TH E

B I O L O G Y O F TH E F ROG

S A M UE L J . H O L M E S , PH D . .

N]
A SS I S T A NT PR O F E SS O R OF Z OO L O G Y I N T H E U NI V E R S I T Y

OF WI S CO N S I N

S E C O ZVD E D I T/ O N

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,
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A ll r ig /”Us r es er v ed
 C O PY RI G H T , 1 90 6 ,

B Y TH E M A M I L L AN
C C O M P A NY .

S et l
u p a n d e e ct ro t y p e d P ub li h
s ed M ay , 1 90 6 Rep r i n t ed

ly
. .

Ju ,
1 90 7 .

Nurmoun 19mm
J . S . C ash in g Co B er w ck
. i
S m it h 0 0 .

Nor w o o d, M as s , U S A . . . .
 P R E F A CE

TH E p re se n t boo k i s th e o utgrowth of a cou r s e of

lectu res del ivere d d uri n g th e p as t s i x ye a r s at th e Un i
ve rs i ty o f M ichig an Th is co u r s e w i th th e accomp a n yi n g
.

l abor a tory w ork w as b ase d o n th e f rog w h ich w as chos e n

,

a s a co n ve n ie n t for m with w h ich to i n trod uce s tude n ts to

a k n owledge of th e morph ology ,

phys iology an d li f e h i s
,

to ry of verteb r ate an i m a ls I n w ri ti n g th i s book I h a ve

.

h ad in m i n d th e n eeds of s tude n ts s uch as mo s t of thos e

,

t a ki n g thi s cours e w h o h a ve h ad so me preli mi n a ry w or k

,

in ge n er a l b iology b ut w h o h a ve f orgotte n mos t of w h a t

,

l ittle of th e e le me n ts of ph ys iology th ey m ay h a ve le a r n ed
in th e school s . A ce rt a i n a mou n t of p hys iology of a more
o r les s ge n e r a l n a tu re h a s accordi n gly b ee n i n trodu ced in
add itio n to th e d escri ptio n s of th e s peci a l f u n ctio n s of th e

v a riou s org an s o f th e body . T he book i s more s u it ab le

f o r us e as a text in college or u n i vers ity cl a s s es th a n in
h ig h sch ools a lthough i t i s h ope d th a t i t wi ll p rove of
,

s e rvice to te ach e rs in high sch ools w h e r e th e f rog i s

s tud ie d in th e cours e in z o o l ogy .

C o n s ider able s p a ce h as b ee n d evoted to describ i n g the

h abits an d n atu r a l h i story o f th e f rog an d th e e n de avo r
,

h a s bee n m ade througho ut th e w ork to correl ate t h e study

of stru cture w ith th a t of th e phys iologi c al fu n ctio n s of the

body a n d th e a cti viti e s of th e o rg an i s m as a whole in rel a
t io n to th e e n v i ro n me n t. Th ere i s I beli e ve a v a lue in
, ,
vi P R E F A CE

getti n g a f ai rly a de qu ate ide a of the whole life of an y o n e

org a n i s m w h ic h i s n o t a tt a i n e d by th e us u a l co urs e of a
s tudy of ty pe s .

I n p rep a ri n g th i s w ork i t h a s co s t m uch del ibe r a tio n in

m an y c a ses to decide w h a t m ateri a l to i n clude a n d wh a t
to rej ect an d prob ab ly s e ve r a l th i n gs h a ve bee n o mitted
,

w h ich it m igh t h a ve b ee n de s ir a ble to h a ve ret a i n ed Th e .

li ter atu re de a l i n g w ith th e f r og i s almo s t a pp alli n g in i ts

exte n t . P e rh a ps n o an i m al excep t m an h as bee n the
, ,

s ubj ect of s o m an y s cie n ti fic i n ve s tig atio n s O n e s eldo m .

picks u p a volume of a p hys iologic a l j ou r n al witho ut fi n d

ing th a t th e f rog com e s in f o r a sh a re of a tte n tio n in o n e
o r more a rticle s I t i n deed seem s as i s ofte n rem a r ked
.
, ,

th a t the f rog i s e sp ec i ally de s ig n ed a s a s ubj ect fo r b io

logic a l r es e a rch I n f act mos t of wh a t i s k n ow n in ce rt a i n
.
,

de p a rtme n ts o f phys iolo gy i s d e rive d f ro m a s tudy of th i s

an im a l .

T h e a n atomy of th e f rog h as b ee n mos t exh au s ti ve ly

tre ated in G a u pp s e xcelle n t rev i sio n of E cke r a n d W ie
’

“
de rsch e im s A n a tom ie d es F ro s ch es
’
a n d I a m n atu r a lly
,

u n d e r gre at ob lig atio n s to th i s work M os t of th e s t ate

.

me n ts m ad e in th e p re s e n t boo k reg ardi n g th e a n ato my

of th e frog h o w eve r I h a ve veri fied by p erso n a l ob s erv a
, ,

ti o n M uch of th e m ateri a l de ali n g w ith th e p h y siology

.

an d n a tu r a l h i s tory of th e f rog is h e re b rough t togeth e r

f o r th e fi rs t ti me W h e re n o t oth e rwis e me n tio n ed th e

.
,

st ate me n ts in th i s book ar e m ade w ith refere n ce to th e

commo n leo p ard f rog of North A me ri c a Ra n a p zp z em
' '

, ,

exce p t in c a se s where th ey m ay b e u n de r stood to apply

equ ally w ell to any s peci e s of th e ge n u s .
 P R E FA CE

I am i n debted to P r ofes sor J acob Reigh ard f o r s e ver a l

s ugges tio n s an d criti cis m s in co n n ectio n w i th th e prep ar a
tio n of th is work ; to D r W P L o mb ard fo r re ad i n g an d
. . .

cri ti cis i n g se ver a l of th e ch a pters de a li n g with th e ph ys i

olog y of th e f rog ; an d to D r J E Du e rde n f o r ge n erou s ly
. . .

u n dert a ki n g th e l abo r of cri t ic a lly r e adi n g t he w hole o f

th e m an u scri pt b efore i t w e n t to pre s s .To D r E r n st.

G a u p p a n d h i s publi s he r F ri ed rich V ieweg I w i s h to

, ,

exp r es s m y th a n ks for th ei r perm i s s io n to rep r od uce s ev

e r a l of th e figu r es in G au pp s revis io n of E c k e r a n d
’

’ “
W ieders ch eim s A n ato m i e des F r osch e s .
”
 P R E F A CE TO THE S E CO ND
E DI T I O N

I N th e prese n t issu e o f this work lit t le ch an ge h as bee n

m ad e except t o correct a few errors which in spi t e o f c are
,

ful revi sio n h ad crep t i n to th e previous o n e I hope t h at

, .

n o w wi t h t h e aid of my cri tics wh ose servic e s a r e gr at efully

,

ack n owledged the book h as be e n re n dered free fro m an y

,

serious mist akes .

