul

,
 G EN E R AL ZOOLOGI C AL F AC T S AN D T HEOR E S I 2 95

to th e ra ce a nd l e a ding to e xtin ctio n s t r uctur e s such a s p al e on

,

t o logi s t s t e ll us h a v e a pp e a r e d m a ny tim e s du ri ng th e hi s t or y o f
th e a nim a l kingd o m a nd i llu s t r a t e d by th e I ri sh st a g wh ose
,

a ntl e rs b e ca m e s o e n o r m o us a s t o m a k e it unfit f or e xi s t e nc e .

The r e app e ar s to b e a t e nd e ncy f o r m o dific a ti o n s to foll o w ce r ta in

d e finite p a ths a nd f or ce r t ai n s t r uctu r e s to p rog re s s in p re de
,

t e r mi ne d di r e cti o n s r e g a r dl e s s o f th e i r utility a nd w ith o ut th e ,

a i d o f n a tu r a l se l e cti o n Thi s e vo luti o n in a d e fini t e pre d e te r

.

m in e d di r e cti o n i s kn o wn a s o r th o g e n e s i s .

The re a re a numb e r Of th e o r i e s o f o r th og e n e si s T h e o ne m os t .

in f avor i s th a t whi ch a scri b e s th e co nt r ol o f mo difi ca tio ns to the

di r e ct e ff e cts o f phy sic o ch e mi ca l fa cto r s o n o rg a ni sm s So m e o f .

t h e ar gum e nt s f o r o r th o g e n e s i s a r e d e r iv e d f r o m th e f o ll o wing
phe n o m e n a : th e p re se n ce o f simi la r v a r i a tio n s in sp e cie s b e l ong
ing to o ne fa mily th e de ve l o pm e nt o f di sa dva nt age o us s tr uctur e s
,

l e a ding to e xtin cti o n (e g th e a ntl e r s o f th e I r i sh s t ag) th e limits

. .
,

o f v a r i a ti o n d ue t o th e ch e m ic a l c o mp o s iti o n o f th e b o dy th e ,

influe nce o f one o r g a n up o n a n o the r whi ch limit s v a ri a ti on and

, ,

t h e p e rsi s t e n ce o f v a r i a ti o n s in d e finit e di r e cti o n s so th a t e v olu

ti o n is no t in r a di a ting lin e s a s na tu ra l se l e ction d e m a nd s
Acc or ding t o o r th o g e n e si s c e r t a in lin e s o f d e v e l opm e nt r e m a in
,

st a ti o n ary w hi l e o th e r s a dv a nc e
, .

Th e th e o r y of o r th o g e n e si s in s o m e for m o r o th e r i s a cc e pt e d
bv m a ny sp e ci a li s t s w o r king in wid e ly s e p a r a t e d fi e ld s Of th e se
,
.

m ay b e m e nti o n e d E im e r ( 2 3 9 butt e rfli e s bi r d s a nd liz a r d s)

, , , ,

Whi tm an ( 2 80 pige o n s) To we r ( 2 70 b e e tl e s) a nd R uthve n ( 2 66

, , , , ,

sn ake s) Th e p ap e r s o f the se m e n fu rni sh e x ce ll e nt co ncre te

.

e x a mpl e s o f h o w o r th o g e n e s is is s upp o s e d t o Op e r a t e .

T H E M U TATI ON T H E OR Y A mut a nt i s a n o rg a ni s m th a t dif

.

f e r s f r o m i ts p a r e nt in ce r t a in w e ll m a r k e d ch ara ct e r i s ti cs which it
-
,

t r ansm its to i ts off sp r ing De Vr i e s i s th e for e mos t e xp o n e nt o f

.

th e mut a ti o n th e o r y a nd it i s h i s w o r k th a t h a s giv e n it w or ld wid e

,
-

re co gniti o n . T h e co nclu si o n s o f d e Vr i e s a r e b a s e d o n a s tudy o f

the e v e ning p r im ro se ( ( Enothe r a la m a r cki a na ) T h e mut a ti o n.

th e or y m ay be st a te d in his wor ds a s fo ll o ws : Th e w a y in which

2 96 AN INT R ODU C T I ON To Z OOLOGY

o ne sp e ci e s or igi n a te s fr o m a n o the r h as no t b e e n a de qu a te ly
e xpl ain e d . T h e cu rre nt b e lie f as s um e s th a t sp e ci e s a re sl o wly
change d into ne w typ e s In contradi ction to thi s co nc e ptio n the
.

th e or y o f mut a ti o n a ssu m e s th a t ne w sp e cie s a nd v a ri e ti e s a re

p ro duc e d fro m e xi s ting for m s by s udde n l e aps The se m ay a ri se .

s i multan e o u sly a nd in g ro ups or s e p ara t e ly a t m or e or l e ss wide ly

,

distribute d p e ri o ds
Whi l e di re ctly o pp ose d to Dar win s the ory o f th e origin o f s pe’

cie s in r e g a r d to the k inds of v a r i a tio n s se l e ct e d d e Vr ie s give s ,

Dar win c r e di t f or th e di sco v e ry o f th e g re a t p rincipl e which

r ul e s th e e v o luti o n Of o r g a ni s m s It i s th e p ri ncipl e of n a tu ral
.

se l e ctio n It is th e sifting o u t o f a ll org anisms of min o r wo r th

.

th ro ugh th e s t ruggl e f or life It i s o nly a si e ve a nd no t a fo r ce

.
,

o f n a tu r e ,
no di r e ct c a u se o f imp r o v e m e nt a s m any o f D a r ,

Win s a dv e r s a r i e s a nd unfo r tun a t e ly m a ny o f h i s fo ll ow e r s a l so

’
, ,

h a v e so o ft e n a sse r t e d It i s o nly a si e v e which d e cide s whi ch

.
,

is to liv e a nd wh a t i s t o d i e
,
B y t hi s m e a n s n a tu r a l
.

s e l e cti on i s the o ne dire cting c a u se o f th e b roa d lin e s o f e voluti o n

( 7
2 2
,
p
. D e V r i e s cl a im s h o w e v
,
e r th a t s p e ci e s a,
nd v a r ie
ti e s ha ve o rigina te d by mut a tio n a nd a re a t p re se nt no t kn o wn
, , ,
”
to o ri gina t e in a n o th e r w a y .

T wo sor t s o f sp e ci e s a r e r e cognize d by d e Vr i e s the s y s t e m ,

a tic sp e ci e s o f t h e s y s t e m a tis t a nd e l e m e nt a r y sp e ci e s which a r e

,
“
m o re num e r o u s a nd a r e de scrib e d a s a ny fo r m which r e m a ins
,

”
co n s t a nt a nd dis tinct fro m i ts allie s ( 2 7 2 p E l e m e nt a r y
,
.

sp e cie s h av e unit cha r acte rs whi ch a r e indivi sibl e a nd di s tinct

in inhe rit anc e All o r g ani sm s a re no t mutabl e t o th e sa me de g re e
“ ,
.

a nd t o Ob se r ve t h e o rigin o f sp e ci e s It i s o nly n e ce ssary to

,

h a ve a pl ant in a mutabl e co nditio n ( 2 7 2 p ,

.

Sinc e th e public a ti o n o f d e Vr i e s r e sults biol o gi s ts h a v e e x a m

’

i ne d th e d a t a fr o m a l a rg e numb e r o f e xp e rim e nts but o nly a ,

co mpara tiv e ly s m a ll a mo unt o f p r oo f is a va il abl e to supp or t the

th e o ry fr o m th e z ool ogica l s ide .

P e r h ap s th e b e s t a ttitud e t o t a k e t o w a r d th e th r e e th e o r i e s
o utline d in th e l a s t f e w p a g e s i s th a t o f P ro fe s sor Whitm a n wh o ,
 2 98 AN I NTRODU CTION TO Z OOLOGY

One m or e illustr a ti on m ay b e cite d to Sh ow h ow r e m ark abl e

t h e r e l a ti o n s b e tw e e n ce r t a in pl a nt s a nd a nim a l s m ay b e co m e .

T h e fl o we rs Of th e ge nu s Yu cca d e p e nd up o n a singl e sp e ci e s o f
in se ct th e P r o nub a m o th f or the i r cross p o llin a ti o n This little
, ,
-
.

white m o th vi sits th e fl o we r s in th e e v e ning It scra p e s som e .

p oll e n fro m a s t a me n h o lds it unde rn e a th i ts h e a d a nd ca rri e s it

, ,

to a n o th e r fl o w e r It clings to th e pi s til o f this a nd th r us ting

.
, ,

i t s o vip o sito r th ro ugh t h e w a ll o f t h e o v a r y l a y s a n e gg It th e n ,

.

m o unts th e pi s til a nd fo r ce s th e p o ll e n it h a s b ro ught d o wn int o

,

th e s tigm a tic tub e An o th e r e gg i s l a id in a n o th e r p a r t o f the

.

o v a r y a nd m o r e p o ll e n i s in se r t e d int o t h e s tigm a ti c tub e

,
Th e se .

p r oce sse s m a y b e r e p e a t e d h alf a do z e n tim e s in a singl e flo we r .

T h e a dva nt a g e to t h e fl o w e r i s o f co u r se th e ce r t a inty o f b e ing

, ,

cr o ss p o llin a t e d a nd of p r o du ci ng se e d s Th e se se e ds p rovide a
-
.

s upply o f f oo d f o r t h e l a r va th a t h a t ch f r o m th e e gg s l a id by th e

m o th in th e o va ry T h e se e d s a r e SO num e r o us th a t th e f e w e a te n
.

by th e l a r va m ay w e ll b e sp a r e d

c . P a r asi ti s m

A p ara sitic a nim al i s o ne th a t live s up o n a n o th e r o rg a ni sm .

Thi s 1 5 a l so a d e finitio n o f a p r e dace o us a nim al We m ay .

di s tingui sh b e tw e e n th e t w o h o we v e r by a ss umi ng th a t p a r a sit e s

, ,

a r e a lw a y s c a r r i e d o n o r in t h e b o di e s o f th e i r vi ctim s whe r e a s ,

p r e d a ce o us a nim a l s a re fr e e living T h e fl e a s a r e comm on

- .

e x a mpl e s o f e xt e r n a l p a r a s it e s T h e m a l a r i a p a r a s it e ( p 88
. .
,

F i g 4 2 ) a nd th e r o und wo r m (p 1 6 0 Fi g 8 1 ) a re i nt e r n al p a r a
. .
,
.

site s Th e lif e his tor i e s o f th e s e sp e ci e s a s giv e n o n th e p age s

.
,

r e fe r r e d to will s uff ic e t o illu s t r a t e p a r as itic h a bit s but ce r t a in

, ,

ch a r a ct e r i s tic s r e s ul ting fro m thi s kind o f lif e will b e p o int e d o u t

b r ie fly .

P a ra sit e s a s a r ul e a r e co mp a r a tiv e ly simpl e in s t r uctu r e This .

i s du e t o d e g e n e r a ti o n a nd no t to a lo w p o s iti o n in th e a nim al
kingdo m In m any c a se s th e r e p r o ductive o r g a n s a r e e x ce e d
.

i ngly w e ll d e v e l o p e d wh e r e a s o r g a n s o f l oco m o ti o n dig e s ti on

-
, , ,

e tc.
,
b e com e a lm os t f uncti onle ss .
 G EN E R AL Z OOLOGI C AL FACT S A N D T HE OR ES I 2 99

FI G . 1 61 . l a r va l s t a ge s o f a pa r a s it e S a ccu li na ca r ci na A N a upli us;

T he , .
,

B C ypr i s ; C C ypr i s a tt a ch e d to h o s t ; D E F s t a ge s i n d e ge ne ra ti o n
, , , , ,

d u e to pa r a s iti c h a b it : ( Fr o m Se d gwi ck a f t e r De l a ge )
.

The life hi s t o r y o f th e c r u s t a ce a n S accu li na carci ni a p ara sit e

, ,

o n a n o th e r cr u s t a ce a n th e cr a b C a r ci nu s m a e nas m a y b e co n
, , ,

si d e r e d a g o o d illu s t r a ti o n o f t h e d e ge n e ra tio n o f p a r a sit e s .

S accu li na h a t ch e s as a N au pli u s ( F i g 1 6 1 A) whi ch so o n ch a nge s

.
, ,

t o th e Cypr i s s t ag e ( B ) Aft e r s wimming a b o ut f o r t w o o r th r e e

.

it se lf t o th e se ta o f a cra b by m e a n s o f i ts
a nt e nna T h e b a s e o f th e s e t a is p e n e t ra t e d by th e h oll o w
3 00 AN IN TRODU CTION TO Z OOLOGY

a nt e nn a a nd th e l a r val S accu li na sh e ds i ts app e nd ag e s She ll

, , ,

muscl e s a nd e x cr e t or y o rg a n s Wha t r e m a in s (E ) p asse s

,
.

thr o ugh th e a nt e nn a into th e b o dy of th e cr ab (F ) a nd fin ally ,

r e a ch e s th e b o dy c a vity A s ys t e m of r oo t s no w g r o w s o u t in a ll
.

di r e cti ons Whe n th e c r a b m olt s th e saclik e b o dy o f th e p ar a

.
,

sit e p e n e t r at e s to th e o utside re se mbling a tumo r c o n si sting ,

a lm os t e nti re ly o f r e p r o ductiv e o r g a n s

( )
d S oci al Life

Th a t th e r e is a c o ntinu a l c omp e titio n a m ong individu als and

Sp e ci e s h a s b e e n p oint e d o u t in th e c o u r s e o f o u r di s cus si o n o f

g e og raphica l di s tr ibuti o n (p a nd th e s t r uggl e f o r e xi s t e n ce

.

(p 2 8
. Ho w e ve r th e re i s a n o the r la w o f n a tu r e n a m e ly
, ,

s oci ability no t so f a r re aching p e rh ap s but o f undo ubt e d i m

,
-
, ,

p or ta nce to th e w e lfar e o f m a ny a nd di ve r se sp e ci e s o f a nim a l s .

C e r t ain o f th e so ci a l sp e ci e s h a v e a l re a dy b e e n m e ntio n e d (p .

a nd th e a ctiviti e s Of th e h o n e yb e e a n in s e ct with a v e ry ,

c o mpl e x so ci a l life ha ve b e e n d e scrib e d in so m e de t ail (p

,
.

T h e b e a v e rs R ocky M o unt a in sh e e p a nd p r a i ri e d ogs ( m a m

, ,

m al s) the swallo ws he ro n s a nd se a gull s (bi r ds) a nd th e an ts

, , , , ,

wa sps a nd t e rmit e s ( in se ct s) a re fam o us f or th e i r co mmunity

,

life .

T h e a dva nt a g e s Of living t og e th e r are m any a nd th e live s of ,

m os t o f th e g re ga ri o us a nimal s d e p e nd up o n the i r in stinct to

fo rm b a nds a nd co l o ni e s Da r win r e cog niz e d th e v alue of soci al
.

life altho ugh hi s the o ri e s we re b ase d l a rg e ly up o n th e e ff e ct s o f

,

th e s t r uggl e f o r e xi s t e nc e He s ays th a t th e ind ividu al s whi ch

.

t oo k the gr e a t e s t pl e as ure in s o cie ty wo uld b e s t e sc ap e v ari o us

d a nge rs ; whi le tho se tha t c a re d l e a st f or the i r c o mra d e s a nd ,
”
live d s olit ary w o uld p e ri sh in g r e a te r numb e rs
,
De e r f o r .
,

e x a mpl e wh e nb a nde d t oge the r m ay e scap e f ro m th e i r e n e mi e s

, ,

whe re as s ingl e indi vidual s wo uld b e de st roy e d T h e R ocky .

M o unt a in sh e e p p os t se ntin e l s wh ch wa r n th e m ain h e r d o f th e

'

a pp r o ach o f d a ng e r a nd e n abl e a ll t o s e e k pl a c e s o f s a fe ty
,
.
3 02 AN INT R ODU CTION TO ZOOLOGY

lu ti o n, b o th di re ctly by se cu ring the we ll b e ing o f the sp e ci e s

,
-

whi l e di m ini shing th e was te o f e n e rgy a nd indi r e ctly by fa vo ring ,

th e g r o wth o f int e llig e nc e The re fo r e co mbin e

. p ra ctic e
mutu al ai d Tha t i s the s u re st m e a n s o f giving t o e ach a nd to
.

a ll th e g r e a t e s t s a f e ty t h e b e s t gu a r a nt e e o f e xi s t e nc e a nd p r o g
,

r e ss b o dily inte ll e ctu al a nd m ora l Th a t is wha t n atu re

, , .

”
t e a ch e s u s .

3 . R E F LE X E S , I N S TI N C TS ,
AN D TH E AN I MAL M I N D

a . R efle xe s

In the M e ta z o a th e r e fl e x c arri e d on e ith e r co nscio u sly o r u m

,

c o n sci o u sly i s co n sid e r e d th e physi ol o gi ca l unit o f n e r vo us a c

,

t i v i ty . It re qui re s in i ts simpl e s t fo r m a se n so r y n e u ron the ,

re ce pt o r a m o t o r n e u r o n th e a dj u s t or a nd th e o r g a n s timul a t e d
, , ,

t o a ctivity th e e ff e ct o r ( se e p
,
T h e s timulu s p a sse s f r o m
.

t h e re ce pt or to th e a dju s t or a nd i s r e fl e ct e d to t h e e ff e ct or Thi s .

a pp are nt r e fl e cti o n sugg e s t e d th e t e r m re fle x .

Th e re fl e x a cc o r ding to th e a b o v e id e a i s o p e r a tiv e o nly in the

, ,

c ase of a nim al s with n e r vo u s sy st e m s If w e w o uld in clude .

c e r ta in l o we r M e t a zo a a nd th e P r o to z oa ou r d e fi ni ti o n mus t be ,

ch ange d so a s to t ake into a cco unt th e a cti on a nd not ,

m e cha nism th a t i s r e sp o n sibl e f o r th e a cti on Acco r ding t o .

vi e w a ny simpl e r e sp o n se t o a s timulu s i s a r ef le x Th e avoi

, .

r e acti o n o f P a r a me ci u m ( Fi g 3 3 p 7 4) i s kn o wn a s a r e fle x
.
,
.
,

th e a ctiviti e s o f P a r a me ci a a nd m a ny o th e r l o w e r o r g a ni s ms
s upp o s e d by s o m e t o b e a s e r i e s o f r e fl e x e s t o e xt e r n a l s tim

B u t if e x a min e d c a r e fully th e r e a cti o n s o f l o w e r org a ni sm s

,

no t a s s i m ple a s th e y a t fi r s t a pp e ar F o r e x a mpl e th e a v oiding

“ ,
.

r e a cti o n o f P a r a me ci u m i s no t a lw a y s e x actly t h e s a m e If .

t h e a nim a l b e t a k e n a s a ce nt e r a b o ut which a sph e r e i s d e s crib e d ,

w it h a r a diu s s e v e r a l tim e s th e l e ngth o f th e b o dy th e n a s a r e ,

s ul t o f th e a v oiding r e a cti o n t h e a nim a l m a y t ra v e r s e th e p e riph

e r a l s u r f a c e o f t hi s sph e r e a t a ny poi nt m o ving a t th e ti m e e ith e r ,

b ackwa r d or forwa r d P a r a me ci u m in spite of i ts cu I i OII s limi

'

; ,
 G E N ER AL Z OOLOGI C AL F AC T S - AN D T H EOR ES I 0
3 3

ta ti ons as to m e thod of m ove m e nt i s a s fre e to var y its r e l a ti ons

,

to th e e nvi r o nm e nt in r e sp o ns e t o a stimulu s a s a n o rg a ni s m o f i ts
for m and str uctu re c o uld co nc e iva bly be Such b e havior d oe s .

not f all withi n th e c o nc e pt of a r e fl e x if th e l a tt e r is d e fin e d a s a

uni for m re actio n ( 2 4 6 p ,
.Je nning s h as p o int e d o u t tha t
b e fo r e th e r e fl e x ca n b e co n sid e r e d a simpl e inva ri abl e unit f or
b e havi or the physi ol o gic a l co nditi o n o f th e o r ga ni sm must b e
,

t ak e n into a cco unt Such a unit might b e illu s tr a te d a s foll ows :

.

stim ulu s A a cting up o n a n a nim al who s e physio l o gic al st a te is

,

B give s r e a ction C
,
.

b I ns ti nct
.