 CO NT E NT S

T H E A M P H I I A N GE N E RA L A N D F R IN PA R T I C U LA R
B I OGs

T HE H A I T S A N D N AT U R A L H IS T R Y
B O OF FR G
THE O

E XT E RN A L CH A R A C T E R S O F T H E F R UG

P R E L I M IN A R Y A CC O U N T O F T HE I N T E RN A L S T R U C T U R E
T H E DEV E L O P ME N T O F T HE F R G O

HIS T O L O G Y O F T H E F R OG
T HE D I G E S T I V E S Y S T EM A N DITS F U N CT I O NS
T H E V O CA L A N D RE S PI R AT O R Y O R GA NS
T HE S K I N
T H E E XC R ET O R Y S Y S T EM
T H E REPR D U CT I V E O R GA NS A N D
O TH E F AT B O D I E S
I N T E RN A L SEC R E T I O N A N D T H E D U CT L E SS G L A N DS
T H E S K E L ETO N
T H E M U S C UL A R S Y S T EM
T HE CI R C U L AT R Y S Y S T EM
O

T H E NE R V O U S S Y ST EM
T H E SE N S E O R GA NS
I N S T IN CTS A N D TR O PI S M S As R E L AT E D T O RE F L E X
A CT I O N
X I X T H I N T E LL I G E N CE O F T HE F RO G
. E

IND E X
 TH E B IOLOGY OF TH E F RO G

C H A PT E R I

TH E A M P H IBI A I N G E N E R A L A ND FRO G S I N P A RT I CUL A R

TH E frog is a represe n t ative of th e cl ass o f vertebr ate

an im als k n ow n as the B at rachia or Amph ib ia This group
, .

is i n t erm edi at e in m an y fe at ures of structure an d h abit b e

t wee n t h e fishes an d the reptiles an d prese n ts th e followi n g

ge neral ch aracters : The ski n is u su ally Smooth and moist ,

an d in all th e rec e n t Amph i b i a wi t h t h e exceptio n o f a few

, ,

of the Apod a e n tirely d evoid of sc ales Th e limbs u nlike

,
.
,

those of fishes are typic ally of th e fi ve toed or pe n t ad actyle

,
-

type The skull articul ates wi t h th e first v ertebr a by two

.

o ccipi t al co n dyles an d is u su ally com posed o f few bo n es .

L u n gs are prese n t except in r are i n st an ces as in cert ai n ,

lu ngless s al am and ers where th ey h ave b ee n seco n d arily lost

,
.

The he art h as three ch a mb ers a v e n tric le an d t w o auricles

,
.

Th e aor t ic arches are symm etric al o n the two sides There .

i s n o a m n io n n o r all an toi s al t hough there i s a hom ologue

,

of t h e l at t er in t he uri n ary bl add er M os t o f t h e species

.

u ndergo a m e t a mo rph osis t h e you n g livi ng in th e w ater an d

,

bre athi n g by m e an s o f gills Th e l at ter di s app e a r as th e

.

lu n gs beco me fu n ctio n al except in some o f th e lower A m

,

p hib ia in which they are r et ai n ed t hroughout life

,
.

The me mb ers of t hi s cl ass as a rule are aqu atic o r sem i

aqu atic in h abit a circu m st an c e w hi c h s u ggested th e n am e
,

of th e cl ass E ven th e most t erres t ri al of th e species ge n er

.

B I
 TH E BIOLOGY OF TH E FROG CH A P .

a lly ke ep in a d amp h abit at V e ry few of th e m c an e n dure

.

an y co n sid er a ble a mou n t o f drough t E ven wh e n they live.

far away fro m w at er duri n g m o st of th e ye ar th ese an i m als ,

n e arly al w ays go to the w a te r in the spri n g to deposi t their

eggs .

Amphibi an s are co nfi n ed to th e torrid an d temper ate

zo n es . I n th e temper ate zo n e th ey hibern at e duri n g the
wi n ter cr awli ng i n to th e mu d or o t her sh el t ered loc aliti es
,

out of t h e r e ach o f fros t I n higher l at i tudes where o n

.
,

a ccou n t of the ex t re m e c o ld the grou n d beco mes froze n

in th e w i n ter to a d e p th o f sever al fe et th e Amphibi a are ,

abse n t si n c e th ey could n o t e asily get i n to situ atio n s wh ere

,

t hey could esc ap e bei n g fro z e n .

The Amphibi a m ay be divide d pri m arily i n to th e S t ego

cep h ali or L ab y r in t h odont s an d t h e L issamph ib ia
, ,
T he .

fo rmer group co n sists exclusively o f fossil forms w hich firs t

a ppe ar in the C a rbo n iferous era a n d ex t e n d i n to the U pper

Tri assic The bodies of the S t ego c ep h ali were usu ally
.

covered with sc ales an d bo n y pl ates and th e S kull w as ,

provided with n u merous d erm al bo n es S om e o f th ese .

exti n ct ani m als were of very l arge size an d in m an y speci es ,

th e de nt i n e of th e t e eth w as fo l ded in a very co mplic at ed

m an n er — a fe atu re which c au sed t he n am e L abyri n thodo n t
,

to be give n t o t h e group .

The L issamph ibia l ack t h e exte n sive d erm al S k eleto n of

th eir ex t i n ct pred ecessors Th eir to o t h s truc t ure i s S impl e
.

a n d they n ever a t t ai n l arge proportio n s T h ey are com .

m o n ly divid ed i n t o three ord ers which m ay be determi n ed

by t h e followi n g key

Le gs bs t
a en

Le gs p s t
re en .

T il d m p h ibi s
a e a an

T ill ss m p h i bi s
a e a an Anm '
a .
 TH E A MP H IBI A IN G E N E RA L 3

TH E AP ODA

The Ap oda or Caecilian s are cre atures o f wormlike form

, , ,

e n tirely d evoi d of limbs an d li mb girdles Th e ski n is .

s mooth an d throw n i n to tr an sverse ri ngs I n some forms .

s m all sc ales are fou n d e mbedded in t h e i ntegume n t Th e .

eyes are sm all o r abse n t The Apod a are ge n erally fou nd .

in mois t grou n d in w hich th ey burrow an d they are co n fi n ed

, ,

to tropic al or subtropic al regio n s N0 sp eci es occu r in Nor t h .

Am eric a n or t h o f M exico .

TH E URODE L A , OR TAI L E D AM PH I B I ANS

The t ailed a mph ibi an s occupy a more pri mitive posi t io n

th an t h e t ailless forms A l a rge p ropor t i o n o f the m live in
.

t h e w a te r an d t h e lo w e r m em b ers of th e group re t ai n t h eir

,

gills in the adult s t ate The U rodel a are divided by G adow .

i n to four families as follo w s

] aw s w i t h u t t t h N hi d limbs
o ee . o n S ir em a ce
’
’
.

Bo th j aw s i th t t h F
w d h i d limbs
ee . o re an n

p s t
re e n .

G ills p sis t t
er N y lids m xill y b
en . s o e e or a ar one

G ills usu lly bs t i t h

a adul t M xi ll y
en n e a . a ar

b sp s t
o ne re e n .

Ey s e i t h lids
w

Ey s d
e i d f lids
evo o

The P r ot eidae co n sti tu te th e m ost pri mitive of the U ro

del es A t the side s o f th e n eck th ere are three p airs of
.

ex t er n al gills The spe cies are aqu atic in h abi t There are
. .

o nly thre e ge n er a two o f which Nectur us an d Typhlomolge

, , ,

are co n fi n ed to N o r t h A meri c a Th e re m ai n i ng ge n u s Pr o .
,

t eu s rep rese n ted by a si n gle speci es P a ng u z m z s is fou n d

’

.
, , ,

o n ly in th e c aves of Au s t ri a This speci es occurs in deep .

cool w at er in r egio n s of comple t e dark n ess I t s eyes like .

,
4 TH E BIOLOGY OF TH E FROG CH A P .

those of m an y c ave an im als are r udime n t ary I t s color is ,

.

n e arly whi t e but i f exposed t o ligh t its ski n gr a du ally tur n s

,

d ark an d eve n t u ally m ay becom e n e arly bl ac k .

A c ave s al am an der Yj p /z lo m olg e r a t/t ea m, closely alli ed

,
/

to P roteus w as fou n d o n ly a few ye ars ago in Tex as w here

, ,

i t w as discovered in w ater throw n u p fro m an artesi an w ell .

Th e body of thi s species is S le n der an d provided with a lo n g ,

fl at t e n ed t ail The legs are lo n g an d S le nd er Th e eyes

. .
,

li ke those of P ro t eus are r udim e n t ary an d buried be n e at h

,

the ski n Th e most com mo n rep r ese n t ativ e of the P roteid ae

.

I .
—P r ot eu 5 a ng u m u s . F ro n t i
v ew fo th em uth i th
o n e u pp er

c o rn er .
( A tter G a d ow , Ca mb idg
r e N t u l H is y )
a ra to r .

are th e
“ mud pu ppies wh ich belo n g or w at er dogs ,

t o t h e ge n us Nec t uru s m a m /05 m i s th e most

.

abu n d an t speci es I t occurs in t h e n orth er n an d e aster n

.

p ar t of the U n i t ed S t ates w est of th e Allegh an ies an d is , ,

especi ally abu n d an t in th e regi o n o f the G re a t L akes I t s .

ge n er al color is brow n above m arked with d arker spots an d , ,

a dir t y whi t e or dusky color below I t h as bushy red gill s .

,

w hich ar e kep t movi n g b a ck a n d for t h at freque n t i n t erv als .

L ike most amphibi an s it i s mos t ac t ive at n ight duri ng th e

,

day i t li es co n ce aled o u t of t h e re ac h o f light .

The fa mily S ir enidae is represe n ted by two ge n era S ir en ,

an d P seu dob r an ch u s bo t h o f w hich are co n fi n ed to N

,
o r th
 TH E A MPH I BI A I N G E N E R AL 5

Americ a E ach ge n us co n t ai n s but a si n gl e species Th e

. .

l arge r o f these S ir en la cer lz n a is fou n d in the rivers and

'

, ,

po n ds o f the S outh er n S t ates fro m Tex as to Nor t h C aro ,

li na Th e body is lo n g an d s n akelike in appe aran ce The

. .

fore legs are ve ry short an d situ ate d close behi n d the exter

FIG . 2. S z r en la cer t m a .
(F m ro th e Ca mb idg r e Nat u ra l H ist y ) or .

n al gills ; th e feet are fou r t oed The r e are three p ai r s o f

-

gill sli t s The ge n us P seudobr an chus has o n ly o n e p ai r

.

o f gill sli t s i n s t e ad of thre e and the feet poss ess but th r e e

,

t o es Th e si ngle species P s ir z a tw occu r s in G eo r gi a and

'

.
,
.
,

F lorid a .

The Amphiu midae i n clud e forms of quite divers e ap pe ar

an ce which are so me t i m es pl a ced in di s t i
,
n ct fa milies Th e .

ge n us Amphiu m a i s represe n t ed by a si ngle sp ecies A mea ns ,

.
,

fo un d in th e S outher n S t ates of Nor t h Americ a T he body .

is eel like w ith t he very s mall legs si tu ated far ap art n e ar

-

, ,

t h e two ex t re mi t i es T h ere is a si n gle p a ir o f gill sli t s

.

behi n d t h e he ad n e ar the fore legs Th e le ngth of thi s

,
.

speci es is o ft e n over two feet Th e fe m al e l ays her eggs in .

the l at t er p art o f th e su m mer an d li es coiled about the m in ,

som e pro t ected s p o t u n t il t hey h atch ,

.

The ge n er a Cr ypt ob ran chu s an d Megalob at r achu s are

some times pl aced in a dis t i n ct family the Crypt ob r an chl ,

dw T h e forme r is r eprese n ted by the l arge h ellbe n der ”

.
,
6 TH E B I OLOGY OF TH E FRO G C H AP .

C . e aster n U n i t ed S t ates This speci es

of the .

m ay re ach a le n g t h o f twe n ty i n ch es I t s body an d h e ad .

a re mu ch fl at t e n ed a n d th e sid es ar e bordered wi t h curious

,

fluted folds of ski n Th e eyes are r el at ively very s m all

. .

Th e hellbe nd er i s very sluggish in its h abi t s bu t i t is , ,

n evertheless a very vor a ciou s e a ter

,
I t s vi t ali t y j udgi n g .
,

fro m an accou n t by M r F re ar is c ert ai nly rem ark able

1
.
, .

FI G .
3 . A mp /21 2m m m ean s .
( Fro m t he Ca mb idg r e Na t u ra l H ist ory) .

M r F re ar
. tells of o n e sp eci me n which h ad bee n picked up
a ft er it
“h ad l ai n exposed to a su m mer su n for for t y eight -

hours I t w as t h e n brought i n to t h e museu m an d left a

.
”

day before it w as pl a ced in alcohol Aft er it h ad be e n left .

in t h e alcohol “ for at le ast t w e n ty hours it w as t ake n ou t ”

,

“w he n it beg an to O e n it s big m ou th vigorously sw ay it s

p ,

1 Am . Na t , V ol
. 1 6, 1 882 .
 TH E A M P H IB I A IN G E N E RA L 7

t ail to an d fro an d give o th er u n doubted sig n s of vit ality

,
.

The gi an t s al am an der of J a p an M eg a /obn t r a enn s m a xim ns , ,

is closely rel ated to th e prec edi n g sp eci es Th e l argest .

spe cime n s k n o w n exceed five feet in le ng t h .

The S alam an dridae for m a l arge family which is fre ,

-

qu e n t ly divid ed i n to several di ffere n t famili es by m an y

writers O nly a few of the mor e n oteworthy forms there
.
,

fore will b e described

,
.

The group is divide d by G adow i n to fou r subfamilies as

follows
A . S er i s
e ht s s
of p a al
ptsal
t i l y c
t e et ran ve r e o r o er o r o n verg n g . i
B P sp h i d i t h ut t t h V t b
. a ra en o mw o ee er e ra
e a

—
.

p h i l T s 4 5 c oe o u s . oe A m é b/slom a fln w .

BB P s p h id i t h t t h
. ar a en o w ee .

C V t b .
p is t h c l us
er e T s 5 T gu
rae o o oe o . oe . on e

l g ly f ar e re e D esm og n a t /z z n w
'

CC V t b . m p h ic l us T gu sm l l
er e rae a oe o . on e a

d l g ly f an ar e re e P letnoa o n t in w
’
.

AA S i s. f p l t l t th i t
er e o l gi t udi l
a a a ee n w o on na

s i s di gi g b h i d P s p h i d
er e ver n e n . ar a eno

t h l ss
'
’
t oo e S a la m a n ar z n ce .

Th e subfamily Amb lyst omat in ae is represe n ted in t his

cou n try m ai n ly b y th e t w o ge n er a Ambly st oma and Chon
dr ot us Amblysto ma co n t ai ns qui t e a l arge n u mber of
.

species They are mostly of co n sid erabl e size and fre

.

qu e n tly spotted in color They are very retiri n g in th eir .

h abits an d are n o t oft e n see n excep t in t h e spri n g w h e n

, ,

they go to th e w ater t o breed Th eir eggs are usu ally .

a tt ached to t w igs or ste ms of gr ass an d are fo u n d in ,

rou n ded o r irregul ar m asses E ach egg i s surrou n d ed .

by a very thick co at o f j elly A mblystom as are amo n g th e .

very first amphibi an s t o lay th eir eggs E ggs of a species .

o f A mblystom a ( prob ably lzgmn n m) h ave bee n t ake n n ear

'
8 TH E B I O L O G Y OF TH E F RO G CHAP .

Ann Arbor M ich ig an at th e followi ng d at es : M arch 1 5

, , ,

1 8 9 2 ; M arch 2 6 1 8 9 5 ; M a rch 2 9 ,
18
9 ;6 M arch 1 3 1 8 97 ; , ,

M arch 2 8 1 9 0 5 ,
.

T h e l arv ae o f A izgmn n m w ere for m erly co n sidered a

'

sep ar ate species the axolotl which w as pl ac ed in a disti n ct

, ,

ge nus S iredo n am o n g the p ere nn ibr an chi ate u rodeles

, ,
.

U n der cert a i n co n ditio n s th e exter n al gills o f this l arv a

m ay be re t ai n ed u n t il aft er th e bre edi n g se aso n an d t his ,

peculi arity led to i ts bei n g m ist ake n for a n orm al adult

form I t h as bee n co n te n ded t h a t th e m e t am o rphosi s o f t h e
.

axolo tls could be a cceler at ed if t hey w ere forced t o bre a t h e

air bu t P rofessor P o w ers h as rece n tly show n th a t th e facto r

,

of n u t ri t i o n is prob ably t h e m os t i mport an t o n e al t hough ,

others are i n flu e n t i al in produ ci n g thi s ch an ge si n ce i t

, ,

usu ally fo llo w s in su fficie n t ly m ature l arv ae u po n a sudd e n

dimi n utio n o f th e food supply .