Instinct is th e fa culty o f a cting in such a w ay a s to p ro duce

c e r t a in e nds with o ut f o r e s ight o f th e e nds a nd with o ut p re vi o us
, ,

e duc a ti o n in th e p e r f o r m a nc e ( J a m e s ) Instinctive acts a r e

.

all c o mpl e x The y ar e p e r for m e d in a Simil ar m a nn e r by a ll

.

m e mb e r s o f th e sa m e s e x and ra ce Tho s e Of the s ocia l a nim als

.

a re r e m a r k a bly v a r i e d . Our study o f t h e h o n e yb e e ( C h ap XII) .

has give n u s suffi ci e nt kn o wl e dg e o f thi s inse ct t o w a r ra nt o ur

using it to illu st r a te c e r t a in kind s o f in s tinct s .

T h e qu e e n s nupti a l flight a nd h e r co pul a tio n with th e d ro n e is

’

in stinctiv e Sh e a cts with o ut p r e vi o u s e xp e ri e nc e a nd a e c o m

.
,

pli sh e s th e filling o f h e r sp e r m a th e c a with Sp e r m a t o z o a with o ut

app a r e ntly r e a lizi ng th e pu rp o s e o f th e wh o l e p e r fo r m a n ce The .

se l e ct i o n o f a h o ll o w t r e e by w o r k e r sc o ut s a nd th e cl e a ning o f it
,

in p re p a ra ti o n f o r th e sw ar m i s in s tinctiv e T h e m any a ctivi tie s

,
.

tha t foll o w th e a dv e nt int o a ne w h o m e s u ch a s c r e vi ce chinking

, ,

va rni shing w a x f o r m ing co mb b u ilding e gg la ying ga the ring

, , , ,

a nd st o r ing p o ll e n a nd n e c t a r r ip e ning h o n e y gu a r ding cl e a n i ng

, , , ,

a nd v e ntil a ting th e hiv e a nd ca r ing f o r th e qu e e n a nd y o ung

,

be e s ar e th e r e sult s o f instin cts

, .

Th e in s tinct o f c a r ing f o r t h e y o ung i s o f sp e ci a l int e r e s t Sinc e ,

those a nim al s th a t ca r e f or th e i r yo ung succe e d in r e a ring a l a rg e r

proportio n tha n those th a t all o w th e i r offsp ring to shi ft f or the m a
3 04 AN INTRODU CTION TO ZOOLOGY

se lve s . T hedi vision of l ab or a mong th e work e r s is d u e to i n ~

s ti nct ; al th o ugh e a ch i ndividu a l i s e quipp e d with th e m a ny

str uctu ral fe a tu re s th a t s o w ond e r fully a dapt it to its mo de of

life onl y one s or t o f w o r k i s und e r tak e n a t a ti m e and thi s in all
, ,

c a s e s i s th e thi ng m os t n e e d e d by th e hi ve F or e x ampl e as .
,

s o on a s a ho ll o w t re e i s se l e ct e d a nd cl e an e d th e work e r s p ro ce e d
,

t o s e c r e t e w a x a nd build c o mb ; wh e n e gg s a re l aid p oll e n i s ,

g a the re d a nd s to r e d with whi ch t o fe e d the fo r thco m ing l a r va ;

dur ing the p oll e n a nd n e cta r ga the ring se a so n e ach w ork e r col ,

l e cts e ithe r th e one or the o th e r a nd th e p o ll e n g a th e r e d on one

,

t rip is all o f o ne col o r h a ving b e e n c olle cte d e vide ntly fro m one
,

kind o f flo we r Alt r u i s tic in s tinct s s uch a s th e se a re e sp e ci ally

.

a bund a nt in s o cia l a nim a l s a nd a m ong th e se m a y b e cl a sse d the

,

so ci al in s tinct it se lf On th e o the r h a nd the instincts o f f e e di ng

.
, ,

o f s e lf d e f e n s e
- o f b a ttling f o r t h e sup r e m a cy o f th e hiv e
,
a nd o f ,

pl aying b e fo r e th e hi ve o n b right d ays m ay be said t o be e g oi stic

,
.

The se d e finiti o n s and e x a mpl e s o f in s tinctiv e a cts a r e not

a c ce pt e d by a ll sci e ntist s ; a nd o f re c e nt y e a r s th e r e h a s b e e n a

t e nde ncy to c on side r th e m a s t rOpi c r e sp o n se s t o s timuli Thu s .

o ne a uth o rity a tt r ibut e s a ll t h e a ctiviti e s o f a nt s a nd b e e s t o

r e fl e x r e sp o n se s t o ch e m i ca l a nd o th e r s timuli T h e e mi
n e nt physiol ogi s t Lo e b sa y s “ M y inve s tig a ti o ns o n th e h e li o
t ropi s m o f a nim al s le d m e to a n alyz e in a f e w c a se s th e c o nditions
whi ch d e t e r min e th e app a re ntly a c cide nt al dir e ctio n
m ove me nt s whi ch a cco r di ng t o t r a ditio n a l n o tio n s
, ,

v olunt a ry or in s tin ctiv e Wh e r e ve r I h av e thu s f ar inv e stigat e d

.

t h e ca u s e o f such volunt a r y o r instinctive m ov e m e nt s in

‘ ’

a nim al s I h a v e with o ut e xc e pti o n di sco v e r e d such ci r cu m

,
s t an ce s

a t w or k a s a r e kn o wn in i na nim a t e n a tu r e a s d e t e r min a t e m o v e

m e nts B y th e h e lp o f the se ca u se s it i s p ossibl e to c o ntrol the

.

volunta ry m ove m e nt s o f a living a nim al ju s t a s se cu r e ly and

un e qui vocally a s th e e ngin e e r h as b e e n a bl e to c o ntr ol th e move
m e nts i n in a ni m a te n a tu re Wh a t h as b e e n t a k e n f or th e e ff e ct
.

Of will or in s tin ct i s in r e ality th e e ff e ct o f light of gr avity

’ ‘
, ,

o f f ricti o n o f ch e mic a l f or c e s
,
e tc ( 5
2,
2 p . A cc o
,
r ding.
3 06 AN INTR ODU CTION TO Z OOLOGY

pl as tic i ndividual d e lib e ra tio n s o r a d apt a tions ( 240 p ,

.

Oth e r a uth or iti e s c on sid e r th e p o we r t o l e a r n by indi vidual

e xp e ri e nc e f or e x a mpl e th e h abit f o r mi ng in th e c ra yfi sh ( p
, ,
-
.

a s e vid e nc e Of mi nd And s till o th e r s d e ny th e p r e se nce of mind

.
,

a tt r ibuting e v e r y act to ch e mic o phy s ic a l p r o c e ss e s .

It ce r t a inly i s t r ue th a t no one kn o ws wh e the r anim als a re

consci o us o r not Ou r e vide nce is a ll g a ine d by infe re nce s fr o m
.

b e h avi or Th e c onditi o ns unde r which th e fa cts of b e ha vior are

.

o bt ain e d a r e o f g r e a t imp or t a nc e An e cd o t e s a r e of littl e va lu e

.
,

s inc e th e o bs e r v e r i s no t sci e ntific a lly t r a ine d a nd wi shi ng t o t e ll , ,

a g o o d st o r y t e nd s t o e x a gge r a t e th e a pp ar e nt int e llig e nc e of the

,

a nim al . T h e m o d e r n m e th o d i s th a t o f c a r r ying ou t d e finit e

e xp e r im e nt s th e h abits o f th e sp e ci e s a nd p a st e xp e r i e nc e s o f th e
,

indi vidua l s b e ing kno wn T h e c on clu sio n i s th e n b ase d o n the

.

l e a st c o mpl e x inte rp re ta tion th a t wi ll a cco unt f or th e fa cts of

b e h a vi or o b se r ve d We can ho w e ve r o nly int e rp re t b e h avi or
.
, ,

o n th e a n a l ogy o f hum a n e xp e r i e nc e th e t e nd e ncy b e ing to a s

,

e r ib e hum a n t r a its t o a nim a ls .

Le a r ning by e xp e r i e n ce a p r o c e ss co nsid e re d by m any indic a

,

tive o f intellige nce is o ne o f th e cr it e r i a o f th e p re s e nce o f mi nd ;

,

but th e m o di fi abi li ty of the b e ha vi or mus t be rapid o r tim e ,

e n o ugh f o r a ch a ng e in b o dy st r u ctu re m ay e l a ps e St r uctu re .

i s a distinct h e lp in infe r r ing mind s inc e a nim a ls with a n e rv ,

o us sy s t e m simil ar t o th a t o f M an must b e a ffe ct e d by e xt e r n al

Stimuli in a s imi l ar w ay .

We kn o w no t wh e r e c on scio u sn e ss b e gins in the a ni m al

wor ld We kn o w whe re it s ure ly re side s in ou rse lve
.

kn o w wh e re it e xi t b e yo nd a r e a so n abl e d o ubt i n
s s —

a nim a l s o f st r uctu r e r e se mbling o u r s which r apidly a d a pt

s e lve s to th e l e sson s o f e xp e ri e nc e B e yo nd this p oint .

,

we kn o w it m ay e xi s t in simpl e r a nd simpl e r for ms unt i l

,

th e v e ry l o w e st o f living b e ings ( 2 76 p ,
.
 B HH J O G R A P H Y
'

A fe w of th e b oo k s ti cl e s th a t h a v e b e e n co ns u lt e d i n p re p a r i ng
a nd a r

thi s w o r k will b e f o u nd i n t h e f o ll o wi ng li s t T h e y h a v e b e e n s e l e c t e d .

c hi e fl y with r e f e r e nce t o th e i r a c ce ss i b ilit y a nd v a l u e t o s t u d e nt s a nd ,

a r e a r r a ng e d a l p h a b e ti ca ll y u nd e r t h e h e a d i ngs o f t h e v a r i o u s c h a p t e r s .

T h e i r nu m b e r s co r r e s p o nd i ng t o nu m b e r s i n p a r e nth e s e s i n t h e t e x t
, ,

Wi ll f a cilit a t e t h e fi nd i ng o f a ny p a r ti cu l a r so u r ce o f i nf or m a ti o n .

C H AP T E R I

I NTR ODUC T ON I
As pe cts of th e Spe c esi Q u e s ti on . P a pe rs r e a d a t a sym po s u m a t i
C hi ca go J a n ,
. 1 ,
1 90 8 . 1 908 . Am e r . Nat .
,
v ol .
4 2, pp . 21 —
8 28 1 .

Darwi n, C .
,
1 86 1 . T h e Or i gi n o f Sp e c i e s b y m e a ns of N a t u r a l Se l e c
ti o n . Ne w Y o rk .

Har m e r , S F . .
,
a nd A E Sh i pl e y . . . E d i tors . T h e C a m b r i d ge N a t
u r a l Hi s t or y . 1 0 vol s Lo nd o n . .

I l e rtwi g, R .
,
1 90 2 . M a nu a l of Z l o o o gy . T r a ns l a t e d by J . S .

K i ng l s e y . Ne w Y o rk .

Lank e s te r , E R . . Ed it o r . A T r e a ti s e on Z l o o o gy . L o nd o n .

P ar k e r , G H . . 1 9 08 — . Z l gi loo o ca P r o gr e s s . Am e r . Na t .
,
v ol .
4 2,
p p -I
-
r s I ss
P ar k e r , T .
J .
,
a nd W . A Ha s w e ll ,
. 1 89 7 — . T e x t -b o o k of Z l oo o gy .

2 vol s . L o nd o n .

8 Th om son,
.
J . A .
,
1 90 6 . Ou tli ne s of Z l o o o gy .
4 th e d . Lo nd o n .

9 . Willi s ton , S . W .
,
1 90 8 . Wh a t is a Sp e ci e s ? Am e r N a t . .
,
v ol .
4 2,
- 8 —1
Pp 1 4 94 .

10 . Z i tte l , K . A . vo n, 1 9 00—1 9 0 2 . T e x t -b o o k of P a la o nt o lo gy . 2 vo ls .

T r a ns l a t e d b y C R E a s t m a n . . . L o nd o n .

C H AP T ER I I

P HEN OM E NA OF LIF E

B ii ts chli , O .
,
1 894 . I nv ti g ti e s a o ns o n M i cr OS C Op i c F o a m s a nd o n

P r o t o pl a s m . L o nd o n .

Dah l gr e n, U . W A
, a nd . . K e pne r ,
1 90 8 .
— T e x t -b ook of t he P ri n
ci p le s o f Ani m a l H i s t o l o gy . Ne w Y ork .

3 07
3 08 I
B BL OGRAP H I Y
Le e ,
F S . .
,
1 90 8 . P hys i o l o gy . Am e r N a t . .
,
v ol .
4 2, pp .
- 1
3 94 4 7
Lo cy, W . A .
,
1 90 8 . B i o l o gy a nd i ts M a k e rs . Ne w Y ork .

Lo e b , J .
,
1 90 6 . T h e D yna m i cs o f L ivi ng M a tt e r N e w Yo r k . .

S e d gwi ck , W . T , a nd E B
. . . Wi l s on 1 8 99 G e ne r a l B i o l o gy
,
.
.

2d e d Ne w . Y or k .

Ve rworm , M .
,
1 899 . G e ne r a l P h ys i o l o gy . T r a ns l a t e d b y F . E .

Le e . L o nd o n .

Wi l son E B 1 8 99 , On P r o t o p l a s m i c St r u c t u r e i n t h e E ggs o f
. .
,
.

E chi no d e r m s a nd So m e Oth e r Ani m a l s J o u r n o f M o r p h v o l 1 5 ,

. .
,
.
,

S u pp l p p 1 —2 8 .
,
. .

1 9 00 — The C e ll i n D e v e l o p m e nt a nd I nh e r it a nce
.
2d .

e d . Ne w Y ork .

1 90 8 . B i o l o gy . Ne w Y o rk .

C H APT E R I I I

THE CELL AND THE C ELL THE ORY

Dahlgr e n U ,
W A K e pne r 1 9 0 8 — T e x t bo o k o f t h e P r i n
.
, and . .
,
.
-

ci ple s o f Ani m a l H i s t o l o gy N e w York . .

Far m e r , B — T h e S t r u c t u r e o f Ani m a l a nd Ve e t a b l e
J .1 0
9 3 .
, g .

C e ll s I n La nk e s t e r s T r e a ti s e o n Z o o l o gy P a r t I 2 d f a sci cl e
’
.
,
.
,

L o nd o n .

H e rtwi g O ,
.
,
1 895 . The C e ll . T r a ns l a t e d b y M . C a m pb e ll a nd

e d it e d b y H .
J . C a m p b e ll . L o nd o n .

Lo cy, W . A .
,
1 908 . B i o l o gy a nd i ts M a k e r s . Ne w Y ork .

Wi l son ,
E B . .
,
1 9 00 . T h e C e ll i n D e v e l o p m e nt a nd I nh e it a nce r .

2d e d . Ne w Y or k .

C H AP T ER I V

AM EB A

B e rns te i n, J .
,
1 9 00 . C h e m o t r Op i s ch e B e we gu nge n e ne s Q u e ck i
s i lb e r t r o p f e ns Ar ch f d ge s P h ys i l
. . . . . o , B d 8 0 , pp
. 6 2 —
8 63 7 . . .

B e rth ol d , G 1 8 86 — S t u d i e n i i b e r P r o t Opla s m a m e ch a ni k Le i p
'

. . .
,

zi g .

B ii ts chli , O .
,
1 8 94 . I nv ti g ti es a o ns o n M i cr OS C Op i c F o a m s a nd o n

P r o topl as m . Lo nd o n .

C alki ns , G N , 1 9 0 1 T h e P ro t ozoa
. Ne w ork . . . Y .

1 90 5
— vi
E d e nce s o f a Se x u a cyc
. l l e i n t h e L i f e -Hi s t o r
y of
Am oe b a p r o t e u s Ar c hiv f P r o ti s t , VOL 5 , p p 1 —1 6 . . . . .

1 90 7
— T h e F e r tili z a ti o n o f Am e b a p ro t e u s
. B iol B u ll . .

v ol 1 . 2 1 —2 0 3 , pp .
9 3 .
3 10 I
B B L OGRAP H I Y
B r oo k s , W . K .
,
1 90 4 . T h e F o u nd a ti o ns o f Z o o l ogy . Ne w Yo rk .

C alki ns , G N . .
,
1 90 1 . Th e Pro t o z oa . Ne w Y or k .

1 90 9 . P r o t o z o o l o gy . P h i la d .

a nd S . W . C u ll , 1 90 7 .
— T he C o nj u ga ti o n of P a r a m e ci u m
a ure li a ( ca u d a t u m ) . Ar chiv f P r o ti s t . . .
,
vol . 1 0, pp 3 7 5 4 1 5 .
— .

C ull , S A . R e j u v e ne s ce nce a s t h e R e s u lt
.
,
1 90 7 j .ti o n o f C o n u ga .

J o u r n E xp o o l , v o l 4 , pp 8 5 8 9
.
— . Z . . . .

De lli nge r , O P , 1 9 0 9 T h e C ili u m a s a Ke y t o t h e St r u ct u r e o f

. . .

C o nt r a ctil e P r o t o p l a s m J o u r n M o r p h , v o l 2 0 , pp 1 7 1 —2 1 0 . . . . . .

H am b ur ge r , C , 1 9 0 4 D i e C o nj u ga ti o n v o n P a r a m e ci u m b u r s a r i a
. .

F o ck e Ar c hiv f P r o ti s t , v o l 4 , pp 1 99 2 3 9
.
— . . . . . .

H e rt e l , E , 1 9 04 — U e b e r B e e i nflu ss u ng d e s Or ga ni s m u s d u r ch
. .

L i c ht , e t c Z e i t s ch f a ll g P h ys i o l , B d —
.
4, pp 1 43 . . . . . . .

J e nni ngs , H S , 1 9 0 1 On t h e Si gni fi ca nce o f t h e Sp i r a l S wi m

. . .

m i ng of O r ga n s m s i Am e r N a t6 — 8 v ol
.
, 3 5 , pp 3 9 3 7 . . . . .

1 9 04 . t ti A M e th
t l i h o d o f D e m o ns r a ng t h e E x e r na D s c a r ge
o f t h e C o nt r a c til e l Z l Va cu o e oo Anz , B d 2 7 , pp 6 5 6—6 5 8 . . . . . .

1 9 04 C o nt r i b u ti o ns t o t h e S t u d y o f t h e B e h a vi or o f Lo w e r
.

Or ga ni s m s C a r ne gi e I ns tit u ti o n o f Wa s hi ngt o n
. .

1 906 B e h a vi o r o f t h e L o w e r Or ga ni s m s
. N e w York . .

1 90 8 Ani m a l B e h a vi o r A m e r N a t v o l 4 2 pp 7 5 4—7 6 0
. . . .
,
.
,
. .

1 908 — T h e I nt e r p r e t a ti o n o f t h e B e h a vi o r Of t h e Lo w e r
.

Or ga ni s m s Sci e nce N S v o l 2 7 p p 6 9 8—7 1 0 .

,
. .
,
.
,
. .

1 9 0 9 — H e r e d it y a nd . Va r i a ti o n i n t h e Si m p l e s t Or ga ni s m s .

Am e r N a t . .
,
v ol .
43 , pp 3 . 2 1 —3 3 7 .

and J am i e s o n 1 9 0 2
C . M o v e m e nt s a nd R e a cti o ns o f ,
. T he
P i e ce s o f I nf u s o r i a B i o l B u ll v o l 3 p p 2 2 5—2 3 4 . . .
,
.
,
. .

J e nson P 1 8 93 ,
U e b e r d e n G e o t r o p i s m u s ni e d e r e r Or ga ni s m e n
.
,
. .

A ch
r f d g . e s P hy i l
s .o B d 5 3 pp 4 8 4 8 0
. 2 — . . . . .
, ,

Lyon E P 1 9 0 5
,
. On t h e T h e o r y o f G e o t r o p i s m i n P a ra m oe ci u m
.
,
. .