The Desmognat hinae i n clud e t hree ge n er a of whic h ,

Desm o gnat h u s is the mo st co mm o n I t co n t ai n s o n ly t hre e .

speci es all o f which ,

ar e co n fi n ed to t h e

e as t er n p ar t o f the
U ni t ed S t ates T he .

speci es live c o n
c ealed in th e day

time u n d er sto n es
or in sh el t e r ed n ooks
w here t he air is
moi s t The fem ale
F I G 4 —D m g t/ f
.

e5 o na F m l i t h O f D u ser/5 1a S her
zus u s ca s e a e w
f
. . .

gg m ss ( A ft W i ld )
e -
a . er
y er .

eggs in t w o lo n g
stri n gs which sh e wr aps arou n d he r body a ft er h avi n g resor t ed
t o a sui t abl e h idi n g p l a ce A n o t her represe n t a tive of thi s .

subfa mily i s i lz lo z mfo n gee/e ms a bli n d S p eci es fou n d in a

'
‘
.
,

c ave in M issouri .
10 TH E BIOLO G Y OF TH E FROG CH A P .

th e gills is doubtless depe n de n t o n th e fact th at th ey ar e

worked in for t h e purpose o f absorbi n g food .

The ge n us Tr it on is re m ark able o n a ccou n t o f th e m arked

s exu al d imorphis m wh ich occurs in several of the speci es ,

esp eci ally duri ng th e breedi n g se aso n The m ale of T cr is . .

at this ti me possess es a high ser r ated c r est above the

FI G .
5 . Tr it on cr ist a t u sfm l . 1 e a e ; 2, m la e as he a pp e a r s du i g
r n th e
b di g s s
re e n ea o n .
( A ft e r Ga d ow .
)

he ad an d body an d is m arked wi th co n spicuous colors

, .

Aft er th e breedi n g se aso n the dors al crest beco mes gre at ly

reduced an d th e color atio n beco mes m o re dull The fem al e .

h as n o crest an d is n o t so co n spicuously colored as the

—
m al e al t hough she also becom es duller in color after th e
,

breedi n g period is p ast .

A close rel a tive o f th e Tri to n s i s the co mmo n n e w t

( D z emy ezj ln s of the or h er e ster p rts
’

i
n t n a n d a n a

of this co u n try I t i s a pretty speci es bei n g colored a n

.
,

olive gree n reddish or red dish bro w n above or an ge or

-

, ,

l emo n yellow belo w an d h avi n g a l ater al ro w o f sc arlet

-

spo t s e ach surro u n ded by a bl ack r i n g A v ari e ty m z n z a

' '

.
, ,
 TH E A M P H IBI A IN G E N E RA L I I

in s, which h as bee n described a n d w hich is ch ar ac t erized

by possessi n g a vermilio n r e d color is s aid by G age to be ,

o n ly an i mm at ure form o f this species E gg l ayi n g w as .

fou n d by j o r d an to t ake pl ace n e ar Worce ster M assac hu ,

setts fro m about April 1 0 to Ju n e Th e eggs are l aid in

,
.

sm all n ests at tached to m asses of vege t atio n or wr appe d ,

withi n le aves of aqu atic pl an ts .

TH E ANURA

The Anu ra or t ailless Amphibia h ave a sh ort b r o ad

, , ,

body w ith well developed hi n d legs fi t ted for j u mpi ng

,
-

They are divide d by G adow as follows

A T n gu bs nt
. o e a e Ag lossa .

AA T n gu p s nt ( Pn n
. o e re e a er o
g lossa ) .

B . H a vel s
sh ul d gi dl
of l pp i g
th e o er r e o v er a n

i t h middl li n e e ne

C S c l di p p hys s dil t d
. a ra a o e a e .

D T mi l p h l g s t cl sh p d
. er na a an e no aw a e .

Ribs p s t U pp j ith t t hre e n . er aw w ee

N ibs U pp j ith t t h '

o r . er aw w ee l é z
’
P e o a i ace .

N ibs B t h j s i t h u t t t h
o r .
‘

o aw w o ee

DD T mi l p h l g s cl sh ap d usu lly
. er na a an e aw e ,
a

su p p t i g d h si dis k s or n a e ve

CC S c l di p p hys s cyl i d ic l
. a ra a o e n r a

B B H l s f t h sh uld
. a ve gi dl m t i g i
o e o er r e ee n n

th m iddl li d f mi ge m e ne an or n a e

di b an ar

C S c l di p p hys s dil t d E gy siom a t z da

'

. a ra a o e a e n .

CC S c l di p p hys s cy li d ic l
. a ra a o e n r a R a n ia ae
’
.

Th e Aglossa i n clud e o n ly a fe w aberr an t forms ch ar acter

.

iz e d by t h e abse n ce of a t o n gue an d t h e fact th at th e

E ust achi an t ubes ope n by a si n gle medi an ap er t ure in th e
posterior side of th e p al ate The most n o t e w or t hy me mbe r .
12 TH E BIOLO G Y OF TH E F RO G CH AP .

of this grou p is th e p eculi ar S uri n am to ad P ip a a memea n a

’

, ,

from th e n orth er n p art of S outh Americ a Thi s cre ature .

h as a most grotesqu e appe ar an ce Th e b ack i s bro ad an d

.

fl at te n ed the he ad sm all tri an gul ar depressed an d fur

, , , ,

n ish ed with i rregul ar fl aps n e ar the lips ; the eyes are sm all

an d h ave a rou n d pupil The m ost re m ark able fe ature of

.

the sp ecies is th e mode in which the fe m ale c arri es th e eggs

a n d you n g Aft er the eggs are l ai d an d fertilize d th ey b e
.
,

co m e push ed u po n the b ack o f th e fem ale t o w hich th ey ,

adh ere Th e ski n t he n grows u p a rou n d the eggs i nclosi ng

.
,

them in sep ar a te c avi ties which becom e e n t irely covered

over The t adpol e st age is p assed wi thi n th ese c avities
. .

Whe n th e you n g P ip a is quit e fully fo r med i t bre aks out an d ,

m akes its esc ape .

The D iscoglossidae are n o t represe n ted by an y Americ an

speci es O n e of t h e m ost n ote w or t hy of th e E urope an
.

speci es o f t his family is th e so c alled obste t ric al to ad A ly fes

-

I n th e bre edi n g s e aso n th e m al e cl asps t h e

’

o é s iet r z ea ns .

fem ale in the usu al w ay an d wh e n t h e egg stri n gs are ex

,

t ru ded h e t an gles the m arou n d his hi n d legs an d c arries

,

th e m abou t w i th hi m Whe n the you ng l arv ae are about

.

re ady to esc ape t h e m al e t akes to the w ater an d fre es hi m

,

self o f t h e m ass .

The P elob at idae i n clude bu t two Americ an ge n er a ,

S caph iopus an d S p ea These forms are com mo nly k no w n

.

as th e sp ade foot frogs o n a c c o u n t o f the peculi ar hor n y

-

a ppe n d age which occurs o n th e i nn er si d e o f th e hi n d foot .

This struc t ure is employed in diggi ng in the grou n d where ,

t h e an im al is co n ce al ed duri n g th e day
'

S eap /z z op a s /20/
.

ar o oki wh ich is fou n d in the sou th er n an d e as t er n p arts of

,

t h e U n ited S t at es is very c a priciou s in m aki n g i ts appe ar

,

a n ce . Aft er r ai n s in th e sp r i n g or summ er t h e sp ad e foot -

frogs co me ou t in gre a t n u mb ers an d lay th eir eggs m aki ng ,

 TH E A M P H IBI A IN G E N E R AL 13

a gre at cl a mor with their so n g The n they dis appe ar an d .

,

m ay n o t ag ai n sho w th e mselve s for sever al ye ars 1

.

Th e B u f onidte or to ads co mprise a l arge fam ily which

, ,

is fou n d o n all the co n ti n e n ts of th e globe Th e pri n cip al .

ge n us is B ufo which i n cludes the best k n ow n represe n t a

,
-

t ives o f the family Th e to ads of this ge n us possess a very

.

rough w arty ski n w h ose irregul ariti es are c aused by the

, ,

l arge n u mber o f poiso n gl an d s co n t ai n ed in it Th ese .

gl a nds secrete a whi tish milky flui d of a v ery poi so n ous ,

n a ture E ve n a very sm all qu an tity of this subst an ce w he n

.

i nj ected i n to t h e blood o f a sm all an i m al soo n produces fatal

effects The abu n d an ce o f thi s secretio n affords th e to ad
.

very e ffi cie n t pro t ectio n an d n o t m an y an i m als h ave th e

,

h ardihoo d to att ack the cre atu re I n additio n to th e poiso n .

the ski n secre t es mucus as in o t he r a mphibi an s although , ,

n o t in gre at qu an tity .

Th e color of to ads like th at of frogs m ay ch an ge u n der

, ,

th e i n flue n ce o f differe n t exter n al co n ditio n s Wh e n ex .

posed in a light colore d e nviron me n t th e s k i n usu ally

-

becom es lighter in color I n a d ark e n viro n m e n t it b e .

comes d arker t hu s bri n gi n g a bout a cert ai n ad apt at io n of the

,

colo r o f th e an i m al to th at of its surrou n di ngs This ch ange .

i s effec ted by m e an s o f ch an ges in th e p igme n t c ells of the

ski n in th e s a m e m an n e r as in th e frog which will be more ,

fully described l ater .

To ads are n octurn al in h abi t D uri n g the day th ey lie .

co n ce aled u n d er s to n es or in o ther d amp sh ady loc ali t i es , ,

ve n turi n g out o n ly tow ard eve n i n g Th ey hop about like .

frogs al though wi th much less agili t y O n th e oth er h an d

,
.
,

t h ey climb with co n sider able re adi n ess Th ey fee d upo n .

e ar t hworms s n ails an d all sorts of i n sects Th e l a tter ar e

, ,
.

ge n er ally c augh t by sudde nly t hrowi n g out th e to n gu e and

1 S ee A bb o tt, Am . Na t ,
V ol . 1 6, an d H ar gi t t, Am . Na t ,
V ol . 22 .
14 TH E BIOL OGY OF TH E FROG CH A P .

th e n with dr awi n g it alo n g with the i n sec t to which i t adheres .

A ngle w orms are seiz e d by the j aws an d stuffed i n to the

mou th by th e fore legs To ads are very useful in destroyi ng
.

l arge n u mbers of i nj u r ious i n sects an d he n c e d eserv e all ,

possibl e e n cour age me n t an d pro t ectio n K irkl an d an d

1
.

G arm an who h ave c are fully ex ami n ed the co n t e n ts of t he

,
2

s tom achs o f a l arge n u mber o f to ads fi n d th at the v ari e ty of ,

i n sects devoured is very gre at A n ts were the fo r ms mos t .

commo nly m et wi th in th e stom ac hs an d be e tles bu gs , , ,

moths an d c aterpill ars were fou n d by G arm an to follo w

,

successively in ord e r of frequ e n cy .

To ads keep withi n a c ertai n loc ality for a lo n g pe r iod .

They h ave their p articul ar holes or n ooks wh ere they li e ,

duri ng the day an d to which they r e t ur n a ft er t heir n i ght s ’

j our n ey in se arch of food Their s e n se of loc ali t y is app ar

.

e n t ly quite good as is sho w n n o t o n ly by t h e fact th at t h ey

,

fi nd their w ay ho me bu t by thei r h abit ually visiti n g cert a i n

,

spots in the cou rse of th eir n octur nal w ande r i ngs Th e .

lo ngevity of to ads i s som ewh at u n cert ai n B o u lenger kep t .

o n e speci me n for twelve ye ars Th ere is a record of a .

speci me n which lived to be thirty six ye ars old an d w as th e n -

a ccide n t ally killed C ases are recorded in which a to ad

.

h as occupied a cert ai n retre at for a lo nger p eriod bu t th e

ide n tity of the i n dividu al i s n o t assu r ed Th er e are n u mer .

ou s sto r i es of live to ads fou n d embedde d in rocks or se aled

up in trees ; bu t wh ile m an y o f t h e m see m to be quite well
authe n tic ated t h ey do n o t give evide n c e of su ffici e n tly c a re
,

ful i n vestig atio n to com pel belie f B uckl an d h as show n

3
.

th at t o ads m ay live withou t food wh e n se aled u p in blocks ,

of lim esto n e for over a ye ar to ads i mp r iso n ed in the lime

,

1 K i kl d B ull N 46 M ss A g ic E p
r an ,
. o .
,
a . r . x . S ta .

2 Gar m B ull N 9 K y E p S
an , . O . 1 , . x . ta .

3 B uckl d Cu i s i i s f N u l H is y
an ,
r o t e o at ra to r .
 TH E A M P H IBI A IN G E N E RA L 1 5

sto n e for two ye ars were i n v ari ably fou n d to be de ad We .

should be skep t ic al th erefore abou t accep t i n g stori es abou t

, ,

t o ads h avi n g bee n fou n d a live in situ at io n s where they mus t

h ave rem ai n e d for a much lo nger t ime .

To ads hibern ate u n der rocks or in c avities in the grou n d , ,

whe r e they are protecte d fro m extre me cold O ft e n sever al .

to ads m ay be fou n d huddled together I n o n e hidi n g pl ace .

H ere they lie b e n u mbed an d al most stiff although n o t actu ally ,

froze n u n til spri ng S oo n after their e merge n ce from their

,
.

wi n ter sle ep th ey u su ally bet ake the mselves to w ater to

d eposi t th eir eggs The bree di n g p eriod of a o [en ngz n os ns
°

in M ass achusetts a ccordi n g to K irkl an d is in April

, in ,

I t h ac a New Y ork ( G age ) fro m t h e middle of April t o M ay

,
1
,

in A n n Arbor M ichig an I h ave fou n d this specie s breedi n g

, ,

in the l atter p art of April Th e eggs are embedded in lo n g

.

stri ngs of j elly which are usu ally fou n d amo n g vegetatio n
n e ar the shore The m ales of B len tzgz n osu s are much
'

. .

sm alle r th an the fem ales D uri ng th e breedi n g s e aso n they

.

frequ e n tly u tter a p eculi ar shrill sou n d After this p eriod .

,

a ccordi n g to Alle n th e so n g ch an ges to

,
a shorter lower ,

to n ed n ote th at at n igh t has a p eculi ar weird n ess an d

, , ,

re aches al most a w ail This n ote i s he ard mostly at eve ni n g

.

an d duri n g the n i gh t though I h ave occ asio n ally he ar d i t

,

e arly in the m orn i n g an d l ate in th e after noo n .

”

Wh e n to ads are h andled an d ofte n eve n wh e n app r o ached, ,

they swell t heir bodi es with air S lo n aker tells o f a to ad

2
.

which wh e n appro ached by a s nake would swell u p an d

orie n t i ts elf with its b a ck toward the e n emy The i n fl ati o n .

of the body m akes it m ore d ifficult to ret ai n hold of t h e

cre a ture as an y o n e m ay re adily d etermi n e
, .

There ar e seve r al sp ecies of to ads in North Am e r ic a but ,

1 Ga g e, P r oc A m A ss A av S ci , V o l 47 ,
. . .
’
. . . 1 8 98 .

2 P r oc. ’
[ n a z an a A c S a l , 1 900 . .
16 TH E BIOLO G Y OF TH E F RO G CH A P .

most of the m are co n fi n ed to th e Wester n S t a tes Abou t th e .

o n ly speci es whi ch o ccurs e as t o f t h e M i ssissippi wi th th e ,

exceptio n of a o gn er ez ens which r an ges fro m North C aro

’

li n a t o F lorid a is B Zen tzgz n os n s which is wi dely distributed

'

,
.
,

a n d qui t e a bu n d an t .