Am e r J o u r n P h ys i o l v o l 1 4 pp 4 2 1 —4 3 2
. . .
,
.
,
. .

M ai e r H N 1 9 0 3
,
. U e b e r d e n f e i ne r e n B a u d e r Wi m pe ra pp a ra te
.
,
.

d e r I nf u so r i e n
Ar ch f P r o ti s t B d 2 pp 7 3 —1 7 9 . . . .
,
.
,
. .

M a st S O 1 — T h e R e a c ti o ns o f D i d i ni u m na s u t u m ( St e i n)
,
.
909 .
,
.
,

with s pe ci a l r e f e r e nce t o t h e F e e d i ng H a b it s a nd t h e F u ncti o n

o f T r i ch o c ys t s B i o l B u ll v o l 1 6 pp 9 1 —1 1 8 . . .
, .
,
. .

M a u pa s E 1 8 88 — R e ch e r ch e s E xp ér i m e nt a l e s s u r la s M u lti
, .
, .

p li ca t i o n d e s I nf u s i o r e s cili e s
S
Ar c h d Z o o l e xp ér e t gén . . . . . .
,

2 m e se r t 6 ,
. .

M i tr e ph anow P 1 9 0 5 E t u d e s u r la s t r u c t u r e le d e v e lo ppe m e nt
,
.
, .
,

Ar ch f P r o ti s t
’
e t l e x p o s i t i o n d e s t r i ch o cys t e s d e s P a r a m e ci e s . . . .
,

E d 5 , pp 7 8—9 1
. . .
 I
B BL OGR AP H I Y 3 11

73 . Sch a e f e r , E .
,
1 904 . T h e or i e s of C ili a r y M o v e m e nt . Ana t Anz . .
,

Bd —
. 24 . PP 4 9 7 5 1 1

S ch u b e r g, A —U e b e r C i li e n u nd T r i ch o cys te n e i ni ge r I ntu
74 . .
,
1 905 .

so ri e n . Ar ch f . . P ro ti s t B d 6 pp 6 1 —1 1 0 .
,
.
,
. .

75 . Willi am s L W ,
. .
,
1 90 7 T h e St r u ct u r e o f C ili a e spe ci a ll y i n
.
,

G a s t r o po d s Am e r Na t v ol —
. . .
,
.
4 1 , pp .
545 5 5 1 .

C H APT E R VI

OTHE R P R OT O OA Z
Bl ack man, F F . .
,
1 9 00 . T he Pr m i iti v e Al ga a nd the F l a ge ll a t a .

Ann B o t v ol 1 64 7 — 6 88
. .
,
.
4 , pp . .

Bour ne , G C . .
,
1 90 9 . C o m p a r a tiv e Ana t o m y of Ani m a l s , v ol . 1 ,

2d e d . L o nd o n .

C alki ns , G N . 1 90 9 . P r o t o z o o l o gy . P h i la d .

Ho war d L O 1 — N o te s on th e M o s q u it o e s of th e U nit e d
,
. .
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St a t e s . B u ll . 25, N e w Se r i e s , U . S De p t
. . Agr i c . Di v . o f E nt o

m o lo gy .

J e nni ngs H S — Th e B e h a vi or of Lowe r Orga ni sm s Ne w

80 .
, . .
,
1 90 6 . .

Y o rk .

Ke nt ,
W . S .
,
1 88 1 . M a nu a l of th e I nf uso r a i . L o nd o n .

K e nte n , J .
,
1 89 5 . Di e K e r nt h e ilu ng von E u gl e na vi r i d i s . Z it e .

f . wi s s Z oo l . B d 6 0 , pp 2 1 5
.
, 35 . .
—2 .

Lang, A .
,
1 90 1 L e h r b u c h d e r Ve r gl e i c h e nd e n Ana t o m i e
. de r
Wi r b e ll o s e n T hi e r e ,
Bd . 1 ,
2d pt .

M i nchi n ,
E A . .
,
1 90 3 . I n L a nk e s t e r
’
s T re a ti s e on Z ol o o gy , pt . 1 ,

2d f a s ci cl e .

Ol tm anns , F — M o r h ol o i e u nd B i o l o gi e d e r Al ge n, B d
.
,
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p g .

I . J e na .

We i s m ann, A .
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1 9 04 .
— T he E v o l u ti o n T h e o r y . Lo ndo n . T r a ns
la te d by J . A T h o m so n
. .

C H APT ER VI I

I NTR ODUC T ON T O TH E I M ETAZ OA

Bove ri , T ,
D i e E.nt s t e h u ng1d e s G
89 2
e g e n s a t z e s z wi sch e n d e n
.

G e s chl e cht s z e ll e n u nd d e n s o m a ti s c h e n Z e ll e n b e i As ca r i s m e ga
lo ce ph a la . Sit z ge s f M o r p h P h ys i o l M ii nch e n, B d 8
. . . . . . .

88 . Calkins , G . N , and S C ull , 1 9 0 7

.
— T h e C o nj u ga ti o n o f P a r a m a
. .
~

li a (ca u d a t u m ) Ar ch iv f Pr oti s t Bd —
ci ur n au r e . . . .
,
. 1 0,
pp 3 7 5 4
. 1
5
3 1 2 I
B BL OGR AP H I Y
89 . Dahlgr e n, U .
, and W . A . K e pne r ,
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— P r i nci p l e s of Ani m a l
H i s t o l o gy . Ne w Y ork .

Ha e ck e r , V , 1 89 7 — Di e K imb hne a von C y cl o p s Ar ch M ik r

90 . . . . . .

Ana B d t — 1
, 4 9 , pp 3 5 9
. . . .

9 1 . Ha t sch e k , B , 1 8 8 6 Z u r E ntwi ck l u ng d e s Am p hi o xi s
. B e r li n . . .

92 . H e gne r , R W , 1 9 0 9 —T h e Or i i n a nd E a r l D e v e l o m e nt o f t h e
.
g y .
p .

G e r m -C e ll s i n So m e C h r ys o m e li d B e e tl e s J o u r n M o r p h , v ol . . . .

—
20 , pp . 23 1 2 96 .

93 H e rtwi g ,
R .
,
1 89 7 . G e ne r a l P r i nci p l e s of Z l o o o gy . T r a ns l a t e d
by G . W . Fi e ld . Ne w Y ork .

Kor s ch e lt E and K He ide r 1 8 9 5—1 9 00 — T e xt -b o o k of t he

94
‘

.
,
.
,
.
,

E m b r yo l o gy of I nv t er e br a te s .
4 vo ls . T r a ns l a t e d b y E . L .

M a rk L o nd o n
a nd o th e r s . .

M ar ti n, H N , 8 9 4 1 — T h e Hu m a n B o d y
. 6 th e d .Ne w . . . Y o rk .

M ontgom e ry, T H , 1 9 0 1 — A S t u d y o f t h e C h r o m o s o m e s . . . of the

G e r m -C e ll s o f t h e M e t a z o a T r a ns Am e r P hil So c , v o l . . . . . . 20 .

S t bh r , P
’

,
T e x t -
b o o k o f H i s t o l o gy
. 1 89 8 P hila d . . .

We i s m ann, A , 1 9 04 — T h e E v o l u ti o n T h e o r y T r a nsl a t e d by
. . .
J .

A T h o m so n
. L o nd o n . .

99 Wi l son ,
E . B .
,
1 9 00 . T h e C e ll i n D e v e l o p m e nt a nd I nh it er a nce .

Ne w Y or k .

—1 — St u d i e s on C h r o m os o m e s J o ur n of E xp
1 00 . 1 90 5 90 9 . . . .

Z l v l
oo .
,
o s .
—
2 6 .

C HAP T E R V II I
HYDRA AND C ( E LEN TERATE S I N G ENERAL

101 . B o ur ne , G C . .
,
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C HAPT ER X III
HI S T ORI CAL Z OOLOGY

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3 20 I
B B L OGRAP H I Y
Ha e ck e l , Erns t 1 8 8 3 T h e E v o l u ti o n o f M a n 2 v o l s ,
. . .

H e a d l e y F W , 1 9 00 ,
P r o b l e m s o f E v o l u ti o n
. Lo nd o n . . . .

J am e s W 1 8 9 0
, T h e P r i nci p l e s o f P s ych o l o gy
.
, 2 vols . . . Ne w
Y ork .

J e nni ngs ,
H S . .
,
1 9 04 . C o nt r i b u ti o ns t o t h e S t u d y of th e Be
h a vi o r of Lo w e r Or ga ni s m s . C a r ne gi e In t s . P ub . Wa sh
i ngt o n .

190 6 Be a . h vi o r o f t h e L o w e r Or ga ni s m s N e w Y o r k . .

1 90 8 — T h e . I nt e r p r e t a ti o n o f t h e B e h a vi o r o f t h e Lo w e r
Or ga ni s m s . Sci e nce N S v o l 2 7 pp 6 9 8 7 1 0 — . . . . .
, , ,

J o r d an D S , . .
,
1 90 5 — T h e Or i gi n o f Sp e ci e s th r o u gh I s o l a ti o n
. ;

Sci e nce N S v ol — 62
,
. .
, . 2 2, pp .
545 5 .

and V L . . K e ll ogg ,
1 90 7 . E v o l u ti o n a nd Ani m a l li f e . Ne w
Y ork .

K e ll ogg ,
V L . .
,
1 90 7 . D a r wi ni s m T o -D a y . Ne w Y ork .

Lo e b , J .
,
1 9 00 C o m p a r a tiv e P h ys i o l o gy
. of th e B r a n i a nd C om
p a r a ti v e P s yc h o l o gy Ne w ork . Y .

1 905 . S t u d i e s i n G e ne r a l P h ys i o l o gy, 2 p t s C hi ca go . .

1 906 . T h e D yna m i cs o f L i vi ng M a tt e r Ne w ork . Y .

1 90 7 . C o nce r ni ng t h e T h e o r y of T r o pi s m s . Jo u r n . E xp .

Z loo .
,
v ol .
4 , pp .
5 1 1 — 1 56 .

M e rr i am C ,
. 1 898 . L fe i Z o ne s a nd C ro p Z o ne s o f th e U nit e d
St a t e s U S D e p t o f Agr i c ll B i o ] S u r v e y,
’
. Bu . 1 0, . . . .

M e tca lf M M 1 9 0 4
,
Or ga ni c E v o l u ti o n
. N e w Yo r k .
,
. . .

M ontgom e ry T H 1 9 0 6 Ana l ys i s o f R a ci a l D e s ce nt i n Ani

,
. .
,
.

m als N e w Yor k
. .

M or gan C L 1 8 9 6 ,
H a b it a nd I ns ti nc t
. Lo nd o n
.
,
. . .

1 908 Ani m a l B e h a vi o r 2 d e d
. N e w Yor k ,
. .

M organ T H 1 9 0 3 ,
E v o l u ti o n a nd Ad a p t a ti o n
. .
,
N e w Yo rk . . .

1 90 7 E xp e r i m e nt a l Z o o l o gy
. N e w Yo rk . .

M ii lle r F 1 8 6 4
,
F ur D a r wi n
.
,
Le i pz i g . . .

Os b o r n H F ,
1 90 8 T h e F o u r I ns e p a r a b l e F a c t o r s o f E v olu
. .
,
.

ti o n Sci e nce N S v o l 2 7 pp 1 4 8—1 5 0

.
,
. .
, .
,
. .

P l a te L 1 9 0 3 U e b e r d i e B e d e u t u ng d e r Da r wi n sch e n Se l e c ’
. .
, ,

t i o ns p r i nz i p .

R o m a ne s , J . G .
,
—
1 89 2 1 89 7 . D a r wi n a nd t
af e r D a r wi n .
3 v ol s .

L o nd o n .

R u th ve n, A G Va r i a ti o ns a nd Ge ne ti c R e l a ti o ns hi ps of
. .
,
1 908 .

te
th e Gar r
S na k e s . B u ll 6 1 U S N at M us
.
,
. . . .

Ste ve ns W ,
. C .
,
1 90 2 . I nt r o d u cti o n t o B o t a ny B o s t o n . .

Th om s on J , . A .
,
1 896 . T h e St u d y o f Ani m a l L i f e N e w Yo rk . .

1 9 08 . H e r e d it y . Ne w Y or k .
 G L OS S A RY

KEY T O P R ON U N C I AT ION

o as i n no t, o n.

6 as i n no te , sou l .

6 as in mo v
e ,
s po o n.

6 as i n no r s o ng .

a bd om e b e lly — f ro m a ba o, I l) th e
’ ’
é da m e n (L th e c o nce a
’
n, a éa om m
'

a .
, ,

l owe r
b e ly l .

’
a b s orp ti on, a as drp s/zon wa ll o w d o wn a nythi ng) t h e pr e ss
(L . a bsor bé r e , s , oc

o f t a k i ng u a s by ch m i ca l o r m o l c u l a r a cti o n
p , e e .

( r e t u rn to g r w ) gr o wth by e xt e r na l a d d i
’
a ccr e t i on a é é / n ( L a a t o
’
, r s zo .
, ,
o ,

ti on o f ne w m a tte r .

ach rom at i n é a m ti n ( G r a
, a with
, z
'

ou t ’
e / r o m a c o l o r ) t h e n o n s t a i ni ng
- .
,
z a, ,

s u b s t a nc e o f t h n cl e s e u u .

a cc e lom a t e sé l m at ,( G r a with o u t '

k
'

o l h o ll o w ) with o u t a t r u e b o d y
. a,
’

a e as , ,

ca vity o r coe l o m .

r o I b rin v e yi ng f r o m th e s u rf a ce to th e
’
a ff e re nt e r e nt ( L E
f , a
fi g t o
) c o n . a , ,

ce nt e r .

a m e b a a m é a ( Gr m o zb s c h a nge ) a ge nu s o f r hi o po d o u s P r o t o o a
’
’

, a . a o , ,
z z .

a m i tos i s a m i ta s zs ( G
, with o u t m i l a thr e a d ) d i re ct nu cl e ar d ivi s i o n
’
'

r. a , e s, ,

With o u t t h e f o m a ti o n o f c h r o m o so m e s a nd a m phi as t e r
r .

af nph i a s t e r a st a r ) th e a ch r o m ti c
’
mfi te ( Gr a mp/ i a r o nd a st
, a as r . t , u er , ,
a

P
fi gur e f o r m e d i n m it ti c ce ll d ivi s i o n co ns i sti ng o f two a ste rs co nne cte d
o ,

b y a s pi nd l e .

anabol i s m a n a b a l z m th wi t tiv m e t ab
g p)
'
(
'

, G r a na é al e a r o n z u c o ns ru c e .
, ,

l
ana ogou s , a na l’ag u s ( Gr . a na , s im il a r to ; log os , pro po r tio n) lik e , i n f u ncti o n,
b ut no t i n t
s ru c u re t .

Tire Ce ntu ry D i cfl ona i am e s S to r m o nth M a nu a l

’
a nd Cy clope d za , th e Re v .
J s

f S bi e ntzfi e Te r m s , W ils on s Ce ll i n De v e lofim e nt I nhe r ita nce ha ve

'

and
’
o a nd E B . .

be e n in va l ua b e l i n pr e pa r i ng th i s g os sary l .

3 23
3 24 GLOSSAR Y
a na ph a s e , a n a
’
f a z ( Gr
'

i n) th e l a t e r pe r i o d o f m it osi s
. a na , b a ck or a ga ,

d u r i ng th e d ive ge nc o f th e d u ght e r ch o m o s o m e s r e a r .

a na tom y t o c t u p) th e st u d y o f th e st r u ct ure o f
'
n t m i ( Gr
‘

, a an t nn o . a a er o, u ,

or
ga ni s m s a s m a d e o t b y d i e cti n u ss o .

Anne l i d a a n l i o ( L nn ll a littl e r i ng ; G r
’
r e se m b l a nce
’
e zd os
‘

) a
'

,
e a . a e u s, .
, ,

p h y l u m o f a ni m a l s h a vi ng b d i e s m a d e p o f m ny s m a ll i ngs o u a r .

a s a il ya d ) t h e j o i nt e d f e e l e r s u po n th e
’
a nte nna n te n a ( L
’

, a nt nn . a e a, r ,

h e a d s f i ns e cts a nd
o st a c a ns cru e .

a nt e nnu l e nt nn ) t h e s m ll e r pa i r o f a nt e nnae i n
’
n t n ul ( L d i m
, a f e . . o a e a , a

th e crust ce a ns a .

a nus d nu s ( L anu s a r i ng) th e o r ifi ce th r o u gh whi ch th e r e f us e o f d ige s

'
, .
, ,

ti n i s v i d e d
o o .

Ap i s i p zs ( L p s a b e ) a g nu s o f b e e s
’
’ '

, ( . a z , e ,
e .

a rch e nt e ron / 2 b e gi nni ng i nt e sti ne ) the

’
k nt , n ( Gr
ar e e nte o n
e ro . ar c 1 , r , ,

pr i m itiv e d i ge s tive cavity .

a rt h rob ra nch i a ( Gr r t/t n a j o i nt or ng / a th e gill

j
t/ o o a ng é i , ar zr r a . a ro , a e te ,

o f a fi s h t i t t fi h
) p a f t h e ,r e
p ra o s
y y sr e m o f t hoe c y s s r ra .

A rth ropod a , ( G r t/ o n a j i nt ; p o f o t ) a
phyl u m f . ar zr , o u s, o , o

a ni m a l s h a vi ng b o d i e s co m o s e d o f s e gm nt s s o m e o r a ll o f whi ch b e a r e
p ,

j o i nt e d appe nd a ge s .

As ca r i s ( G r a sé r zs a w o r m i n t h e i nt e s ti ne s ) a ge nus o f r o u nd
’
‘ ’

, s é a s a r z . a , ,

w o rm s .

a s e xua l a s e é s ? a l ( G r with o t ; a nd se x u l) appli e d t o m o d e s o f

'
, z . a, u a ,

r e pr o d u cti o n i n whi c h t h e s e xe s a e no t co nce n d r r e .

a s s i m i l a t i on m lo to m ak e li k e ) th e c o nve rs i o n o f
'
i m i la / n ( L
’ ’

a s s to . a ss z z , ,

d i ge ste c o o d i nt o livi ng p t o pl a sm
1
ro .

a s te r G a s t a ) t h e s t a r -s h p e d s t r u ct ur e s u r r o u nd i ng th e ce n
’
s tér t
, a
( r e . as r , r , a

tr o so m e .

au d i tory , c
’
i d z to
’ “
rz
'

( L . a u d i tor , a h e ar e r ) , p e r a t ini ng to th e se nse of h e a i ng

r .

be h av i or, oé /t au y or t; h o l d h a ve ) th e t ot al of a ll
'
( oe , abo u Iza ooa n, , , su m

th e var i o us m o ve m e nts o f a n a ni m l a .

“
il
b at e r a l o l, t'
e r l ( L ozs t w i ce ;
z a f a s i d e ) h a vi ng th e s i d e sym
a .
'

,
o ,
s

m e t i cal r .

b i oge ne s i s o j n e s zs ( G r oi os li fe ge ne s zs o r g n) th e ge ne s i s o f livi ng
'
' ' '

, zo e .
, , i i ,

b e i ngs f r o m livi ng b e i ngs .

“
b i o ogy o ol oj z ( Gr bi os li f e
l '
l go d i s co u r s e ) th e s ci e nc e o f li f e a nd liv
' '

, z . , o s, ,

in g thi ngs .

“
bi ra m ous o r d nt us ( L oi tw o r a m u s a b ra nch ) d ivi d i ng i nt o t wo
’
, z . , ,

b r a nch e s .

b la s tocml blas to sé l ( Gr bla sto a b u d

’
k o llos a h o ll o w ) th e cavit y o f th e
'

, . s, , ,

b last u la .
3 26 GLOSSAR Y
ch oa nocyt e
’
ko a ( G nao u e f u nne l ; é u tos , vit y) ll ar e d
" ’

, no s zt r . e , a a ca , a co

ce ll of th e s po nge .

c r om a t n,
h k ’
i l ) th e d e e pl y st a i ni ng s u b sta nce
( Gr nr om a
’

r o m a t zn . e , co o r ,

o f th e nu cl e a r n tw o r k nd f th e ch r m o s o m e s co ns i sti ng o f nu cl e i n
e a o o , .

ch rom os om e s m (G / on co l o r ; b o d y) t h e d e e pl y
'

,
s n o r. e zr r a , o ta , ,

st a i ni ng b o d i e s i nt o wh i ch th e c h r o m a ti c nucl e a r ne tw o k r e s o lve s its e l f r

d u i ng m it o si s
r .