The H ylidae o r tree frogs form an exte n sive an d widely

, ,

distributed fa mily Th e ti ps o f th e to es are fur n ishe d with

.

sm all adh esive disks w hich e n able the an i m al to cli mb up

the tru n ks of trees M an y sp eci es are able to cli mb u p a
.

vertic al surface o f smooth gl ass This i s re n dered possible .

n o t so much t hrough th e suctio n o f th e di sks as by a s t icky

secretio n which is p r od u ced by th e gl an ds of th e ski n at

these poi n ts .

M ale tree frogs are u su ally able to m ake a n oise which i s

a sto n ishi ngly lou d for cre a tures o f so sm all a size I n H yl a .

an d its alli es th e voc al s ac of the m ale is c ap a ble o f gre at

diste n sio n an d whe n fully i n fl ated beco m es much l arge r

, ,

th an th e h e ad The voice is he ard most o fte n in the b r eed

.

in g se aso n but it m ay also be h e ard duri n g most of th e

,

s u mmer especi ally abou t dusk The n ote o f the tree frog
,
.

i s ofte n reg arded as i n dic a tive of appro achi n g r ai n It .

is h e ard freque n tly i m med i ately before a shower The .

i n te gu me n t of th e cre ature i s e asily affected by ch an ges

in moisture S itu atio n s wh ere the air i s d am p ar e alw ays
.

preferred an d i t i s n o t u n natur al th at th ei r so n g sho u ld

,

be h e ard whe n the atmosp here app r o aches the poi n t o f

s a tur atio n .

A gre at m an y species o f tre e fro gs h ave a re m ark abl e

power of ch an gi n g th eir color u n der differe n t exter n al c o n
dit io n s . Wh e n amo n g gre e n l e aves their color is u su ally
gree n bu t wh e n o n the b ark of trees o r o n the grou n d thei r
,

color m ay ch an ge to a br o w n or gra y .

The North A meric an species of this family n o r th of

1 8 TH E BIOLOGY OF TH E FRO G CH A P .

an d where th ey u n dergo develop me n t as far as th e t adpole

st age .

Th e E ngyst omat idae co n t ai n bu t o n e North Americ an

sp eci es E ngys /o m a ea r o/z n en se which is fou n d in th e S ou t h
'

, ,

e rn S t ates fro m S outh C aroli n a to Tex a s

,
Th e l arge fa mily .

Cyst ign at h idae i s r eprese n ted o n this co n ti n e n t by o n ly th ree

sp ecies w hi ch are co n fi n ed to M exico an d F lorid a
,
.

Th e Ranidae or t ru e f r ogs i n clude the F irmister ni a with

, ,

cyli nd ric al s acral di apophyses The fa mily com prises n u .

m e ro u s ge n er a o n ly o n e of w hich the typic al ge n us Ran a

, , ,

is fou n d in North A m eric a This ge nus co n t ai n s abou t .

o ne hu n dred an d fo rty speci es which are fou n d in all of ,

t h e co n ti n e n ts o f th e globe al t hough occu rri n g o n ly in th e

,

extreme n orth er n p arts of S ou th A m eric a an d Aus trali a .

There are four t e e n North Am eric an sp eci es t w o of whi ch , ,

R femp o r a r z a an d R ag z lz s occur also in E urop e O n ly a

' ' '

. . .
,

fe w o f th e better — k n ow n forms are here tre ated o f F ull .

d escri ptio n o f t h e speci es m ay be fou n d in C ope s B atr ach i a ’

of Nor t h Americ a .
”

R a n a ca f esé z a n a the bullfrog — This is by far the l argest

'

.
,

of Nort h Am eric an sp ecies of Rana an d o n e of the l argest of ,

th e ge nus I t att ai n s a le n g t h of five to eigh t i n ch es I t is

. .

wid ely distribu t ed thro u ghou t the U n i t ed S t a tes e a st of th e ,

Rocky M ou n t ai n s fr o m M exico to C an ad a
,
Th e color of .

t h e upper surfa ce v a ri es fro m gree n to olive brow n m arked -

w i th sm all d ark er spots Th e h e a d i s u su ally brigh t gree n

.
,

a n d t h e l egs ar e m arked with blo tch es o f d arker color .

Th e d erm al plic ae beh i n d th e eyes are i n dis t i n ct Th e tym .

p an u m is very l a rge esp eci ally in th e m ale The to es o f

,
.

t h e hi n d feet are bro adly webbe d th e web exte n di n g to the ,

tip of t h e four t h t o e .

This sp ecies r arely g o es far fro m t h e w at er I t i s u s u ally .

fou nd ei t her p ar t ly i m merse d in th e w at e r or si t t i ng o n t h e

 TH E A M P H I BI A IN G E N E R AL 19

b an k o f som e po n d or stre am I t m akes for th e w ate r very .

qu ickly whe n al arm ed and usu ally skims alo n g the surface
,

for sever al y ards before divi n g below Accordi n g t o K alm .

,

i t m ay le ap to a dis t an ce of t hree y ards but Abbo tt who , ,

experime n ted with sever al specim e n s fou n d n o n e th a t could ,

j u mp qui te seve n feet The m ales h ave a very lo ud ho arse

.
,

b ass voice which h as bee n co mp ared to th e ro ari n g o f a

,

bull Whe n a n u mb er o f the m are cro aki n g n e ar by the

.
,

n oise as K alm observes i s

,
so loud t h at t w o people t alki ng
,

by th e side of a po n d c an no t u n d erst an d e ach oth er They .

cr o ak all togeth er ; th e n s top a lit t le an d begi n ag ai n I t , .

seems as if th ey h ad a c apt ai n a mo n g th em ; for whe n h e

begi n s to cro ak all the others follo w ; an d whe n he s t o ps
, ,

the o thers are all sile n t Wh e n this c apt ai n gives th e sig n al

.

fo r stoppi n g you h e a r a n o t e like p o o p

,
co mi n g fro m ‘ -

hi m I n d aytime they se ldom m ake any gre at n o ise u n less

.
,

the sky is covered B u t t h e n ight is their cro aki n g ti m e ;

.

an d whe n all is c al m you m ay h e ar the m th ough you are

, ,

n e a r a m ile an d a h alf off

”
.

B ullfrogs fe ed n o t o nly u po n the cre atu r es devoured by

other species of frogs b u t t h ey freque n t ly c ap ture oth er
,

an im als which their sm aller rel at ives ar e u n a ble t o sw allo w .

Th ey ofte n d evour f ull gro w n speci me n s of o th er speci es of

-

Ran a t he you n g o f ducks an d other w a ter fowl an d eve n

, , ,

sm all chicke n s wh ich ve n t ure too n e a r th eir h au n ts .

The bullfrog requires t w o ye ars to complete it s m e t am o r

phosis I h ave o ft e n c aptured its l arge t adpoles be n e ath the
.

ic e in midwi n t er .

A very closely allied speci es o f bullfrog R g ry lz o has

'

, ,
.

rec e n tly be e n d escribed fro m F lorid a by S t ej n eger I t h as .

somewh at lo n ger to es and a d ark er color th an ea z esé z a na

'
‘

an d is s aid to h ave a q ui t e differe n t voic e .

R a n a ela mz fa ns This species has a n e arly u n i fo r m gree n

'

.
20 TH E B IOLO G Y O F TH E FR O G C H AP .

or brow n ish color above m arke d o nly wi t h sm all i rregul ar ,

bl ack spo ts The derm al plic ae are co n spicuous Th e hi n d

. .

legs are short an d the web exte n ds well out o n th e to es .

Th e most co n spicuous fe atu re o f this species i s t h e very

l arge tymp an u m which in the m ale co n sider ably excee ds t h e
,

di am eter of the eye I n the fem ale th e t ym panu ni is c o n .

sider ab ly s m aller bei n g about three fourths th e di a m eter of

,

the eye an d dist an t fro m the l atter by n e arly h alf i ts o w n

di ameter This speci es i s wi dely distributed fro m t he
.
”

E aster n S t a tes to M issouri an d M i n n e sot a an d fro m C an ad a

to F lorid a a nd M ississippi I t is closely co n fi n ed to w ater .

like the bullfrog I t m ay re ach a le n g t h of thre e i n ches

. .