Ch rom otropi s m é o m bt c o l o r ; tr p a t u ni ng)

’
pi m ( Gr o/z m
’

th
’

,
, r ro z . ro a, o e , r , e

r e a cti o n o f a n a ni m l t o c l a ~ o o r.

il
c i um ,s zl’
z um (L
’

i li m a n ye li d with th e h i s gr o wi ng o n it ) a m i n t e
’

. c u , e a r , u

h a i li k e pro ce ss o f a ce ll
r .

cl a s s Has ( L l s z a n a sse m b l y o f pe o pl ) a n m b e r o f a ni m a l s e g a d e d
'

, . e as s, e , u r r
.

a s a co ll e ctiv e u nit b e o f t h r e se n o f c e t a i n co m m o n ch a ra ct
ca s e e ce e r
us p r

t h m vi i n o f th f e tili e d e gg
.

cl e a va ge é ZE vdj
'
, ,
,
s o e r z .

cl it e l l u m é lz t l pa ck sa d d l e ) a gl a nd u l a r r i ng a o nd
’
m (L
’

, , li t /l ae u . e e ee , ,
r u

th e b o d y o f th e e ar thw o m r .

cni d ob l a s t ni a o alast ( G r é n a ne ttl e ol stos a b u d ) a ce ll i n whi ch a

' ’
’
'
’
, . za e ,
a , ,

ne m at o cys t i s d e v e l o pe d .

cni d ocil ni l ( G r l ni a a n ttl e L ci li u m a n e ye li d ) a ti ff h i li k e

' ’ ’
' '

ao sz . e e, e s a r
, .
, ,

r o ce s s proj e cti ng f o m a c ni d b l a s t r
p o .

coa gu l at e urd l e ch a nge f r o m a fl i d i nt o a

’
t o g u lat ( L a u la e
)
'

u
, g a c . co r , ,

c u r d li k e o r thi c k ne d m a ss e .

C (B le nte r a se l n t , ( G é l s h
e o ll w ;
'
e n ete n i
ra n t e s ti n ) a h
p y rl m .f , oz o
'

,
o ro ,
e ,
u o

a ni m a l s w ith a s i ngl e b o d y c a vit y whi c h O pe ns b y a s i ngl e ifi ce th or ,

e

m o uth .

C(B lom sE lonz ( G r k t l s h o ll o w ) th e t r u e b o d y cavity l yi ng b tw e e n t h e

’
’

o a e
, .
, , ,

d i ge stive t a ct a nd t h e b d y w ll a nd li ne d with e pith e li m

r o a , u .

cce lom a t e se lo n i t ( G , é l h ll w ) h avi ng a c oe l m

’
f c r. oz os ,
’

o o ,
o .

comm e nsa l i s m n m e n om t o g th e tab l e ) th livi

’
l n (L ; m n
'

, o sa ze r . c , e r e sa , ,
e

t o ge th e r o f tw o spe ci e s f a ni m al s o r pl a nts b ut n ith e r a t th e e xpe nse o , e

th e o th e r .

conj uga t i on t n ] g a / on ( L e nj g a t n to u nit e ) a t m po r a ry o r

’
, a 0 s z . o u ur , , e

ne nt u ni o n o f tw o c ll s f r r e p d u cti o n e o ro .

cont ract i l i ty é n t t us a d a wi ng t o ge th e r ) th e
’
l t zl ti ( L
' '

, nto ra z . co rac ,
r ,

e r ty o r f o r c e b y whi ch b d i s h i nk o r c o nt a ct o e s r r .

cop u l at i on k op a M t on ( L u nit e ) th e a ctive t a ns m i s i n

'
, pu l s/ . co are, ,
r s o

s pe r m f r o m th e m a l e t th e f e m a l e o .

cort i ca l e o a k l ( L r te x b a k r i nd ) b e l o ngi ng to th e e xt e rna l co v e r i ng

’
,
r a . co , r , ,
.

cu t i cl e k u t i k l ( L e u tze u la d i m o f u t s th e s k i n) th e o u t e r m o st c o v e r i ng
’
' '

, . , . c t , ,

o f t h e b o d y o f c e rt a i n a ni m a l s .

cyclos i s s é lo s i s ( G r k u é l s a c i r cl e ) a ppli e d t o cu r re nt s withi n th e b o d i e s

’
‘

, z . o , ,

o f P r t o zoa o
 GLOSSAR Y 3 27

cys t, s i st ( Gr l a d d e r ) a t e rm a ppli e d to ce rt a i n Pr o t o zo a wh e n
lzusti s , . a b ,

th e y su r ro u nd th e m s e lve s with a w a ll a nd p ss i nto a r e sti ng st g a a e .

cyt oph ar ynx i to f i nge s ( G Au t s a ve ss e l ; pn ngx g ll t ) t be’

‘

s ' ar r. o ar u u e
, , , , a u

l e a d i ng f r o m th e b o tt m o f t h e o r l gr v e i nto t h e b o d y o f a Pa am ci m o a oo r e u .

cyt op l a s m s i to pl m ( G r k t s a ve ss e l pl s n a nythi ng f r m e d
’
,
az
) th e . u o , ,
a za , o ,

s u b s t a nc e o f t h e c e ll b o d y a s o pp se d t o th a t o f th e nu cl e u s o .

d i a s ta s e ,
’ ’
ai a s tas
'

( Gr . a
’
i a s ta s i s, a se
pa r a ti o n) , a s ub s a nce t whi ch h a s th e
pr o pe r ty o f co n e r v ti ng st a rc h i nt g
o s u a r.

d ia s t ol e , a
’
i as
’
to le ( Gr d i a stole , . se
p a ra ti n) th pe i d f ti m e d u i ng whi h
o ,
e r o o r c

a r h yth m i ca ll y pu l sati ng v e sse l 15 r l ax d

e d il t e d
e or a .

j
'
(L ige sti o th e d i ss lvi ng o f f o d ) th e p o c s s o f pr e
’
d i ge s ti on,
’
ai es ty o n . a ,
o o , r e

p ar i ng f o o d p ti o n fo r a bs o r .

d i e ci ou s i twi e a h us e ) h mmg th e s e x e s d i s ti nct

’
i e s/ s ( G r é
’ ’
, a ié zu . a s, c o os , o , ,

a p li e d t o S p c i s th a t co ns i s t o f m l e n d f m l e i nd ivi d u l s
e e a a e a a
p .

d i pl ob l a s t i c ip l bl ti n( G r
“
' b l ; bl t h avi ng tw o
’
ipl d a b ud )
’
, a o as . a oos , ou e a s os , ,

g e r m i na l l a ye r s .

d i s s i m il a ti on ’
/ on ( L i s i m i la e m a k e u nli k e ) th e p ro ce s e s
’ ’
i i n i l t ,
a s 7 e s i . a s r , , s

by whi ch p t pl a s m i s b r o k e n d w n i nt o s i m pl r p d u cts
ro o o e ro .

d i s ta l d i t l ( L i t I s t nd a pa rt ) s it u a t d w a y f m th e pl c e f a tt a ch
’ ’
,
s a . a s o, a , e a ro a o

m e nt .

d or s a l d o s l ( L d o s u m th e b a ck ) o f o r p r t a i ni ng t th e b c k
’ ’

,
r a . r , , e o a .

e cd ys s , e i na i ’
sis ( G r . e k a u s i s,
’
e m e rg ng) , i s h e d d ng i or mo lti ng th e e x e r na t l
co ve i ng r .

e ct od e r m , e té to o e r /n ’ '
’

( Gr . e k tos, o u s t id e ae r
’
ma, sk i n) , th e ou e r t ge r m l ay e r .

t
e c osa r c, e nto s ar é ( G r
’ ' '

. e é tos, o u ts i d ; e sa r x , fl e sh ) , th e ou e r t l a ye r of ce r

t i n P r ot o
a z o a.

e ff e re n t , e
f
’
e re nt (L . e
f for e x , o ut
f e r o, I ca rr y
) , co n e v yi ng tw a r d ou .

e ge s ti on é j , e s e/
zon
’
( L ege sti o v o i d ) th e a ct o f v o i d i ng
.
, , th e re f us e f d i ge sti o n o .

l t
e e ct r o r opi s m , e le é tr o tr o p i zm ( G r e le ctr o n a m b e r
’
' ’

.
, tr o p a t r ni ng) r
e , u , e

ac ti o n to th e e l e ct i c r cu rr e n t .

“
e m b ryo, e m br i ’
0
( C r . e n, i n or u o, b u d ) , th e e ar l y s ta ge of an a n i m a l wh e n

it i s withi n th e e gg m e m b ra ne .

e nd opod i t e e n a p o a t ( Gr e n n withi n ; pou s a f o t ) th e nne r o f th

’ ' ’ ’
'

o z ao o ,
r e
, .
, ,

t w o m a i n d ivi s i o ns o f a t yp i ca l cru st a ce a n a ppe nd a g e .

e nd osa r c e n r e ndo n withi n fl e sh ) e i nne r pr o t o pl a s m

’ ’
s ar é
( G h
'

t
’

, ao sa r x .
, , ,

o f ce rt a i n P t a r o o zo .

e nte r on e n te r on ( G r e nt
,
'
n i nt e sti n ) t h d i ge stive t a ct th a t i s pr i m i
. ero ,
e , e r

t i v e ly d e iv d f r o m th e e nt d e m r e o r .

e nt od e r m e n t m ( G r nd o n with i n
’
s k i n) th i nne ge r m l aye r
’ ’
rm
”

, o ae r . e , ae a, ,
e r .

e nzym e e n z i m ( G r e n i n
,
'
zu m e l ve n) n u n rga ni e d f me nt .
, ,
e a , a o z e r .
3 28 GLOSSAR Y

p i ge ne s i s th e o ry ve l o pm e nt
’
e , e
p ij e n e si s ( Gr . e i
p, u
po n ; ge ne s i s ) , a of de
( se e p .

p i pod i t e , e p ip pou s f o o t ) l
’
( Gr ep i
’
de ve pe d
'

o a it u po n
e .
, ; , ,
a p r o ce s s o u
po n
th e b asa l j o i nt o f som e of th e a ppe nd a ge s o f ce r a t i n st e a ns cr u ac .

e p i th e l i u m , ep i n
t e li ’
u m ( G r ep i .
, po n ; tn
u a llo, I ro , a g w ) t h e l ye r of ce ll s
i
f o r m ng th e s u r fa c e o f a ll th e i t l
n e r na m e m b ra ne s o f t h e b o d y .

e vo l ti
u on, e v o lu s/
’
zo n
( L . e , o ut ; v oluo, r o ll ) , a th e o ry o f de v e l o pm e nt
p .

e x cr e ti on, e ns é ’
r e s/
zo n (L . ex, o ut ; e r e t u s, se
pa r a t e d) ,
th e e li m i na ti o
u se l e ss o r h a rm f u l s u b s a nce s t f r o m th e b o d y .

e x opod i t e , e é so p
’
o a
'

’
ft ( Gr . e x o, t id
ou s e ; p o u s, fo o t) , th e ou e r t of th e
ma n d i ivi si o ns of a t ypi l ca cr ust a c e an a
pp nd g
a e e .

e xos k e l e t on, e lz so s k e l ’
e t on
( G r . e x o, ou s t id e ; sk e le to n
) , a ny s r u c ur e t t
d u ce d by th e h a r d e ni ng o f th e i nte gu m e nt .

fa ce s , f e ’se s ( L .
f a’x , d r e gs ) , u nd ige te d parti l s
s c e ca s t o ut by an a n m a i l .

f a m il y, f a m ’i li ( L f a m i li a .
, a h h o l d ) th e
o use ,
na m e of a
gr o u p abo ve a

ge nus a nd be l w o an o rd e r .

f a u na , f i na
c
’
(L . Fa u u u s , o ne of th e go d s o f th e fi ld )
e s , a ll th e an im l a s
pe
c u li a r to a c o u n ry, a r e a , o r t pe r i
od .

f e r m e nt , f er m e nt ( L f ’
. e r m e nt u m , l e a ve n) , a s u b s a nc e t whi h t c r a ns f o r m s a n

o rga n c s u b s a nc e i t i nt o ne w c o m po u nd s .

f e rt il i za i on, t f
e
r
’
ti l i za s
’
no n ( L f e r ti l i s .
, fru fu it l ) , th e un on o i f a S
p e rma

to z o o n W i th an e
gg .

fla ge l l u m , fl j a ’ f
el u n (L .
fl a
g e llu m , a i
w h p) , th e w h i pli k e ap
p nd age
e of

m a ny P r o to zo a .

f ol i a ce ou s , f ni u s ( L f oli u m l e a f ) r e s e m b li ng
li d ’
l f
'

o s .
,
a e a .

f u ncti on f u ng é sn
’
on
(
, L f u nct i o
p e r f o r m a nc e
,
) t h e .
, , m o de o f ac ti on
p r o pe r

to a n o r ga n o r s ru c u r e t t .

ga me t e , ga m e t
’
( Gr .
g m e te ,
a a wi f ) e , a re p ro d u c tive ce ll whi ch un it e s with
a no th e r re
p rod uc tiv e ce ll to fo r m a z ygo e t .

ga ng l ion g , a ng gl i ’
on
( G r g a nggli on,
. a t u m o r u nd e r th e sk i n n ar t e nd o n) e a ,

a m ass o f ne r vo u s ti ss ue co n a t i ni ng ne ve r ce lls a nd g ivi ng r i t o ne rv e se

fi b e rs .

ga s t ri cg
, ( G r
gas
’
a s te r , s o m a c
)
tr i l'
, o f o r p e r a n ng to th e s . t h t i i t o m ch a .

li ’
ga s t ro th , ga s tr o li tlz ( Gr ga ste r , s o m a c li tlt os, s o ne ) , a
’

. t h t s t o ny co ncr e

oti
n i n t h e s o m ac o f c e r a n c r u s a c e a ns t h t i t .

ga s tru a , l g as
’
tr o la
’ ’

(d i m . of G r ga ste r ,
. t
s om ac h) , an e m b ryo c o ns s i ti ng Of

t w o ge r m l a ye r s i n l s i ng a
c o ce n tr l a c av it y .

ge n i ta l j e n i t l ( L
,
’
a .
ge ni ta li s, se r vi ng t o b e ge t) , p e r a t i ni ng to th e or
g a ns o f
e ne r a ti o n
g .
33 0 GLOSSAR Y

i nt ra ce ll u l a r, i n tr a se l
’
z? la r ( L i ntra, . withi n) , e x s i ti ng o r d o ne i nsi d e of a

ce ll .

t
i n us s u s ce pt i on, i n t u '
s u se p
’
s/
zon
(L . i ntu s, withi n ; s u s ce
pt u s, ta k e n up ) ,

th e add iti o n of ne w pa r ti cl e s a m o ng t h e p e e xi ti ng r s
pa r ti cl e s o f pr o t o

p l as m .

i rri ta b i l i ty ,
’
i r i ta oi l i ti '
( L i r r i ta r e ,
. e xc it e ) , th e pr o pe r y t of re s
po nd ng to i
sti m u li .

k a ryop l a s m , n ’
ar i o pla zm ( G r é a r u on, nu c l e us ; pla s m a , thi ng fo rm e d) ,
'

. a

th e s u b s a nce t of th e nu c l e us .

k ar yos om e , é a r i (7 s ome ’
( Gr . k a r u on, nu c l e us ; s om a , b o d y) , nuc l e li whi h
o c

t i n with nu cl e ar d ye s
s a .

k a ta b ol i s m n ’
a t o o li m
( Gr é t l le a th r o wi ng d o w n) th e pr o ce ss by
'

, a z . a a o , ,

w hi c h pr o t o pl as m b re a k s d o w n i nt o S i m pl e r pr o d u ct s .

l a rva , lar v a
'
’ L la r va ,
g o s o r m as k ) , th e yo ung o f a ny
( . a h t an i m a l which
i vl
d u r ng i ts d e e o pm e n i s u n k e i ts pa r e n t li t .

l ’
la te ra , la t e r a l ( L la tu s, S d e ) , o f o r p e r a n ng to th e S d e
. i t i i i .

m ric e a thw wo r m s
’
Lu b u s , l u m or i é us ( L l u m br i cu s ,
. r o rm ) , a ge nu s of .

macroga m e t e , m a k '
ga m e t
’
( Gr m a k r os, l o ng ; m e te , w f e i ) la
‘

ro .
g a , a r
ge te

pr o d u c tive ce ll , th e e gg .

m a crom e r e , m a é l o ng ; m r s a pa rt ) th l a ge r e ll s
’
r o m Er
'

( G r m a é r os, . e o , , e r c

i n t h e e r l y e m b ryo ni c s t ge s o f ce t i n a ni m l s
a a r a a .

m acr onu cl e u s m nr o u nk le u ( G r m t r os l ng n le ) 3 l arge nu cl e us

’
a s . a ,
o uc us , .

i n c rt a i n P t o o a
e ro z .

m at e r i pe n) t h e r i pe ni ng o f th e e gg by
’
u r a i
t on m a t 2 r
, t sl o n ( L m tu 7 e t . a rar , ,

th e f o r m a ti n o f po l a b d i s o r o e .

ma x ill a m é,s i l ’
i ( L m a x i ll
a a jaw )e
’

a n a ppe nd a ge ne ar th e m o uth o f
. a, ,

a r th ro po d s .

max il l i pe d m te s i l i pe ( L m x i ll a ja w pe a e s f o t) a f o o tli k e a ppe nd

’ ’ ’

,
a a . a a, , o ,

a ge n a r t h e m o th o f ce t i n
e th o p d s u r a ar r o .

m e d us a m e , ( L M u s a m yth l g i ca l w o m a n. wh se heai r w
’
a s t u r ne d a, o o o a

i nt o s nak e s ) a j e ll yfi sh o r e pr d ctive o o i d o f ce rt a i n coe l e nte r at s

, , r o u z e .

m e rob l a s t i c m e r o bl s ti é ( G m e r s a pa t ; ola t
’
a ge r m ) a ppli e d t o
‘

, a r . o , r s os , ,

e ggs o nl y t f whi h e c t i nt ll d i n l v a e
p a r o p c e s u g c e a g c ar u u o r .

m e s od e rm m e s a ér m ( G r m e sos m i d d l e ; d m
' ’
Sk i n
) t h e m i d d l e g rm
’

o er a, e
, . , ,

l aye r o f t i pl o b l sti c a ni m a l s
r a .

m e s ogl e a m e s oglé ci ( G r m e os m i dd l e ; gloi a gl e ) th e ge l ati n u s s b

’’ ‘

s , u o u
, .
, ,

s t a nc e b e tw e e n t h e e ct o d e m a nd nt o d m o f coe l e nt e a t e r e e r r s.

m e ta b ol i s m m e ta a li m ( G r m e t / le ch a nge ) t h e pr o ce sse s co nne cte d

’ ‘

, o z . a fo , ,

with th e m nufa ct ur e a nd d e str u cti o n o f prot o pl s m

a a .
 GLOSSAR Y 33 I

m e ta m e re , m e t a m e r ( G r m e ta , ’
p t) ne o f a l o ngit u d i na l f te m e r os ,
'

. a r a ar ,
o

s e r i e s o f par t s whi ch a e s e i a ll y h m o l o go u s with o ne no th r r r o a e .

m e t a m orp h os i s m t m or f o s i s ( G r m e t ch a nge m o pn sha e

p ) a
’ '

, e a e . a, r , ,

m k e d ch a nge i n f o m o r f u ncti o n
ar r .

t
m e a ph a s e m t’a d
f z ( G r m e ta
, a f t e r ) th e m i d d l
e s t a ge o f m it o s i s d i ng
.
, , e ur

whi ch o ccu rs th e s plitti ng o f th e h o m o o m e s i n th e q at r i a l pl at e c r s e u o .

m e ta pl a s m m t a pl zm ( G r m e t a f t e r ; pl m
, e
’
a thi ng f o m e d
a
) pp a li e d . a, as a, r ,

t o li f e l e s s i ncl u s i o ns i n p o t pl as m r o .

M e ta zoa n t a z o a ( Gr m e ta a f t e r ; z o n a n a ni m a l ) m a ny-ce ll e d a ni
’ '

, i e .
, o , ,

m a ls .

m i cr oga m e t e , m i k r fi ga m ( G m i é r os , ll ; wi f e ) ll
’
ét r . sm a g a m e te , a , a sm a

re p r o d uc tive ce ll , th e s
pe r m a o zo o n t .

m i crome re , m ’
i kr o
’

m ér ( Gr . m i k r os, ll ; m e sm a r os , a
p ar t) , th e s ma ll e r ce ll s
i n th e e m b ryo ni c t
s a ge s o f ce r t a i n a ni m a l s .

“
m i cr onucl e u s , m i k m u nk le
’
us ( Gr . m i cr os , sma ll nu cle u s ) , th e sm a ll e r o f
tw o nu c le i in t i n P ot o a
ce r a r zo .

m i tos i s , a th re a d i i re ct l e ar d ivi si o n
(
’
G r m i to
)
'

mi t si s
a . nd s, , nu c .

nos
“
m once ci ou s , m o n ’
E slzu s ( G r m onos, . o ne oi , a h o u s e ) h avi ng , bo th ma le
a nd fe m a le se xu a l o r ga ns .

m orph ol ogy, m or f ’
ol o
j i ( G m orpne , f o rm ; log os , d e scr p i ti o n) th e i nce
'

r .
, sc e

of for m a nd s r uc ur e t t .

m ou l t , m olt ( L m u ta r e , . h ng ) c a e , th e p r o ce ss of c as ti ng o ff t e
g u m e n t a ry ,

ti cu l ar o r
cu , e xo s k e l e t a l st u ct r ur e s .

u t i
m a t on m ? td ’
sl ,
z zo n
( L m u ta r e ,
. c h a nge ) ,
a th e o ry o f th e or i gi n o f s pe c e s i
( se e
p .

ne m at ocys t ne m t s i st ( Gr ne m
,
th r e a d ; l u ti s a b ag) a st r uct ur e co n
'
a
‘

o . a ,
'
s , ,

ta i ni ng a th r e a d ch a ra ct e r i s ti c o f coe le nt e at e s ,
r .

ne ph r i d i u m nef r i i m ( G r nep/r s a k i d ne y) o ne o f t h e e xcr e to ry

, a
"
u . z o , ,

o r ga ns o f th e e a thw o r m r .

ne ur on n u
,
’r
on ( G r n u on a n e rv e ) a n e rv e. ce ll wit hea ll i trs pr o l o n g ti,
o n s
,
a .

nu c e o us nu é IE lu s ( L nu cle u s a s m a ll nu t) a d e e pl y st a i ni ng b o d y o ft e n
l l '
'

, , o . , ,

pr e s e nt wit h i n a nu c l e u s .

nu c e us n k u s ( L n cle u s a s m a ll nu t) the d yna m i c ce nt e r f a c e ll

l ’ le ’ o
’

i ,
z . u , ,
.

t i
nu r m e nt nu t i m e nt ( L n
’
,
u tr i r e nr o u i s h ) th a t whi h p ro m o t e s .
t h e
,
r ,
c

g orw t h o r r e p a i s t h e w a s t e o f a ni m a l b o d i e s r

t ,
’
nutr i i on nu t i s on ( L n tr i
l r r n o u i s h
z ) th e p o c e s e b y whi
.
c h o g
u
a n e ,
r ,
r s s r

i s m s t ak e i n f o od a nd a d d it t th e i r livi ng ti ss u e s o .

oce ll u s ,
”
’
o se l u s (L . oce ll u s , a littl e ) i m pl e e ye O f a n i ns e c t
e ye , a s .

(E s op h a gus , e so
f
’
a
g us
( Gr . o s i opl gos th e gull e t ) th e ca na l th ro ugh whi ch
za , ,

fo o d a nd i
d r nk pa ss e s to th e s t o m a ch .
33 2 GLOSSAR Y
om m at i d i um , om a tzd i
’
’
um
( Gr . di m . of om m a , e ye ) , par t of th e c o m po u nd

ye o f a n a r r o po d
e th .

’
ont oge ny, on toj e ni ( G r . o n, i
b e ng ; ge nna o, I pro d u ce ) , t h e d e v l o pm e nt o f
e

an i nd ivi d a l u o r ga n s m i .

oocyt e ,
’
o o si t
'

( G r . oon, a n e gg) , a s a ge t i n the m a t u ati o n f

r o an e gg .

r i gi n i gi n
( )
’
ne s i s , b aj e n e si s G oon, an gg ; ge ne s is th e a nd
'

ooge r . e , o , or

de ve l o
pm en t of th e e gg .

oogoni u m , o o go ni u m
' ‘

’
( Gr . o on, an e gg ; go nas, ge ne ra ti n) o , an e gg j stu

be fore ma t u rati o n t ak e s pl a c e .

or ga n, or ’g a n (L . o rga nu m , an i nst ru m e nt ) , a
pa r t of a o r ga n s m i h avi ng a

s
pe c ifi c f u nc ti o n .

ort h oge ne s i s or t/zoj e n’e s i s ( G nos t i ght ne s i s ) , th f

’

r or t s ra a
, .
, ge e o ry o e vo

luti o n ( pse e .

os cu um ,l ’
os k u l u /n (L . oscu lu m , a littl e m o u th ) , an o
p neng i th ro u gh whi ch
w at e r i p s e x e ll e d f ro m a s po ng e .

os ti u m , os t i u m
’
(L . os ti u m , a n o pe i
n ng ) , an o
pe n ng i n i va i o u s r o r ga ns o f th e
bo d y .

ova ry,
’
5 va r i (L . ov u m , a n e gg ) , t h e o r ga n o f th e fe ma le in whi ch e ggs
de ve l o
p .

ov i d u ct ,
’
o v z cl u é t
'

( L . ov u m, an e gg ;
’
a uct us , le d ) , t h e d u c t of th e o va r y .

ovu m , o v u m
"
(L . ov u m , a n e gg ) , an e gg .

ox i d at i on ok i a nn , s a
’ ’
s o
( G r . ox u s , s h a r ) , th e
p co m b i nati on o f a s ub s a nce t
with o xyge n .

pa l e ontol ogy, pa i nt i
’
( G r pa la i os
’
le on tol o j i b e ngs ; logos, di s
’ '

.
, a nc e o nta ,

co u rs e ) , th e sc e i nce of f o ss il o r ga n s m s i .

p a ra s i te , pa r
’
a si t
( G r pa r a s i tos . th e s e xpe n e ) a n , o ne who e a s t at a no r
’
s ,

a ni m a l th t live s i n n o r a t th e e xp ns e o f a n th e r a ni m a l
a , o , e o .

p ar i e t a l p a ,ri t l ( L p a i s
’
w ea ll ) p
a e t a i n i n g t t h e . w a ll r e , , r o s .

t h n n i n j i ( G / ’
vi i I '
'

a nn
'

p a r e og e e s s p t e n n e s
, s p t rn a g n g e oa o e r . ar ze os , r ,

p r o d c e ) u e
p ro d uc ti o n ,b y mr e a n s o f u n f e tili e d e g g s r z .

p e ll i c l e p e
, l i é l ( p
L e
’
l li u l a s m a ll s k i n )
. a thi n o t e r c o v
c e i ng o f s kai n o r, , u r

c uti cl e .

p e r i ca r d i u m pe i é ar a i u m ( Gr p i r u nd b o u t k a a i a t h e h e a rt ) a
’ '
’ ’
, r . er ,
o a r , ,

m e m b r a no u s s c s u nd i ng th e h e a t urro ar .

p e ri s t a l t i c p d wi ng t ge th e r a ll r o u nd )
'
i t l t i é ( Gr p i t lti é
,
er s a . er s a f os , ra o ,

a p li e d t o th e w a v f c nt a cti o n nni ng d w n th e a li m e nt a r y ca na l
p e s o o r ru o .

pe ri s t om e p r i t m ( Gr pe i r o u nd a b o t th ) th e pa t
’
a m
‘

, e s o t nz . r ,
u s a a, ou ,
r

whi ch s u nd s th e m r ro u th o r o r l o p ni ng ou a e .

p e r i t o n e u m p i t n,o um ( G rer
’
p i t n i oo n
"

wh t i s st r t h e d
"
r o u nd o r . er o a , a e c

o ve r ) th ,m e m b a n li ni ng th e b o d y cavity a nd co ve i ng m o st o f the
e r e r

o r ga ns i n th e ab d o m e n .
334 GLOSSAR Y

p rox i m a l p ol ,
r z si
’
mal ( L .
p r ox i m a s, ne a r e s t) , s itu ate d ne ar th e pl a ce o f
a tt h m nt
ac e .

“
p se u d op od i u m , su do p i
o a
’ ’
um ( G r pse u a e s
.
’
, fa l se
po u s , f o o t ) , a t e m po r a ry

pr o tr us i o n t pl a s m i n ce r ta i n ni m a l s
of th e p r o o a .

p p p p (
u a , ? ’
L
i
p p a d ll
a
) a s t a g e i n. t h e li f e
u
yc l o f e rt a i n i ns e ts
a, o , c e c c .

py l o r i c p lo
’
,i é ( G t
p u lo u
"

ar
g a t p ) o f o r pe t a i ni ng to th
-k e e e r r . ifi ce r s, e , r e or

b e tw e e n t h e s t o m a c h a nd th e i nt e sti ne .

py r e n o i d p i re n i , ( G r p u en t h e s
’
t o ne o ao f f r u it ;
’
ci s f r m ) a s m ll
. r , ao
’
, o , a

co l o l e ss m s o f pr t e i d
r as o .

i
r e a ct on, r e a k s/zo n, th e r e s po ns e t a s ti m u l us
’
o .

r e ca p i t u l at i on e a pi t nla / n ( L , a ga i n
r
pi tu lu m a h d ) a
a
’
s zo . re, ca , e a ,

th e o ry th at h l d s th a t th i nd ivi d l i n i ts d ve l pm e nt pa ss e s th gh
o e ua e o rou

th e a nce s t l st g s f t h e r a ce ( s e e p ra a e o .

r e d u ct i on e a ué / n ( L r l d ) th e h a lvi ng o f th e nu m b r
’
b a ck ;
’ ’
,
r t s zo . e , a u c us, e , e

o f c h o m o s m e s i n th e g
r m n l e i d u i ng m t
o ti n e r uc r a u ra o .

r e fle x éfle és ( L
, r
’
e b k fl x u b nd ) b e nt b a k ( s e e p
. r ,
ac e s, e , c .

r e ge ne ra t i on a g i n; o I b e g e t ) t h e r e ne w a l
’
e f n e a / n ( L , r
g n e r f s zo . re , a e er , ,
o

a
po r ti n o f l o t o r e m ve d ti u e
o s r o ss .

re s p i ra t i on b a k ; p i r o I b r e a th ) th e a b s o rpti o n f
'
e pi a a n (L r,
r s r s a . e, c s , ,
o

o x yge n a nd e xcr ti o n o f c a b n d i o x i d e r o .

r os t ru m r os tr u m ( L r ostr u m a b e ak ) a b e a k li k e st u ct u r e
’
, .
, ,
r .

sa l tat i on s l l t / n ( L s lt ti d a n e ) a n a b r pt t ra ns iti o n o r h a nge

,
a e
'
s io . a a o, c , u c .

s a p rop yt ch i p jr o i t’
i é (
, G r s p
sa s p u t i d ;
'

py i /0 a p l a nt ) livi ng n d e . a ro ,
r t s, ,
o

c a yi ng s b s t a nc e s u .

s ca ph ognat h i t e s é f og na t/ t ( G r s np/ w a b a t ; g n tli os a j a w ) a plat e

’ a a o a
, zz .
, , ,

whi ch b l e s t h e w t e u t o f th b r n hi a l ch m b e r f th e cr a yfi h
a a r o e a c a o s .

s chi z ogony se i l v o d u cti o n b y s plitti ng

’
i ( G l i I )
'

g n , o c
zo e a roe p r r . sc z z ,
e ,
.

s e cre t i on se ar e n s e pa r a t e ) t h e pr o ce s s o f s e pa ti ng s b
'
, on ( L s et s s . e cr u , ,
ra u

s t a nce s b y gl a nd l a ctivit y u ar .

s e ne s ce nce s ne e n ( L
’
n x o ld ) t h e co nd iti o n o f g r wi ng Old
'

,
e s s . se e , ,
o .

ns u s s e ns ) ca pa b l e f e ce ivi ng o r t r a ns m itti ng i m
’
s e ns ory s n o i ( L
”

, e s r . se ,
e ,
o r

p r e ss i n s f o m with o u t
o r .

se pt u m s p t u m ( L eptu m a pa rtiti n) a w a ll s e pa r a ti ng t w ca viti e s

’
, e . s o ,
o .
,

se t a, s e ’
ta ( L s t
'

thi n sti ff h a i r ) br i stl e s o r s ti ff h i s

. e a , a , , ,
a r .

se x au l s e é
,
’
sfi l ( L se x u s s e x) o f o r p e t i ni ng t o s e x ; d o ne b y m e a ns o f
a .
, ,
r a

th e tw o s e xe s m a l e a nd f e m l e , a .

s i nus s i n s ( L s i nu s h o ll o w ) c vity o h o ll o w i n ti ss u e
'
, u .
, , a a r .

s om a t i c s m a t i é ( G
,
o som b d y) o f th e b o d y ; a ppli e d t o ce ll s th at d o
’
r . a , o ,

no t t k e pa r t i n r pr d
a ti o n e o uc .

s om i t e o m t ( Gr s m '
b o d y) a s e gm e nt o f a n a ti c l t e d bo d y
‘

, s z . o a, r u a .
,
 GLOSSAR Y 335

s pe rm at id , s
p er
’

m a ti a
"
( Gr spe r m a .
, se e d), a ce ll whi ch is v t
co n e r e d i nto a

spe r m a o z o o n. t
s pe r m a tocyt e , s p
’
er m a to s i t ( G r . spe r m a , se e d), a s a ge t i n th e m a t urati o n o f
a s pe r m a o z o o n t .

s pe rm a oge t ne s is , s
p
'

er
'
m a toj e n e s i s
‘
’
( Gr . s e r m a , se e
p d ; ge nna o, I pr o d u ce ) ,
th e o r i gi n a nd de ve l o pm e nt o f s p e r m a o zo a . t
“
s pe r m a t ogoni u m , sp
’
er m a to "
g o ni u m ( Gr spe r m a , s e e d ; ganos, ge ne r a . ti o n) ,

a ge r m ce
ma us le ll j t b e f o r e m a u ra o n ak e s p a ce t ti t l .

"
s pe r m a t ozoon, spér m a to zo on
'
'

( G r .
p
s er ma, se e d ; zoon, a n imal) , a ma t ur e
ma le ge r m ce ll .

i
sp er m ’ e
e, p r m
s i ( Gr .
p
s e i r a o, wi nd r o u nd
) , th e t
s a ge in m ito ti c nu cl e a r
d ivi s i o n wh e n th e ch ro m a ti n i s i n th e f o r m of a th re a d .

s por e , s
p or
'

( Gr spor a .
, se e d), a sm a ll r e pr o d u ctive b o dy o f ce rta i n a ni m a l s .

s t a ti o na ry ; au sti s a b l a d d e r
( Gr ) th e o rga n o f
’
s t atocys t , s ta t o s i s t
‘

. s ta tos, , ,

ili
q u b r a o n o f th e c r a y s
e ti fih .

t t li
s a
’
o t h , s ta t o li t/z ( Gr s ta tos,
'

. t ti o na ry
s a li tli os, t
s o ne ) , a so li d par ticl e
withi n a sta t cyst o .

s te a ps i n tép s i n a f e r m e nt whi c h r e s o lve s f at s i nt o f att y a ci d s a nd gl yc

’
,
s ,

e ri n .

s te r eil s t,r
’
i l ( L st r i li s b a r r e n) no t r e pr o d u ci ng i ts k i nd
e . e , , .

s t e r nu m s t r nu m ( G r ste r n n th e b r e a s t ) t h e v e nt ra l pa r t o f a s o m it e o f
'
’
, e . o , ,

a n a r th
po d ro .

s t i gm a s tig m a ( G r st gm a a m a k ) a ppli e d t o s po t s o f c o l o r o r to s m a ll
’ ’

, . i ,
r , ,

o e ni ngs
p .

s t i m u l u s st i m ’a l s ( L ti m u lu s a go a d ) s m thi ng whi ch e vo k e s s o m e
'

,
u . s , ,
o e

f u ncti o na l o r t o phi c r e a cti n i n th e ti ss u e s n whi h it a ct s

r o o c .

s ym b i os i s m oi os i a livi ng t o ge th e r ) t h e livi ng t o ge th e r
'
i m bi o s i s ( G r
, s . su s, ,

o f t w o d i ff e r nt spe ci e s o f o ga ni s m s
( se e p e r .

s ys tol e s i s to le ( G r su st le a d r a wi ng t o g e th e r ) t h e pe r i o d o f ti m e d u r i ng
' '

’
, . o , ,

w hi ch a r h ym i cally pu l s a ti ng v e sse l i s c o nt ra ct e d .

te l oph a s e , te l oj az ’
( Gr te los, nd ) , th e l ast ph as e of m it o ti c ll d ivis i o n
'

. e ce ,

d ur ng i whi ch t h e nu c l i e a re re -f o r m e d .

te l ’
s on, te l s o n ( Gr . te ls o n, th e e nd ) , th e l ast j o i nt of th e ab do m e n Of th e
c ra y s fih .

t e s t i s , te s ti s ’
(L . te s ti s , a witn e ss ) , th e m a le
ge r m g l a nd .

th e r m ot rop i s m , tlzer m ot '

r opi zm ( Gr . t/ze r m o, ea h t tr ope t ur n) , , re ac ti o n to
h e at
th i gmotropi sm , tai g m ot
’
ro
'

p i zm
( G r . t/zigm a , t o uch ; tr ope , a t u rni ng) ,

r e ac ti on to c o n t a ct .

th orax , ’
tao r a k s ( Gr . t/zor a x , th e b r e as t) th at
, pa r t of th e t runk s it uat e d
be twe e n th e he ad or ne ck a nd th e abd om e n .
33 6 GLOSSAR Y
t i s s ue , ti sn ’o
(L . te x e r e , t o w e ave ) ,
a n a ss o c a i ti o n o f si m il ar ce ll s with s
pe c ali
f u nc ti o ns pe r f o r m to .

tra ch e a , tr a n
’
e a ( Gr tr a cne ia ,
'

. t he wi nd pi pe ) a br a thi ng tu b e o f a n i ns e ct
, e .

tri ch ocy t s , tr i é
'
o s i st
'

( Gr . t/zr i x , h i ; n tzs a b l a d d e r ) a sm all h a i r lik e

a r us
'

, ,

t
s r u c u re t o f ce r a t i n P ro t o zo a
( se e p .

tri plob l a st i c, th re e fo l d ge rm ) , h avi ng

'
tr ip lo bla s ti k ( G r tr iploos , ola stos,
‘

. a

th re e ge m l aye rs r .

t ropi s m t o pi m ( Gr tr ope t ur n) a t ur ni ng ca u s e d b y s ti m u l u s
'

,
r z .
, , .

typh l os ol e tif lo s l ( G t pal s b li nd ; sole n t ub e ) a thi ck

,
’
' “
o r. a o , , , fo l d of th e
i nt e sti ne o f ce rt a i n a nne li d s .

u ni ce ll u l a r u ni se l u l,
’ '

ar (L . u n u s, o ne ; ce llu la , a ce ll ) , c o ns s i ti ng of a

s i ngl e c ll e .

i
u n ra m ous , a ni r é
’
m us (L . u nu s , o ne r a m u s, b r a nc h) , ha vi ng o ne b ra nc h .

u ropod ,
’
zi r opoa
'
( Gr . ou r a ,
t a il p o u s, f o o t ) , th e l as t a
ppe nd a ge of th e c r a y

fis h .

t
u e ru s , u te r u s
'
( L . u te r u s, th e wom b) , a s
p e c a i l se c ti o n o f th e o vi d uct .

va cu o e l , v a le
'
u 07 ( L d i m . . of v a ca ns, e m p y) , t a m i nu t e ve s i cl e in ce r a t in
P ro t o z o a.

va gi na , vetj i na
'
h e ath ) th e pa s age l e a d i ng f ro m th e u te r us
( L . va i n
g a, a s , s

to th e ge nit a l o ifi e i n c e rt a i n f e m a l e s r c .

ve ntra l , ve n t a l ( L v nt r t h e b e ll y)
'
r f o r pe rt a i ni ng t o th e u nd e r s i d e o f
. e e , , o

th e b o d y .

v ita l i s m , i ’
v ta l i zm (L . v i ta , li fe ) , a b i o l o gi ca l doc t r i ne ( se e p .

z oo ogy, l zo o
‘

l aj’
i
( Gr . z o o n, a n i mal logos , d i s c o u r se
) , th e sc e i nce of an i m als .

c o nj uga ti o n o f
’
z ygot e zi
( Gr yo k e ) th e b o d y f o r m e d by th e
‘

, got . zu
gon, a ,

tw o ga m e te s .
33 8 IN DEX
Ari st otl e 8 , , 267
* —2 6 8 . B i o b i o ti c f au na, 2 77 .