R a n a sy /va nea th e wood frog — U nlike th e t w o preced

'
“

, .

in g speci es R sy lna nea is u su ally fou n d in d amp woods

'

.
,

oft e n far from w ater I t occ asio n ally occurs at a co n sid er .

able el ev atio n o n e spe cim e n h avi n g b ee n t ake n by M r

,
.

Alle n n e ar the top of M ou n t B artlett New H am pshire at , ,

a n alti t u d e o f twe n ty fi v e hu n dred fee t This frog s ays M r -

.
, .

“
Alle n is co m mo n es t in th e be ech woods an d so closely
,

rese mbles in color the d e ad be ec h le aves th at n o t in fre ,

que n tly eve n afte r h avi n g see n o n e j ump it i s wi t h diffi

, ,

cul t y disti nguished fro m t he b ackgrou n d Wh e n frigh t en ed .

it t akes prodigious le aps in an erra tic course arid u su ally ,

esc apes i nt o so m e hole or u n der a log At n ight whil e .

,

w alki n g in a d a mp spot in th e woo ds I fou n d n u mbers of ,

t he m co n greg a te d in t h e p at h where th ey h ad prob ably ,

co m e to feed Ra rely h ave I h e ard the m u t t er a

.

sou n d in the su m mer t hough occ asi o nally whe n in the , ,

woods at n igh t I h ave de t ected th eir fai n t raspi ng cr aw

, ,

au k
’
aw -

R a n a p zp z en s ,
th e l eop ard frog This i s perh aps the m ost
’ ’

co m mo n of all t h e Nor t h Americ an species of Ran a I t s .

grou n d color is gree n m arked wi t h r ath er l arge bl ack

 TH E AMP H I BI A IN G E N E RA L 21

blotches edged wi t h w hitish The legs are crossed above .

with bl ack b ar s whic h m ay o r m ay n o t be i n ter r upte d in th e

middle There are usu ally two irregul ar rows of bl ack spots
.

o n th e b ack be twe e n th e pro mi n e n t derm al plic a e ; th e

,

lower sid e of the body is p al e Th e tymp an i are sm alle r .

th an th e eyes an d there i s n o bl ack e ar p atch Th e vomer .

in e teeth lie betwee n t h e pos terior n are s Th e legs are .

lo ng so th at wh e n th e heel i s brough t forw ar d i t exte n ds in

,

fro n t o f t h e ti p of the s n out .

C ope di s t i n guishes four v arieties o f th is speci es for a ,

descriptio n of wh ich the re ad er m ay be r efe r r ed to this

author s B atr achi a o f Nor t h Americ a
’

Ra n a p a ins ir z s t h e p ickerel fro g —This species re

.

'

se mbles the precedi ng o ne I t i s usu ally brow n ish in colo r .

,

with t w o ro w s o f l arge rect an gul ar d ark brow n blo tches

betwee n th e derm al plic ae There is a brow n spo t a bove .

e a ch eye and a d ark li n e betwee n th e eye an d the n ostril .

The body is whi t ish below bu t th e lo w er side of the hi n d ,

legs is yello w E xter n al voc al s acs are abse n t

. .

This spe cies i s qu ite commo n in the e aste rn p art of the

U n ited S t ates I t is s ai d by C op e to pre fer cold sp ri n gs
.

an d stre a mlets but is of all our frogs th e most frequ e n tly

,

se e n in the grass .
”

RE F E RE NC E S
A b b ot t , C C . . A N t u lis t s a ra
’
mbl s b u t H m d d
Ra e a o o e, 2 e .
, 1 8 94 .

Allen , G M . . No t es on th R pt i l s
e d A m p hibi
e s fI t e an an o n erval e ,

Ne w Ha m pshi re . P c ro . B os . S N t H is t V l 9 9
o c. a . .
, o . 2 , 1 01 .

B ou l enger, G . A . T he T i ll ss B t chi s f E u p 8 9 7
a e a ra an o ro e, 1 .

B reh m Thi l b E d 7
, A C . . er e en, . .

Cop e E D B t ch i f N t h A m ic er a. A rt “A m h ibi a in t h e
, . . a ra a o or .
p
River sid N t u l H is t y
e a ra or .

D m é il et B ib on
u rE p é t l gi G é é r . r o o e n ral e o u H is t i c m plet
o re o e des
R p t il s
e e .

Diirigen B D u t sc h l d Am p hibi
, . e an s en u n d Rept i ien , l 1 8 9 7.
22 TH E BIOLOGY OF TH E F RO G CH A P .

F isch e r -
S ig w a rt , H b ch t u g
. u s
B i l gisch
Am o o e B eo a n e n an n e re n

p h i en . V iert elj a h h d N t f G s ll
rsc . Z u ich L X I I J h g 8 9 7
. a ur . e e . r , , a r . 1 .

G adow , H . A m p hibi d R p t il s V l 8 f th
a C mb idg
an e e . o . o e a r e

Nat u ral H is t y or .

H ay , O P . . T h e B at ra ch i s an an d il s f t h S t t f I di
Re p t e o e a e o n a n a,

1 8 9 2, 1 7t h A n n Re p . . D pt G le . eo . an d N tu l R s u c s
a ra e o r e .

H off m a nn, C K . . A m p h ibi n e ,

”
i B
n Cl ss
ro n n s dO d u g
’
a en u n r n n en

des T h ie r r eich s, B d V I , . 2 .

H olb ook , r J . E . No rt h A m e ric an H e rp e t o o g l y .

J ordan D , . S . A M an u al o f th V e e rt e b rat e A im ls n a of the No rt h ern

U it d S t t s
n e a e , 9 th e d , . 1 9 04 .

L ey dig, F . Die a n u re n B at rac h ie r des d u tsche en F un

a a, 1 8 77 .

Rfis el V O I1 Rosen h of . H is t i o r a n at u lis ra r a n aru m n o s t rat iu m , 1 75 8 .

S p all an z an i, L . E xp erie n ces p o u s i

r erv r a l H ist o ire de la
’
ge e n ra

i
t o n , 1 78 7 .

cc u t s f th
A o n o e g en eral a n at o my of t h e f ro g are c o n ta n e i d in t h e
f ll i g k s
o ow n w or

B ou ne, r G C . . A n I n t ro duc t i on to the S t udy of t h e Co mp arat ve i

A n at o my of An i m ls a , 2 vo ls 900
.
, 1 .

E ck e r , A . A n at o my o f t he F g ro , t ran sl at e d by G eo r g e H a sl ma ,

Oxf d or ,
1 8 89 .

E c e k r u nd W iedersch eim . A n at o mi d F e es ro sc h es, au f Gr u d n

e ig e n er U n t er suchu n ge n du ch us r a n eu b b it t
ea r e e vo n Dr E rn . st G au
pp,
1 8 9 6—1 90 4 .

Ho w es , l m t y Z t my 9
G B . . At l s a of Pr c t ic a al E e e n ar oo o , 1 02 .

H xley and M rt in
u G l B i l gy 889 a . e n era o o ,
1 .

M ar h ll A M T h F g
s I t duc t i
a ,t A t my 6 d 8 96
. . e ro : an n ro on o na o , c .
, 1 .

M ivart S t G eor g Th C mm , F g 8 74 . e . e o on ro , 1 .

P r k r an d P ar k r
a e A El m t y C u s i P c t ic l Z l gy e . n e e n ar o r e n ra a oo o ,

1 900 .

V og t u nd Y u ng . Le h buch d
r er p ra k t i sch en ve rg e l ich d en en An at o
m ie, 2 Bd .
11 HA BI T S A ND N AT U R A L H IS T ORY OF TH E FROG 23

C H A PT E R I I

TH E HA BI T S A ND N AT UR A L H I STORY OF TH E FRO G

H abit at The h abit at of Ra n a p zp zens lik e th at o f m ost

' '

.
,

species of frogs i s usu ally in or n e ar t h e w at er I n d a mp o r

,
.

we t we a ther however this species freque n tly w an ders for a

, ,

co n side r able dist an ce fro m it s aqu at ic ho me I t i s li able to .

be fou n d al most an ywhere n e a r the sh ores o f l akes po n ds or , ,

stre ams in the wide terri t ory over which it is distributed .