A r te mi a 3 2 , . i h it
B o c e m s , 23 .

t
Ar e ry : C r a y fih s ,
20 2 —20
3 ; E ar thworm ,
B i ge n ti c l w 2 8—23 2
o e a , 2 .

1 71
—1
7 4 ; H o ne yb e e ,
—
2 4 2 2 43 . B i l ogy
o , 1 .

Ar throb ranchi ae , 204 . B i p h or

o 88 ,
2 .

th —
Ar r o po d a , 6 7 , 2 2 5 23 3 ,
. B i m o s appe nd age
ra u ,

1 60— B i d s and be e
*
Ascar i s lu m br i coi de s , 3 2 , 1 63 , 1 61 r 59 s, 2 .

B l a to l
s 85 coe , 1 .

l
Ase xu a re pro d u c o n, 9 7 ti . B l t od e rm
as : C ay fi h , 1 10 r s ,
21 5; Ho ne y
i il ti
Ass m a o n 1 3 , 1 4 , 4 6 , 4 9 , 1 7 1 ,
. be e , 25 2 .

i ti
As so c a o n ne u ro n, 1 7 9 . B l t
a s om e r e , 1 0 8,

t
As e r , 3 1 . B l as o t ph ore , 1 84 .

t l
As ra r ay s, 3 1 . B l as o t po e r , 1 86 .

tt h
A a c m e n ce 25 t ll ,
B l as u a ,t l 1 0 8, 1 1 0, 2 29 : Ea r th
t t
Au o om y, 2 20 . wo m r ,
1 85 *
Gr a nti a , 1 49 ,

vi i
A o d ng r e ac o n : E u gle na, 8 7 , ti 88 Hydr a , 1 3 7 .

P ar a m e ci u m , 7 3 76 . B l oo d , 1 1 1 : C r ay fih s , 20 1 —20 2 E art h
w o rm , 1 71, Hone ybe e , 24 2 .

t i
B a c e r a , 2 9 , 4 6 , 66 , 8 1 , 25 9 . B l o od v l e s se s , 1 1 1 : C r ay fi sh , 20 2 ;
*—
B ae r, K l
a r E vo n, 2 7 1 , . E ar thw m or , 1 7 2 1 75 Hone ybe e ,

B ala nti d i u m coli , 8 2 . 24 2 .

l
B a f ou r , F ra nc s M , 2 7 1 , 2 7 2 i . . Bo dy vity (
ca se e C oe lorn) .

B ar na c e , 2 2 5 l . B ody w ll 4 a , , 5 : Asca r i s, E ar th
i
B a rr e r , 2 7 8 . wo rm , 1 68 .

l i
B a sa d sk 1 1 7 , 1 23 , . B om bus , 266 .

i
B a s po d e , 1 9 7 , 1 9 8 it . B o ne s , 1 01 .

B a e so n, W , 2 90
t . . t
B o a ny, 1 .

v
B e a e r , 3 00 . Bou on t ,

B e e : ca r pe n e r , 2 6 4 ; e a f cu ng t l tti ,
2 64 ; B ra c hi o po d a , 9 .

i i
m n ng, 2 6 5 —2 66 so c a , 266 il so li B ra in : C r ay fih s , 20 5 ; Ear thw o rm ,

t a y 6 4— 6 6
r ,
2 2 . Hone ybe e , P la 4

B ee gl 56 ue ,
2 . na r i a , 1 56 .

B e hive : l ni ng
e c ea , 25 7 ; gu a r d ng , i 25 7 i l h
B ra nch a c a m b e r , 2 0 4 .

25 8 ; nu m be r of i ivi
nd d u a s i n, l 25 8
°
B r a nc a hi l fil
am e n s 2 0 4 t , .

v ntil ti ng 5 7
e a , 2 . B r a nc a a , 2 2 5hi t .

B ee l se 5 9 ou , 2 . i l
B r a nchi ocar d a c ca na s , 203 .

B e m il k
e 53 , 2 . Bra n hi o
c di c g ve s ca r a r oo , 1 95 .

B e e m o th 5 8— 5 9 , 2 2 . B ra nchi o t git e 95 s e , 1 .

Be e s t 58 co u s , 2 . B r a ula cce ca 2 5 9 .

Be e t e e 58 r ,
2 . B r e e d ng a b s, C ray s , 2 1 1 —2 1 4
i h it fih
B s a nd fl owe rs 2 6 — 6 4
ee , 2 2 . i
B r e e d ng se aso n, E a r o rm , 1 8 2 thw .

B ees i n ge n r l 60 — 6 6 e a , 2 2 . B ro n, 3 4 w .

B tl 3
ee e , 01 . B ryo z oa , 5 , 7 .

B h avi o : Am b 5 3 —5 8 ; C yfi h
e r e a, ra s ,
221 l
B u cca po u c , 1 6 9 h .

22
4 E a thw m 8 6— 8 9 ; E gl n r or , 1 1 u e a, 87 i
B u dd ng, 1 5 , 2 7 7 : Gr anti a , 1 4 8
Hydr a , 1 2 7- 1 3 2 ; P a r a m e ci u m , 73
—
79 . 13 3
—1
3 4 ; P r o ozo a , 80 t
B e ne d e n, Ya n, 3 6 . 1 52 .

iht
B c a , 27 1 . l
B u m b e be e , 2 6 2 , 2 6 6 ; gue s t , 29 7

il t l t
B a e r a sym m e ry 4 , 5 , 6, 7 , 1 5 8 1 86 , ,
. i
o r ch d , 2 6 2,

i
B na ry divi i s on, 9 6 , 1 1 2 : A me ba 5 1 *
, tt fl
B u e r y, 2 7 9 , 3 0 1 .

E u gle na 8 7 ; H ydr a 1 3 3 *
,
.

P a r a m e c i u m 6 7 *—68 . Ca lcif e rou s gland s , 1 6 9—1 7 0

 I N DEX 339

C a m bar u s afli ni s , 1 9 3 . be a r e r of he re d it ary q ualitie s, 3 2 ; in

C a m bar us pe llucid us te s ti i 2 1 9 , .
g e rm p a sm le o ry , th 2 88 ; i n Hydr a ,

1 —
C a m bar us vi r ili s 9 3 2 2 4 ,
. 1 3 4, 1 3 6 ; nu m b e r i n a n m a s, i l 3 2 te

l
C a na sy s e m , o f spo ng e , t duc tion
i n nu m be r ,
ill
C a p a ry , 1 7 1 , 1 7 5 , 203 . 1 0 7 ; u n on i n f e r i z a o n, 1 07i til ti .

C a r a p a ce , 1 9 5 h t
C ro m o r o p s m , 5 4 , 5 6 i .

h
C ar bo yd ra e 2 2 1 7 0 t , , . h li
C ry s a s 2 5 4 , .

C a rb o n i n p ro o p a sm , 2 1
,
t l . h l
C y e 25 3 , .

C a rbo n d oxi d e , 5 0 i . h
C y m e 2 40 , .

C ar ci nus mae na s , 2 9 9 . ili

C a 2 8 1 5 4 , 1 5 5 ; P a r a me ci u m, 6 1 —6 2,
, ,

i
C a r d a c s om a c t
200 , 20 1 h , .

ttl
C a e o f P a ragu a y , 2 8 1 . C ili t 8
a e , 2 .

Ce ll2 6—
, 33, de n on 3 5 ; d v fi iti , i i C i l ti n 4 6
rcu a o , , 4 9, 1 1 1 : C ra y fih s , 203

i 4 E rthw
—1
s on 29 ,
f o rm 2 9 m or ,
20 a o rm , 1 7 1, 1 74 7 5 ; Ho ne y
p h o lo gy, 2 6 —29 ; num b e r , 2 9 ; o r g n, i i b e e , 2 4 2—2 43 .

33 , p h i l y s o o gy . 2 6—2
9 ; i
s ze . 29 ; C i rcu m oe so p h a g e a l co nne c e , 20 5 tiv .

it
un ,
1 2, 2 6 ; w ll a , 2 8, 35 . C i rcu m ph a rynge a l co nne c e, 1 77 tiv .

C e ll the o y 3 4—3 6 r , . i
C r rus 1 5 6 , .

C ll lose 93
e u , . l
C am , 6 .

C e m e nt gl nd s 3 a ,
21 . l
C a ss 3 , .

C nti pe d e 6
e 5 , , 22 , 2 27 . C a ss ca o n, 3 —7 Ho ne ybe e , 2 60 —2 6 1
l ifi ti .

C e nt l n rv ra y t e ous s s e m : C ray fih s , l w
C a ,
5 E rthwo m l v i i l
—1 —
7 8 ; Ho ne y be e , d sco d a 1 0 8 *
20
°
a r , 1 77 C e a a ge 1 0 7 1 1 0 , , ,

24 1 2 44
—2
45 . equ l p ti l
a 8;, p fi i l ar a , 10 su er c a ,

t
C e n r o so m e , 2 8 3 0, 3 , 1 , 1 04 . t tal un qu l 08 o , e a , 1
*

t h
C e n r o sp e r e , 28 3 0 , . C l e v g c vity
a a e a , 1 10 .

h l th
C e p a o o r ax 1 9 4 , 1 9 5 , . C lit e ll m 65 82 83
u , 1 , 1 , 1 .

C e r a ti na d u pla , 2 64 . C l ca 1 4 5
oa , ,

vi l
C e r ca gr oo e , 1 9 5 v . C ly p e 3 u s, 2

t
C e s oda, 1 5 8 . C ni d bl t o 3 as
24 , 1 20 , 1 2 , 1 .

h t
C ae opo d a , 1 90 . C ni d il oc 3 , 1 20 , 1 2 .

hl
C e a , 1 98 . C o g l ti n
a u a o ,
2 2 , 20 2 .

hl
C e i pe d 1 9 6, 1 9 8
, . C o o n : E thwo m
co H o n yb ar r , e ee ,

h i
C e m cal co m p os o n o f iti living o rga n s m s i 253 ; P la nar i a , 1 56 .

1 1 —1 2 l t t —1
. C oe e n e ra a , 5 , 7 , 1 1 6, 1 1 9, 13 9 43 , 1 5 8,
C he m tro p i o sm , 53 : A m e ba , 5 5 *
C ray
fi sh , 2 2 2—2 2 3 ; E a r o rm , thw 1 8 7 —1 88 l t
C oe e n e r o n, 13 9 .

Hydr a , 1 3 2 ; P a r a me ci u m, 78 . l
C oe om , 4 , 5 , 6 , 1 1 1 1 3 8, 1 58 1 59 , ,

hl
C i o pod a , 2 2 7 . A sca r i s , 1 60 , 1 6 2 1 63 ; C ra y s 200 ; ,
fih ,

iti
C h n, 1 9 4 , 2 3 4 . E ar orm thw 1 68 1 86 ; Ho ne y , ,

Chla mydomo na s 93 94, 9 6 , , . b e e 2 3 9 ; Le e c

,
1 90 ; P la na ri a , 1 5 8 h ,
.

C hlo rogo ge n ce s 1 69 ll , .
l t
C oe o m a a , 5 , 7 , 1 1 1 .

hl h l
C o r op yl I 7 . 1 8, 8 4 . 8 5 . 88. 93 . 9 4 , 9 7
. . l i fl i
C oe om c u d , 1 7 1 .

hl l t
C orop as , 2 8 . h
C o n, 3 5 .

h
C o a no cyt e , 1 47, 1 5 2 . l
C o o n 2 40 , .

h
C o rd a , 6 . ll ll
C o a r ce s, 1 47, 1 48 .

h t
C o r da a , 5 . ll i
Co o d, 2 2 .

h i
C o r on, 2 5 1 . j ll
C om b e y 5 , .

h t
C r om a i n, 3 0 , 3 1 3 2 ; co nt nu y o f , , i it li
C o m m e nsa sm 2 9 7 , .

33 it li
C o m m u n y f e o f be e s 2 6 6 , .

C h o m a to ph
r o re , 81 , 84, 85 , 9 3 . C om p a r a tiv e a na o m y , s o ry o f , 2 7 0 t hi t
C hr om osom e , 6 9, 1 03 , 1 04 , 1 0 5 , 1 0 6 ; 27 1 .
3 4 0 IN DEX
ti
C o nd u c o n, 1 7 8 . i
C uv e r,
C o nd u c tivit
y, 5 6 . C yclops 1 2 5 , 23 2, .

j ti
C o n uga o n, 1 1 2 , 1 1 3 , 1 1 4 : o f c r o m o h l i
Cyc os s 66 , .

so m e s, 1 03 ; P a nd or i na , 9 4 ; P a r a Cypr i pe di u m ,

—
me ci u m 6 7 68 7 1 , Cypr i s ,
, 7 2. 7 3 ,

C o nne c e tiv ti
ssu e , 1 1 1 . t
C ys , 9 6 : Am e ba , 5 2 ; E ugle na , 86
i hi
C o no d a r s, 24 8 . 87 .

i
C o nsc o u sne ss 5 7 , . Cy toph ynx ar , 5 9,
ti it
C o n nu y o f ge rm p a sm , 2 88 l Cy t p lasm
o , 2 6, 2 7 , 2 8 .

2 89 .

t
C o n ra c e til
e o ry of o com o o n th
of l ti D arwin C h arl e s 8 , , , 2 6 2, 2 73 2 7 8,

A m e ba , 4 2—4 5 . 2 93 , 2 94 . 2 9 5 . 2 9 6

t til v
C o n r ac e a cu o e : Am e ba — 1
39 4 , l , De a th 9 8 , .

50 ; Chla mydomo nas 93 ; E u , De e r 3 00, .

gle na, 8 1 , 8 2 , P a r a me ci u m, 6 0 D e lli nge r, O P 4 3 4 5 . .

, , .

63 6 7 6 8 ; Volvox, 9 7 , . D e rm a l e p ithe li u m 1 4 6, 1 5 1 ”
,

C o pe E D , 2 7 0
, . . . De rm a l po re s 1 5 2 , .

l ti
C o pu a o n : C ra y s 21 2
*—
2 1 3 ; Ear fih , th De t e r m i na nt , 2 8 8 .

wor m , 1 8 2 Ho ne yb e e , 2 5 0—2 5 1 . D e v i lfi sh 6 , .

l t
C o p u a o ry o rga ns o f Ho ne ybe e , 2 3 9 , D i a ph r agm 2 4 2 , ,

Di a st a se 1 7 0 , .

l
C o ra , 5 1 3 9 , 1 4
, 43
—
2 1 . D ia to m 4 6 , .

l
C o rp u s c e s 1 7 1 , 2 0 2 , . Didi ni u m ,

C o xa , Dige stion 1 3 1 4 4 6 ; e xt r ace ll u l a r

, , , , 1 27

it
C oxo po d e , 1 9 7 , 1 9 8 . i nt r a ce ll l u ar , 1 27 : A me ba , 4 9 ; C ra y
C ra b 6 , , 2 25 . fi sh , 20 1 E ar ; thw o rm 1 70 ; Gr a nti a ,
,

fih
C r ay s , 1 93 — 2 2 4 , 2 7 6 2 7 7 , 3 0 6 ; a d a p , 1 48 Hone yb e e , 2 40 ; Hydr a, 1 2 7 ;
t a t i o ns 2 8 2 ; a na om y 1 9 4—2 1 1 ,
, t , P a r a me ci u m, 6 7 .

a ppe nd a ge s 1 9 5
*—
2 00 a u o o m y,
, , t t Dige s tive gl a nd ,
20 1 .

2 20 —
— h i i tiv t —
DC fi h
*
2 2 1 ; b e a v o r , 2 2 1 2 2 4 ; b r e e di ng g es e sy s e m : As car i s , 1 60 1 6 1

h it
ab s , 2 1 1 , ci r cu a l y ra s ,
2 00 —20 1 ; Ea r thw o rm ,

tion, 203
—20
4 ; d ge s e sy s e m i tiv t 1 6 9 —1 7 0 ; Hone ybe e , 2 40 ,

2 00 — 20 1 ; e m bryo o gy , 2 1 4 , 2 1 5 2 1 6, l 24 2 ; P la na r i a , 1 55, 1 58 .

e xcr e o ry s y s e m , 20 5 ; m u scu t t D i oe ci o u s 209 , .

lar sys e m , t ne r o us sy s v D i p l obl a stic 4 5 7 1 1 0 1 5 9 , , , , , .

te m, 205 ; r e ge ne r a o n 219
*
ti , D i plo po d a 2 2 7 , .

2 20 ; r e p ro d u c e sy s e m 2 09 , tiv t , D i r e ct ce ll d i v i s i o n 2 9 3 2 , ,

re sp r a o ry sy s e m , 2 0 4 ; i t se nse t Di sco nti nu o us d i st ri bu ti o n 2 7 8 2 7 9

—
,
.

o rga ns 20 5 2 0 8
—
,
a scu ar sy s e m , 20 1 v l t Di sco nti nu o u s va r i a ti o n 2 8 1 , .

2 04 . D i s e a se s o f b e e s 2 5 9 , .

C ro p, 1 69 1 7 0 , . i
D pe s
s r i o n 2
—
7 7 27 8 ,
.

C r o ss po na o n 2 4 6lli ti , ,
26 2, 264 . D i s si m il a ti o n 4 6 4 9—5 0 6 7 , , , .

t
C r u s a ce a , 1 2 5 , 2 1 9 , 2 2 6, 2 2 7 , 2 2 9, D is t a l 1 1 7 , .

23 2 . D i st r i b u ti o n o f a ni m al s : i n Sp ace ,
275

tl
C ry s a , 2 8 . 279 ; in tim e ,
279

t ll i
C rys a o d , 2 2 . Do rsa l , 59 ; a bd o m in l a t
a r e ry,

t h
C e nop o r a 5 7 , , . 20 3 ; b l oo d ve sse l ,
1 6 8, 1 73 ,
24 2

C ule x, 88 9 2 , . po re , 1 68 .

ti l
C u c e , 1 0 1 : Asca r i s 1 6 3 C r ay s , , fih Dr one hone yb ee ,
23 4, 2 54 ; ce ll
,

1 9 4 ; E ar o rm 1 68 ; thw
E u gle na , ,

8 2, Ho ne yb e e 2 3 4 ; Hydr a , 1 1 9 ; ,
Duj ar d i n 1 9 , .

P ar a meci u m, 5 9 , 61 . Dy se nt e ry a me bi c, 8 2 ; , of be e S. 25 9
3 4 2 IN DEX
t
Ex e nsor m u s c e , 209 l . F oo t Hone yb e e , Hor se , 2 93
"

i ti
E xt nc o n o f a n m a s , 2 9 5 i l . Hydr a , 1 1 7, 1 1 8 .

Ey e : C ra y s , 2 0 5 —20 8 ;fih
Hone y Fore 3 05 l , .

*
b e e . 23 5 . 24 5 Fo rm , o f o r ga n sm s, 1 1 i .

E ye br u s , 23 7 h . il
F o ss , 2 , 2 7 9 ; nu m be r , 5 , 6 .

Eye s po . 8 1 . t
84 . 9 4 . 9 7 ; P la nar i a . l
F o u b r oo d , 2 5 9 .

i
F u s o n nu c e u s , 7 1 , l
l
Ga a p agos I s a nd s, 2 7 8 l .

F ac o r s o f ab a , 2 7 5 —2 7 6
t h it t . l
Ga e n, C a u d u s 2 68 l i , .

F ae-
ce s , 1 3 , 1 4 , 1 5 5 , 2 4 2 . Galle r i a me llone lla , 2 5 8—2 5 9 .

F a m y, 3 il . lt
G a o n, F r a nc s, 2 8 9 i .

ti
F a gu e , a s m u u s , 1 6 ti l . t
Ga m e e . 90 . 9 3 . 9 4. 9 6. 1 13 .

F at , 2 1 , 2 2 1 7 0 ,
. t
Ga m e oge ne s s, 1 1 3 i .

F a u na : o f s a n d s , 2
—
8 5 2 86 ; il t abu l at e d , G a ng o n, 20 5li .

27 7 . Ga s i n p ro o p a sm , 2 1
,
t l .

Fe m a e , 9 8 l . t
Ga s r ae a , 2 2 9 .

F e m ur , t
Ga s r a ca l vit
y,
t
Fe rm e n , 2 5 1 70 , . t
Ga s r a e p l ith li
e u m , 1 4 6,

Fe r tili ti
z a o n 1 04, , 1 07 ; h
c r om o t
Ga s r c m i ill
2 00 ,
.

s o m e s i n 1 0 7 ; nu c , li
e in ,
1 06 : As ca r i s , Ga s r ot lith
—
200 20 1 , .

1 6 1 ; C r ay fih s , 21 4 ; Ear thw orm 1 85 ;

, t
G a s ro as cu a r cav l vit
y, 5 , 1 1 9, 1 27 ,

E ud or i na , 9 6 ; Hone y Gr a nti a , 1 4 9 ; 1 39 .

b e e , 2 5 1 ; Hydr a , 1 1 4 , 1 3 7 ; P ar a me Ga s t l ru a, 1 0 8, 1 1 0, 1 1 1 ,
1 3 7,
oi u m 6 7 , 7 1 , 1 1 2 ; P la nar i a , 1 5 6 ;
,

P la s m odi u m , 9 1 , 1 1 3 ; Volvox 9 8 , ,
t l ti
Ga s r u a on, 1 10 .

1 1 4 . l
Ge m m u e 1 5 2 , .

Fe v er e s tiv o -au u m na t l 89 ; ma l ilar a , t l t

G e ni a a pe r u r e ,
1 61 , 2 1 0, 21 1 l
c o aca ,

88 , 8 9 , 9 1 —9 2 ; p e r n ci o u s , 8 9 ; uar i q 5 6 ; p ore ,
1 1 5 4,
t a n, 8 9 ; e r a n, 8 9 ; y e o , 80t ti ll w . G e nu s, 3 .

i i
F s s o n, 1 5 A me ba , 5 2 ; E u gle na , Ge ob i ot i c f au na , 2 7 7 .

8 7 ; Hydr a , I 3 3 ; P ar a m e ci u m ,
*
hi l i t i ti
Ge o gr a p ca d s r b u on, 2 9 3 .

6 7 —6 8 ; P la na r i a
*
, 1 56 . Ge gr p c
o a hi l i l ti
a s o a o n,
2 8 —
5 2 86 .

l ll t
F ag e a a , 8 2 . l i l i
G e o og ca p e r o d s 2 7 9 , 2 80 ,
.

Fl a ge ll m u , 80 , 8 1 , 85 , 86, 8 7 93 , ,
t i
Ge o r o p sm 5 4 : P a r a me ci u m , 78 ,
.

Ge r m b a nd , Ho ne yb e e , 2 5 2 .

Fl m ll
a e ce ,
1 55, ll
Ge r m ce s 9 8 1 0 2—1 0 7 1 1 4 1 1 5 , , ,
.

F lat w m —1 60 l ifi c ti o n, l
Ge r m aye rs, 1 0 8 , 1 09 1 1 0 , 1 1 1 , 1 5 4,
or s, 5, 1 53 ; c ass a

1 58 ; co n r a s e t t d with Hyd r a, 1 59 .

l
F e a 298 , . G e r m p a sm 9 8 l ,
1 02 .

l
F e xor m u s c e 209 l , . Ge r m p a s m l
e ory , th 2 8 8 —2 8 9 .

F g li ht 23 8 , . Ge r mi na es c e, lv i l 1 06 .

lw
F o e r s a nd b e e s 2 6 2 2 64 , , . Ge ryoni a , 1 4 2 .

l t ti
F u c u a ng a r a o n, 2 8 1 v i ti . G e sne r , C o nr ad , 2 6 9 .

Fo l 3 6 ,
. i tfi
G an b e rs, 1 7 8 ,
li
F o a ce o u s a ppe nd a ge , Gill 95
s, 1 , 20 4 .

F oo d : Am e ba , 4 6 ; C r a y s 2 0 1 ; E a r fih , th Gi d
z z ar , 1 69 .

w
or m , 1 70 ; E u gle na 85 ; Gr a nti a , , Gl nd c ll
a e s, 1 69 .

1 4 8 ; Ho ne yb e e , 2 4 0 , 2 5 6 Hydr a 1 2 5 ; , Good sir , 3 6 .

P ar a me ci u m 6 6 ; P la na r i a 1 5 5 , , . ti
Gr a f ng, 2 4 : E ar thworm ,
Hydr a,
F ood v
a cu o e , 1 4 8 : A m e ba l
47
*
,

P a r a m e ci u m , *—
66 67 . Gr a nti a cili a ta , 1
—1
44 4 9 , 27 7 ; t
a na o m y
 I N DE X

1 45 . 4 8 ; e m b ry o ogy ,
1 l Ho l oz o i c nu t r iti on, 8 5 .

1 49, nu t iti r o n, 1 4 8 ; r e p rod u c Hom o 3 , .

ti o n, 8 —1 Ho m o logou s st r u ct u re s
1 4 49 .
, 29 1 ,

Gr a nu l ar the o y r of pr o t plo a sm c i s t r uc Ho ne y 2 5 6—2 5 7 ; ce ll s

, , 25 6 ; co m b, 25 4,

t ur e , 20 . 25 6 ; flavo r, 2 5 7 ; s a c, 2 4 0,
—2 2 2
Gra vit
y, 5 4, 221 . 257 .

G re e n g a nd , l 20 5 . Ho ne yb e e , 1 5, 23 3
—2
59. 275. 27 7. 2 84 ;
G ro wth , 1
—
3 15 ; by a ccre tio n , 13 ; by ac tiviti e s of w o r k e r s, 2 5 4
—2
58 ; ad ap a t
i ntu ss u s ce p ti o n, 1 3 : A me ba , 50 ; ti n 8 ; a n t om y 3 4
o s, 2 2 a , 2

P a ra me ci u m , 6 7 . di 59 ; d n
se a se s , mb y 2 ro e , e r

lt
G ul e : E u gle na , 8 2, 84 ; P ar a me l gy 25
o o , e n mi 5 8— 59 1 e e s, 2 2

oi u m, 59, 67 . i n ti nct 3 3 —3 4 ; m t am ph i
s s, 0 0 e or os s,

Gymna me ba , 8 1 .
5 4 ; q ue e n 33 2 . 2 .

Gyna nd ro m o r p h , 2 60 , 2 6 2 . re p d ti
ro n 4 8 —
5 uc wo k e o , 2
*
2 1 , r r,

H it
ab ti
f or m a fi sh 2 23 4 o n, C r ay ,
—2 2 . Hoo k e 3 4 , .

Ha b it a t 2 —
7 5 2 7 7 2 84
, , . Hor se a nce st ry 3 ; e vol utio n
, , ,
29 2

Ha e ck e l 2 2 9 ,
. Hu ggi ns 2 2 4 , .

Hmm at o chr o m e 8 4 , . H u nge r 1 6 , .

Hmm oglobi n 90 1 7 1 , , . H u xle y 1 9 1 9 3 2 6 2 , 2 7 0

, , , .

H a i r o f Ho ne y be e , Hyd ra di ce ci a 1 1 6 , .

Hali ctu s 2 66
,
. Hydr a f u sca 1 1 1 1 1 6—1 3 9 , , ,

Ha ll e r Alb re cht 2 7 2 —2 7 3
, , .

H a l ob i o ti c f a u na 2 7 7 , . 2 7 8, 2 84 , 29 7 ; a na o m y , t 1 1 7
—1 25 ,

H a r ve y Wi lli a m , 27 2 , . h vi 7 be a o r, 1 2 ,
1 3 2

Ha t chi ng : C r ay fi sh 2 1 6 2 1 7 *
Ho ne y , , co nt st e d with P l n i
ra a ar a, 1 5 9 ; nu ti r

ti n 7 ; e ge n ati
— *—
b e e 2 5 3 ; Hydr a 1 3 7
, , . o , 1 2 r er o n, 1 3 7 1 39;
He a d Ho ne yb e e 2 3 5 23 6
,
*—
, . rep d tio 3 3 3 7
n
ro
— uc ,
1
*
1 .

He ari g Ho e yb e e 2 4 7 2 4 8
n n ,
—
, . Hydr a gr i s e a 1 1 6 , .

He a rt : C rayfi sh 20 2 ; E a rthw o r m , ,
Hydr a vi ri di s 1 1 6 1 1 7 1 2 7 1 4 3 , , , , .

1 7 3 ; Ho ne y be e 24 2 ,
. H yd r oge n i n p rot op l a sm 2 1 , , .

He m o spo r i d i a 8 2 , . Hyd r oz oa 1 3 9 1 4 1 , ,
.

He p ati c d u ct 20 1 20 2—20 3 , , . Hym e nopt e r a 2 60 2 6 1 , ,

.

He re d ity 2 8 4—2 8 5 2 8 7 —2 90, a cq ui r e d ,

Hy po st om e , 1 1 7 1 1 8 *
,

c ha ra c e r s , t 2 8 7 —2 88 ; a nd e v l uti o o n,

2 7 5 —2 9 7
i n P a r a me ci u m, 7 9 ; ; me th I d 288 .

od s o f s u d y , 2 8 5 t . I d 1o p la sm , 2 88 .

h
H e r m a p r od e , 1 4 9 1 5 6 it , . l
I e um , 2 40 ,

He rmi cr ab , t 29 7 . I m ago , 25 2 .

He ro n 3 00 . it ti —
I m a o ns, o f Am e ba , 4 5 4 , 4 7 49 6 — .
,

He rt wi g 3 6 ,
. I ncur r e n ca na , 1 4 5 , t 152 l .

He t e rom or pho s i s 1 8 9 2 20 , , . I ncu rr e n po re I 4 5

*
t ,
.

Hi be rna ti o n 2 7 6 , . i
I nd re ct c e d v s o n 2
—
9 3 2, ll i i i ,

Hi ghway s o f d i spe r sa l 2 7 8 , . I nd ivi

dua 3 , 1 5 7 1 l , , ,

Hi r u d i ne a 1 90 , . l t
I nf e ro a e r a ne r e , 20 5 l v .

H i st ology, 2 ; hi st o r i ca l 2 7 1 ,
i
I nf u so r a , 80 , 8 1 , 1 0 1 .

Hof e r , 4 9 I nge s o : Ame

n ti b a 6 — C r ay
. 4 49 , ,

Ho f m e i st e r 3 5 , . fi sh 2 0 1 ; E a rt
,
o rm 1 70 ; Gr a nti a , hw ,

—
Hol m e s S J 2 24 , . .
, . 1 48 ; Hydr a , 1 2 5 1 2 6 ; P a r a me ci u m,
Ho lob l a sti c e gg 1 0 8 , ,

Ho l ophyti c nu t r iti on, 8 5 . In h it ance

er q i ed h : ac u r c a r a c e r s, t 2 87

Ho l o t r i chi d a 8 1 , . 2 88 ; a nce st r l 2 89 ; g a ,
e rm p l a sm in .
3 44 I N DE X
2 88 ; M e nd e l ’
s la w of , 2 89 of i
L p s, o f Ascar i s , 1 60 .

i nsti nct s , 3 05 . Li r i ope , 1 4 2 .

I nse ct a, 6 , 2 2 5 , 2 2 7 , 2 3 3 , 2 6 1 . iv
L e r , 20 1 .

ti —
I ns nct , 3 0 3 3 0 5 ; Ho ne yb e e , 3 03
—
3 04 . iv fl
L e r uk e , 1 5 8 .

t i
I n e r br e e d ng, 2 86 . t
Lo b s e r , 23 0 2 7 7 , .

t ll i
I n e r ce ular d ge s o n, 1 5 5 ti . ti
Loco m o o n, 1 7 , 1 8 : Ame ba , 4 1 —4 6 ,

I nt titi l ll
e rs a ce s, 1 1 9, 1 21 , 1 23 , 13 4 . C r ay s , 2 2 1 ; E u gle na 8 5 ; fih ,

I nt on l fi b
e rz a e r s, 3 1 .
*—
Hydr a 1 2 9 1 3 0 ; P ar a me ci u m, 64 ,
,

I nt t i n
es A e : sca r i s , 1 60 , 1 6 2, 1 63 ; 66 ; P la na r i a , 1 5 4 .

C ay fi h r s , 20 1 ; E ar thw o rm , Loe b , J 3 0 4 .

6 9 ; Ho ne yb e e ,
1 2 40 , 25 2 . Lu m br i cu s te r re str i s , 1 6 4— 1 90 .

t ti
I n e s no — e gu m e n t t a ry b l ood v e sse l , 1 72
*

1 73 . M a cr oga m e t e 90 9 6 9 8 , , ,
.

t ll l i
I n r a ce u a r d ge s o n 1 4 8 , 1 5 5 ti , . M a croga m e t o cyt e 90 9 1 , , .

t ti
I n u s su sce p o n, a m e o d o f gro th wth , 50 . M acro m e r e ,

v t t
I n e r e br a a , 5 . M a cr onu cle u s 6 7 68—7 1 8 1 , , , .

it ilit
I rr ab y, 1 6 1 7 , 5 6 ,
. M a l a costr a ca 2 2 6 2 2 9 , , .

l ti
I so a o n : ge o gr a p ca , 2 8 5 —2 8 6 ; hi l M al ar i a 88 8 9 9 1 9 2
, , , ,
.

l
o ogi ca l 286 ,
. M a le 9 8 , .

M al p ighi 2 4 2 2 7 1 , , .

J e lly fi sh 5 1 3 9 , , . M a l p ighi a n t u b ule 24 2 253 , , .

J e nni ngs H S , . .
, 3 8 , 4 2 . 4 6. 5 3 .
M a m m ali a 3 ,
.

M a nd ibl e : C r a y fi sh 1 96 1 97 , , ,
21 6;
K y op l m
ar as ,
26 . H o ne yb e e ,
23

K yoso m
ar e , M a stigo pho ra 80 8 1 8 2 , , .

K t b li m — M a t u r a ti o n 1 0 3 1 0 5
a a o s ,
1 3 , 4 9 0 5 .
, ,

Ki ng b cr a , 2 25 , 2 2 8 . M a t u r ity 1 4 7 1 7 2 , , , .

KOlli k e r 3 5 ,
. M a u ch a m p she e p 2 8 1 ,
.

Kr e i d l 2 0 8 ,
. M axilla 1 9 6 1 9 7, , ,

K r o po tk in , 3 01 M a xill ar y p a l p u s 23 ,

M a xilli pe d 1 96 1 97 , , ,
1 98 .

La b i l p lp
a a u s, 2 3 M e chani sti c the ory o f li f e ,
2 3 —2 5 .

La b i m 3
u ,
2 M e d u sa 1 3 9 , , 1 42 .

La br u m , 2 1 6 , 2 3 M e gacki le acu ta 2 6 4 , .

Lam ar ck , 2 7 3 2 8 7 , . M e la ni on 90 , .

v
L a r a : C r ay s , 2 1 6 , 2 1 8 ; H o ne y fih M e m br ane ll 6 2 , ,

be e 2 5 2, , M e nd e l Gr e gor 2 7 4
, ,
.

t l l v *
La e r a b oo d e ss e I 7 2 , 1 7 3 l , . M e nd e l s la w o f i nhe ri t a nce
’
,
2 89

t l li
La e r a ne , 1 60 , M e nt u m 2 3 5 , .

t l v
La e ra ne r e , 2 0 5 . M e r ob l a sti c e gg 1 0 8 *
,

v
La e ra n, 8 8 . M e r o z o it e 90 1 1 3 , ,
.

h
Le e c , 1 90 , M e so b l a sti c b a nd s ,

fih
Le g : C ra y s , 1 98 ; Hone ybe e , M e so d e r m 1 1 0 1 1 5 o rga ns , ,
a rI SI ng f ro m,
*—
2 6 3 2 8 3 . 1 1 1 E ar thw orm , 1 86 ; Hone ybe e ,
ih
L c e n 29 7 , . 25 2 ; P la nar i a , 1 54 .

i i
L e b g, M e so gle a 1 1 9 1 25 , , .

i
L fe i i n 8— n m n 8 —2 h M e sohi ppu s 2 9 2
or g
, , 1 0 ; p e o e a , 5 .
, .

L f e cy c e , 1 4—1 5 , 80 ; P a r a me ci u m , 7 1 —7 3
i l . M e so m e r e ,

i l
L gu a , 2 3 M e sothora x 2 3 6 ,
.

Li m nob i o t i c f au na 2 7 7 , . M e t ab oli sm I 3 I 7 46 7 2 , , , , .

i i
L nn , M e t age e si s 4 1 4 2
n 1 1 —
, .

Li nna u s, C a r 4 , 2 70 l , . M e tam e re 4 1 65 , , .
3 4 6 IN DEX
Ord e r , 3 . Pe r i ca d i al sinu s
r , 20 2 , 2 4 2 .

Orga n, 1 1 1 . P e r i patus , 2 25 , 2 27 .

v l ti
Or ga ni c e o u o n 2 2 8 . Pe r p i he al ne v ry te m r ous s s 1 77
-
1 78
Orga n sm c ar a c e r s cs 1 0 —
, , .

i h t i ti , 16 ,
. Pe r s i t lti c c nt ti n 7 5
a o rac o , 1 .

Og
r a n z a o ni ti
f
o b o di e s , 1 2
-
13 . Pe r i t m 59
s o e , ,

Orga noge ny , 1 0 8, 1 1 1 . Pe r it ne um 68
o , 1 .

i i
Or g n o f o e r s, 2 4 6 fl w . Pe r ivi sce al i nus 3 r s , 20 .

Or ohi ppus 2 9 2 , . P h a ry nx, 81 , 1 1 1 , 1 59 : Ascar i s , 1 60 ,

th
Or oge ne s s , i E ar thw o rm , 1 69 , 1 70 ;
Os ci lla r i a , 4 6, P la nar i a , 1 55 .

l
Oscu um , 1 4 4 1 49 , , P hot ynthe i os s s, 2 8, 84 .

Osm os s, 1 7 1 i . P h t t op i m
o o r s , 53 : A m e ba , C ray
ti
Os a , 1 4 5 1 5 2 20 2 2 4 2
, , , . fi sh , 2 23 E ar ; thw orm 1 88 ; , E ugle na ,

v
O a ry : A sca ri s C r ay s 21 I
*
; ,
fih , Hydr a , 1 3 1 .

Ear thw
orm , 1 80 1 84 1 85 , , P y h l a, o f a n m a s, 5 i l c ha r a ct e r s i tics
Ho ne yb e e 2 5 0 Hydr a , ,
1 19 p ara ng, 4
se ti .

1 3 5 ; P la nar i a 1 56 , . h l
P y o ge ny , 2 2 9 .

i
Ov d uc : C r ay s t E ar or m fih , thw , P hys ali a , 1 4 2 ,
1 6 7 , 1 80 H o ne y b e e ,
,
250 h i h i
P y s co c e m cal e o ry o f fe , th li 23 —2
5 .

P la na r i a 1 56 ,
. h i l i
P y s o o g cal co n nu y b e ti it twe e n ce ll s ,
*
Ovu m 1 0 6 ,
.
97
w ih
O e n R c ard 2 7 0
, ,
. h i l i l i l ti
P y s o o g ca so a o n, 2 8 6 .

i ti t
Ox d a o n i n m e ab o s m , 1 3 , 1 4
,
li . h i l i l t t
P y s o o g ca s a e s , 7 8 , 1 8 9 .

livi
Oxyge n i n ng m a e r 1 1
,
i n m e t ab o tt , h
P ys o o gy, 2 ; i l hi t i l
s o r ca , 2 7 2—2 7 3 .

li sm , 1 3 1 4 ; i n p r o o p as m , 2 1
, t l . i
P ge o ns, 2 93
t
Oy s e r , 2 8 3 . i
P nc e r , h
1 96 1 98 , .

P i the ca nthr opus , 3 .

P ala mon 2 20 ,
. P la na r i a lacte a , 1 5 7 .

P a e o n o ogy , 2 , 2 7 9—2 8 0, 2 9 2
l t l *
. P la na r i a m acula ta , 1 53 1 65 , 1 71 ,
l
P a e o z oo o gy 2 l ,
. 1 7 6. 277 ; a na o m y . t 1 53
—1
5 6.
P a nd or i na m or u m, 9 4 , 9 6, 1 1 0 , 1 1 3 .
ge ne r a on,
re ti
P a r a me ci d ce , 8 1 . P la nar i a polychr oa ,
P a r a m e ci u m ca ud a tu m, 1 5 , 5 9—7 9 8 1 8 2 , , , P anl t d
s , co m p a r e s, 1 7 8 with an m a i l -1 .

8 7 , 1 0 8 , 1 2 7 , 2 7 5 , 2 7 7 3 0 2 ; a na o m y ,
t , P la s m odi u m f alci par u m , 8 9 .

—
5 9 6 4 60
*
be a o , 7 3 7 9 ;
,
r — e re d h vi h P la s m odi u m m ala r i a 8 9 ,
.

*—
i t y, 7 9 ; f e cy c e , 7 1
— li
7 3 ; o co m o o n, l l ti P la s m odi u m vi va x, 8 2 88 9 2, 27 7 ,

64 , 6 ;6 n u r o n , 6 6 —6 7 ; re p r o t iti i
d sco e ry o f 8 8 ; v
fe s o ry
*—
89 9 1 ,
li hi t ,

r e p ro d uc o n 90 — t i i
*—
ti
d u c on 6 7 7 1 , , 91 ; ti
r a ns m ss o n by ,

P a ra m y um , l 84 . q t
m o s ui oe s, 8 8 8 9 90 , 9 1 , 9 2 , ,
.

iti
P a r a s c P r o oz o a , 8 0 , 8 2 t . l ti
P as d , 28 .

P a ra s s iti
m , 29 3 00
8— . P latyh e q i nth e s 5 7 1 5 3 , , , .

P a raz o a , 5 . l
P e o p od , 1 9 8 1 99 ,

P ar e i o pod 1 98 , . l
P e u r ob r a nc a , 2 0 4 hi .

it l l
P a r e a b oo d e sse 1 73 v l , . l
P e ur on, 1 95 .

P a rt h e no go ni di a , li
P ny 2 6 8 ,
.

t
P a s e u r, 9 . P li ohi ppus , 2 9 2 .

Pe a h, yb r d , 2 8 —
9 2 90 i . P o d ob r a nc a , 204 hi .

lli l
Pe c e 5 9 ,
61 , , P odocoryne ca r ne a 1 4 3 ,
.

P e na u s ,
23 2 . i
P o son g a nd Hone yb e e l , ,

i l t
P e n a se a 1 6 0 , . l
P o ar b o d e s 6 8 9 1 i1 0 6 , 1 1 3 , 1 1 4,
, , ,

Pe n s i ,
1 5 6,

ti
P e r ce p o n, 1 7 8 . Po ll e n, b ask e t ,
23 7
—2 8
3 b r us h ,
 IN DEX 3 47

co m b , the ring R e ca p it l ati n th e o ry , 2 2 8—2 3 2 .

*
23 6 23 8; ga , u o

25 6 . R e ce p t or 7 9 , 1 .

P oly cha t a , 1 90 , 1 92 . t
R e c um , 1 6 1 ,
l
P o yp . 5 . 1 3 9 . R e di , 8 .

i
P o r f e ra 5 , 7 , 1 4 9,
. i
R e d u ct o n, o f c r o m o so m e s, 1 04 h
P o r o cy te , 1 4 6 , 1 4 8 . 1 06 , 1 0 7 , 1 3 4 .

P or os por a gi ga nte a , 80 . R e e se , 1 1 7
P o r u gu e se M a n-o i -Wa r , 1 4 2 ,
t R e e x, 1 7 8—1 7 9 , 3 0 2—3 0 3
fl
t i
P o s e r or , 5 9 . R e ge ne r a on : ti C y s ,
ra 21 9
*-
2 20 fih
ii
P r a r e d og, 3 00 . t w *—
E a r h o rm , 1 8 9 1 90 ; Gr a nti a , 1 5 2
P re f or m a o n ti
e o ry 2 7 1
—27 2 th ,
.
*—
Hydr a , 1 3 7 1 3 9 ; P la na r i a ,
i t
P r m a e s, 3 . R e m ak , 3 5 .

i itiv t
Pr m e s re ak , 2 1 6 . R e p ro d u c o n, 1 5 —1 6 ; ti a s e xu a , 15 ; l
i
P r ob osc s o f P la na r i a ,
,
1 5 4, 1 55 . se xu a ,
—
15 16 : l Am e ba 5 0 —5 3 , ,

P ro nu b a m o , 2 9 8 th . 1 1 2 ; Ascari s , 1 6 1 C hla mydomona s ,

*— —
h
P r o p a se o f m o s s, 3 0 3 1
, it i . 2
9 , 93 ; C r ay s ,
2 0 9 21 1 ; E ar fih th
li
P r o po s , 2 5 6 . w o rm , 1 80 —1 8 5 ; E ud or i na , 9 6 ; E u
P r oso pyle s 1 4 5 , 1 52 g le n a 8 7; G r a n ti a 1 8 —1 49 ;
,
.
, , 4
t
P ros o m um , 1 6 5 , i 1 78 . H e yb e e ,
o n 2 —
48 25 0 ; Hydr a, 1 1 4,

P ro t ide ,
21 ,
2 2, 1 70 . 13 3
*—
13 7 ; P a nd or i na . 9 4 .
Pro th r o ax, 2 3 6, 9 6, 1 1 3 ; P a r a me ci u m , 6 7 7 I *—
,

1 1 2 ; P la nar i a 1 5 6 ; P la s modi u m , 8 9
*
P r otohi ppus , 29 2 .
,

t l
P r o o p as m , c he m i c l a co m po s ition ,
21 9 0 —91 , 1 13 ; S pondylom oru m, 94 ,
23 ; b r d ge s i o f , 29 , p pe tiro r e s of, Volvox, *-
97 99 1 1 4 ,
.

—2 ifi ity
1 9 3 ; s pe c c of, 23 ; st ct ru ure of , R e pr od u ct e or ga ns , iv
1 59 : Ascari s
19 C ray s , 209 , 21 1
*
fih
P ro t pod ite
o .
1 95. 1 97. 1 9 8. 1 99 Ear thw
orm , 1 6 8, 1 80 —1 8 5 ,

Pr o t o oa 5 z , , 7, 29 , 3 1 ,
66 , 80—99, 1 00 , 1 0 1 , Hone yb e e , 2 4 8 , 250 ; Hydr a ,
1 1 9 ; P la na r i a , 1 56 .

P r o t oz o aaa , 23 1 . v i
R e se r o r , o f E u gle na , 84 .

v ti l
P r o e n r cu u s, 1 6 9 . i ti
R e sp r a o n, I 4 : A m e ba , 5 0 ; Ea r or m thw ,

i l
P ro x m a , 1 1 7 . 1 75 ; Gr a nti a , 1 4 8 ; P a r a me ci u m , 6 4 ,
P se ud OpOd i ospo r e , 5 2 , 1 I 2 . 67 .

P se u d o po d i um , 4 1 , 80 , 8 1 , 1 24, 1 3 6, Re sp i t ra o ry sy s e t m : C r ay fih s ,
204

1 55 . Ho ne yb e e ,
2 43
*

t li
P ya n, 2 5 . Re tic l the u ar o ry of p ro t pl
o a smi c s r ue t
lv ll
P u i u s, t ur e ,

P upa. 2 5 2 . 25 4 . R h o t ro p i m
e s ,5 4, 1 3 2 .

l i
P y o r c s om a c t h , 200 , 20 1 . R hi p d
zo 8 o a, 0, 8 1 , 8 2 .

P y re no d , i
8 5 93 , . R h um b le r, 4 8 .

i —
R ob n, 2 83 2 8 4 ,
29 4 .

Q u e e n ce ll 2 5 5 , . R o ck y M o u n a n s e e p, 3 00 t i h .

Q ue e n ho ne ybe e .
23 3 .
249 . 2 5 4 . 2 5 5 R oss , 88 .

t
Ro s r u m ,
R a ce s , o f be e s , 2 6 1 . t t i
R o a or a , 5 , 7 .

i l
R ad a ca na s , I 4 5 , *
1 52 l . R o u nd o rm , 5 , 1 60—
w 1 6 3 , 29 8 .

l
R adi a sym m e ry. 4 . s. 7 . 1 5 9 t . R oy l j e lly
a , 253 .

ti
R adi a ng ca na s , o f P ar a me ci u m , 6 3 l . R u thve n A ,
. G .
,
295 .

R a e : o f r e ge ne ra o n, 1 90 , 2 20 —2 2 1 ;
t ti of

re sp r a i ti o n, 2 44 . S acculi na car ci ni , 2 99
* —3 00 .

Ra y J hn
, o , 2 69 —2 7 0 . Sa liva y gl nd r a s, 2 4 2, 2 5 3 .

Re a ct i o n s, 1 6 , 5 3 —5 4 (se e also B e ha vi or ) . Sa ltati n 2 8 o ,

1 .
3 4 8 IN DEX
t
Sal s, i n p r o o p a sm , 2 1 t l . t l
So m a o p e u r e , 1 86 .

S a pi e ns , 3 . it
Som e 4 , 6 1 6 5 , ,
.

h ti
Sapr op y c nu r on, 8 5 t iti . ll i
Sp a a nz a n , 9 .

Sar cod e , 1 9 . i l ti
Spe c a cr e a on, 8 .

h t t
Sca p ogna hi e , 1 9 7 , 20 4 . i
Spe c e s, 1 —3 .

h l
Sc e e , 5 2 . th
Spe r m a e ca : E a r orm , 1 80 , 1 81
*
thw
h
Sc iz ogony, 90 , 1 1 3 . 1 8 2 ; H o ne y b e e , 2 5 0 2 5 I
*
, .

hi t
Sc z o n , 90 . Spe r m a d , ti
1 04 1 3 5 1 84 , , .

h i
Sc l e d e n, 3 4 3 5 , . t t
Sp e r m a o cy s , o f Gr a nti a , 1 4 9 .

h lt
Sc u z e M ax 1 9 , 3 6 27 1 , , , . t t
Spe r m a ocy e , 6 8 , 1 0 3 *
E ar or m thw ,

Sc hw a nn, 3 4 , 3 5 2 7 1 ,
. 1 84 ; Gr a nti a , 1 4 9 Hone yb e e , 2 5 0 ;
l l t
Sc e rob a s , 1 4 6 1 48 , . Hydr a , 1 3 4—1 3 5 .

i
Sco r p on, 6, 2 2 5 2 2 8 ,
. Spe r m a oge ne s s 1 0 2 t 1 04 : C ray i , ,

h
Scyp oz o a , 1 3 9 . fi sh , 2 0 1 ; E a r o r m , 1 84 Gr anti a , thw
Se a a ne m one 1 3 9 297 , , . 1 4 9 ; H o ne yb e e 2 5 0 ; Hydr a , 1 3 4—1 3 5 , .

Se a cu cu m b e r , 6 . Spe r m a ogo n u m , t 1 3 4, 1 49, i

1 84 ,

Se a gu , 3 00 ll . 250 .

Se a lil
y, 6 . Sp e r m a t oz oa , 1 5 , 9 1 , 9 9, C ray fih s ,

Se a s u r , 6 q it . 21 1 ; Ea r thw or m , 1 83 ,
1 84 ;
Se a u r c n, 1 04hi . Gr a nti a , 4 9 ; H o ne yb e e , 2 5 0 ; Hydr a ,
1

Se a a nu , 5 w l t . P la na r i a , 1 5 6 ; P las m odi u m,
ti
Se cre on 4 6 5 0 , , .

t
Se gm e n : C r a y s , 1 95 ; Ea r fih th i l
Sp cu e . s. 1 44 I 4 7 I 4 S. 1 4 9 . 1 5 2 .
*—

wor m 1 6 5 ,
. i
Sp d e r , 6 , 2 2 5 2 2 8 ,
.

t ti
Se gm e n a o n, 4 6 , . i l it i
Sp nd e i n m o s s 3 0 ,
*
,

Se m na i l r e ce p a c e : C r ay s , 21 2 ; t l fih i i l
Sp nn ng g a nd , 2 5
Ear thw
orm 1 6 7 , 1 8 0, 1 8 1
*
H o ne y
,
i
Sp r a c e , l 2

b e e , 2 5 0, i l th
Sp ra p a , o f E u gle na , 8 7 ; o f P ar ame
Se m na i l v il
es c e : Asca r i s 1 6 1 ; E a r ,
th oi u m 6 5 , 66 ,
.

wo rm , 1 82 ; Hone yb e e , 2 4 8 , i
Sp re m e ,
P lana r i a 1 56 ,
. l h
Sp anc no p e u re 1 8 6 l ,
.

*—
Se nse o r ga ns : C ra y s —
20 5 2 0 9 ; E a rt fih ,
h S pondylom or u m 93 9 4 1 00 , , .

wor m , Ho ne y b e e 2 4 5 2 4 8—
, . Spo nge . s. 1 44
*—
1 53 .

Se nse s o f C r ay s
,
222 fih .

Se ns itiv
e p an 1 8 l t , . Spo ng n, 1 5 2 i .

Se nso ry ne u ro n I 7 9 *
,
. t
Spo n a ne o u s g e ne r a o n, 8 9 ti .

Se p u m t , t
Spo n a ne ou s m o e m e n s, 7 3 v t , 7 8, 1 28
*

S e r tula r i a 1 4 2 , . 1 29 .

t
Se a , 1 6 5 *
Spo r ob a s 91l t ,
.

l
Se xu a re p r od u c o n 9 6, 9 7 ti , . Spo roga ny, 1 13 .

h ll
S e , a n a ccre o n I 2 ti , . Spo rozo a , 80 , 8 2 8 8 , 8 9 1 0 1 , , .

hi
S r m p 20 8 , . Spo r o z o e , 91 , 1 1 3it .

Si e b olt v on 2 60 , ,
. t
Spo r 2 8 1 , .

i
S nu se s o f cr ay s , 203 fih , . l ti
Spo r u a o n 1 1 2 : Ame ba , 5 1 , 5 2
,
.

i
S z e , o f o r gan sm s 1 0 i , . Sp u r ,
23 7 .

lt
Sk e e o n o f spo nge s 1 5 2 , , . q i l
S u rr e 2 7 7 , .

l i i
S e e p ng S ck ne ss 8 2 ,
. t h
S ar c 1 7 1 8 , , .

ll fih
Sm e : C r ay s , 2 2 2 ; Ho ne yb e e 2 4 7 , . t fih
S ar s , 6 .

Sna il 6 2 87
, , . t t t
S a o cy s , 1 9 7 20 8 , 2 2 2 , .

Soci a i f e , 2 9 7 —3 0 2 ; o f b e e s, 2 64—2 6 6
ll . t t lith
S a o , 20 8 .

ti ll
So m a c ce 9 7 , 9 8 1 00 1 1 4 1 1 5 , , , , . t i
S e ap s n 1 7 0 ,
.

ti l
Som a c p asm , 1 0 2 . t l t
S e r na a r e ry 2 03 , .
5 5° IN DEX
Var i a ti o n , co n tinu ous o r fl t u ting
uc a , 28 1 Wa t e r , ca rr e i d by b e e s, 25 6 ; i n p ro t o

di scon nu o us, ti; i n P a r a me

—
28 1 28 2 l m
p as , 21 .

ci u m, 7 9 . W te r fl
a e a, 2 2 5 .

Va s d e f e r e ns : As car i s 1 6 1 ; C r ay s ,
.
, fih W alt . 2 5 4g
.
255 ; g l a nd s. 23 9. 25 4 ; p nc i h
E ar or m , 1 6 7 , thw
1 8 2, 1 84 ; e rs, 23 8 .

Hone yb e e , 2 4 8, P la nar i a , We i sm a nn ,
A, . 2 7 4 , 2 8 7 , 2 88 .

1 56 . Whale ,
1 0, 2 7 7 .

Va s cula r sy s e t m : C r yfi h a s ,
20 1 Whit m a n , C O . .
,
2 9 5 , 29 6.

2 04 ; Ea rthw m 7 or , 1 1 , Hone y Will 3 0 4 ,

.

be e , 2 4 2—2 44 . Wil so n E ,
. B 3 5 .
, .

Ve u m l ,
23 7 . Wi ng : of Hone yb e e , 23 6, 23 8; ve ins ,

Ve n ral , t 59 ; a bd o m in l a t
a r e ry , 23 8 .

2 00 ; b lood ve sse l ,
1 68 , 1 73 ; Wol ff , F K —
2 7 1 27 2 . .
, .

ne r ve cord , 1 7 7, 205 , th ora c c i Wo lve s 3 0 1 , .

t
a r e ry , 2 03 . Wo rk e r hone ybe e , ce ll
,
25 5
”

V nt i l
e r c e
4 ,
2 2. Worm s flat 5 ; j o int e d , , , 6; ro und , 5 .

Ve rt b t 5 6 7
e r a a, , , .

V ti gi l gan
es 9 a or s, 2 1 . Ye llow f e ve r p r it e 8 a as , 0 .

Vi r h w R d l ph
c o 3 6 , u o , 1 0, , 27 1 . Ye k s R M
r e 4 ,
. .
,
22 .

Vi c ln v
s e ra y t m er ous s s e , 20 5 . Y lk gl nd s
o , 5 6 ; pyra mi d s
a , 1 ,

Vi si on C ay fi h — 8
7 : r s , 20 20 , 222 ; Ho ne y sp he re 4 5 s, 2 1 ,
2 1 .

be e , 24 6—24 7 . Y th 4 7 7
ou ,
1 ,
1 , 2 .

it i
V al sm , 2 3 24 2 5 , , . Y cc p lli n ti o n by m oth 2 9 8
u a, o a , .

it lli
V e ne m e m b r a ne 2 5 2 , .

Volvocace ce , 8 1 , 9 2 , 9 6, 1 00 . Z oae a , 23 2 .

Volvo ci na , 8 1 . l
Z oo ch ore a , 1 43 , ll 297 .

Volvox, 8 1 , 9 2 9 7 9 9 , 1 00 , 2 7 5 —2 7 9
*-
, 1 1 0, 2 28 Z o oge ogr a phs'. 2 . .

3 01 . i
Z oo d , 1 3 9 .

Vr i e s Hug o d e
, , 2 7 4, 2 8 2, 29 5 , 296 . l i t i tion l 2 ; v l utionary
Z oo ogy, d s r b u a ,
e o ,

2 ; hi t o i l 6 7 74 ; s
o f t o —
d ay 7 4 ;
r ca ,
2 -
2 ,
2

Walki ng : Ho ne yb e e , 23 8 . sy t e m ti c
s a , 1 .

Wallace A R 3 0 1 , . .
, . Z oo phyt e 39 , 1 .

Wa nd e ri ng ce ll 1 4 7 , . 1 57
*
Z yg t o e o 93 94. 96 98 7r. 3 . . . .
10 . 1 1 .

Wa sp 2 6 1 2 64 3 00
, , , . 1 1 4, 1 1 5 .

P r i nte d i n t h e Uni te d S ta te s of Am e ri ca .