I t s r ange as give n by C op e is fro m

“ Ath ab asc a L ake in th e ,

n or t h,
to G u at e m al a i n clusive in the sou t h an d fro m th e
, ,
”

A tl an tic co ast to the S ierr a Nev ad a M ou n t ai ns I t has .

,

therefore th e wides t distributio n of any of the North A m eri

,

c an speci es o f Amphibi a although i t i s n o t k n ow n to occu r

,

o n th e P a cific slope .

Th a t Ra n a p zjmens is co n fi n ed to the n eighborhood o f

'

w ater depe n ds in gre at m e asure o n th e fact t h at the ski n

must be kept moist in ord er th at cut an eous r espiratio n m ay
t ake pl ace As soo n as the i n tegume n t becomes dry as it
.
,

quickly does i f th e f rog is exposed to a w ar m dry atmosphere ,

it is n o lo nger c ap abl e of servi n g as an org an o f r espiratio n ,

and the an im al soo n perishes The frog u nless i t i s amo n g

.
,

w e t gr ass or weeds or in a m oist a tmosph ere

,
must r e m ai n ,

where i t c an moist e n th e ski n by an occ asio nal plu nge i n to

t h e w ate r A n oth e r circumst an ce w hich se r ves to ke ep th e
.

frog in close proximity to w ate r i s the me an s thus afforde d

o f esc api n g fro m e n em ies A n y o n e w ho has w alked alo n g
.

t h e m argi n of a po n d o r st r e am must h ave observed th a t

24 TH E BIOLO G Y OF TH E FROG CH A P .

wh e n a frog is st arted u p i t almost i nv ari ably m ak es a j u mp

for the w ater I n this w ay th e cre ature has a re ady mod e of
.

esc api n g n o t o nly fro m m an bu t fro m a n u mb er o f other

, ,

e n emies which migh t e asily overt ake i t in a fair field Aft e r .

i ts fi r st plu nge i n to th e w at er th e frog usu ally swi ms so me

dist an ce u n der the surfac e an d th e n com es up exposi ng ,

o n ly the tip o f it s s nout above t he w ater to get air F re .

qu e n tly i f th ere is gr ass or w eeds n e ar t h e w at er s edge the

,
’
,

frog will swim a few stro kes aw ay from t h e shor e and the n
tur n b ack an d quietly co m e to th e surface a mo n g t h e vege
t at io n wh ere i ts adve n t would usu ally n o t b e suspected by
,

th e observe r .

D uri ng th e breedi ng se aso n in t he spri ng frogs are more ,

closely co n fi n ed to t h e w a ter t h an at o t her ti m es of th e ye ar .

I n th e su mm er th ey w an der fa rth er fro m th e w ater in se arch

o f food D iffere nt sp ecies v ary gre atly ho w ever in this
.
, ,

respect Th e wood frog Ra n a sy /va lz ea i s co m mo n ly

'

.
, ,

fou n d in woods m il es aw ay fro m an y po n d or stre am M ost of .

t h e oth er Nort h Am eri c an species of Ran a are mo r e closely

co n fi n ed to an aqu ati c h abi t at I n E urope the w a t er fr o g

.
,

R esr n /en fa is decid edly aqu at ic in its h abi t s wh ere as o th er

.
, ,

spe cies co m mo n ly sp o ke n o f as the gr ass frogs sc atter

, ,

t hrough the m e adows and woo dl an ds a fte r th e breedi n g

se aso n .

F ood — Th e food of frogs co n sists of e arth w orms in

.
,

sects spi ders in fact o f al most an y ki n d o f an i m al s m all

, ,

e n ou gh to be seiz ed an d s w allowed L arge fr o gs h ave n o .

se n ti me n t al scruples ag ai n s t d evouri n g their sm aller rel atives .

Th e l arge bullfrog is an especi ally d angerous e n emy to o t her

m embers of i ts ge n u s I h ave oft e n fou n d t h e stom ach o f
.

t his an i m al gre at ly diste n ded from i ts h avi n g sw allowed

n e arly full grow n speci m e n s o f R a n a arth w or m s

’ ’

ppz z en s E .

ar e a favorite ar t icle o f die t a hu n gry frog will d evour sev

26 TH E BIOLOGY OF TH E F RO G CH A P .

which the to n gue is covered e nable s i t to adhere to th e

obj ects it st r ikes ag ai n st so th at they m ay be co n veye d to
,

the mouth .

The frog has an i n sti n c t to s nap at sm all movi ng obj ects

th at come su fficie n tly n e ar Thi s actio n is dete r mi n ed m ore.

by th e motio n an d size of the obj ects th an th ei r form Un .

less a thi n g is m ovi ng th e frog p ays little at te n tio n to i t

,
.

F rogs m ay oft e n be c augh t by d angli ng sm all bi ts of red

y ar n before th e m o n a hook Wh e n the y ar n is seized th e .
,

an im al m ay be j erked ou t of th e w a ter Accord i n g t o .

K nau er frogs and t o ads h ave th e powe r of ej ec t i n g i n diges t

,

ible bodies fro m th e stom ach by w ay of th e m outh B i t s of .

gr ass or moss accide n t ally sw allowed with the food are gotte n
rid of in thi s w ay .

P rot ru sion of t h e Ton gu e — Th e frog is able to throw .

out its to ngue wi th rem ark able rapidi ty but th e m e thod by ,

wh ich this fe at is ac
co mplish ed w as u n t il ,

rece n tly but in ade ,

q u a t e ly u n d e r s to o d .

1
H artog an d G au p p
2
.

h ave fou n d th at th e
protru sio n is brought
a bout by th e pressure

o f th e lymph in th e
FIG 6. Figu h m g t h t g u f t h hu ge subli ngu al ] ym p h
. re s ow e on e o e

f g i t h diff nt p si t i s (A ft
ro n re e e re o on er
h I be a
.

W i d h im )
e ers e .
sac T l na y . b
.

re adily show n i f we cut

off the upper j aw o f th e frog an d i nj ec t air or liquid through
th e mylohyoi d muscle whi ch exte n ds be n e at h th e to ngue
, .

The lymph sp aces beco me filled an d this c auses t h e to n gu e ,

1 H ar t o g , Ann . Na t . H u t ,
M ay, 7 ,
1 90 1 .

2 G a u p p A n at A n z 1 9, 1
, . .
, 90 1 .
11 HA BI T S A ND N AT U R A L H IST O RY OF TH E FROG 2
7

to be r aised u p an d th r ow n forward s ays H artog

.
“I f
, ,

“w e i nj ect wi th melted coco a butter colore d with car

mi n e or alk an et an d keep u p the pressu r e u n til the m ass
,

sets we fi n d th at i t fills an e n ormous lymph sac b et w ee n the

,

mu scle an d th e body o f the hyoid exte ndi n g through the

,

m edi an i n termuscul ar fissure i n to th e to n g u e itself se n di n g ,

bran ch es betwe e n th e f an sh aped r amific atio n of the in t rin

-

si c m uscles at the edges of th e to n gue an d i n to i ts termi nal

d il at a tio n s
. Accordi ng to H artog th e co n tractio n o f the
”
,

mylohyoid muscle expels the lymph fro m the subhyoid sp ace

i n to the to ngu e and t hu s effects the protrusio n of th is org an
L ocomot ion —Th e loco motio n of th e frog is effected by
.

le api n g and swi mmi ng and in both o f th ese oper atio n s the
,

lo n g hi n d legs pl ay th e chi ef p art I n th e ordi nary resti n g

.

posi tio n th e body is i n cli n e d upw ard in fro n t bei ng sup ,

ported o n th e fore legs which are held in a peculi ar twist so

,

th a t th e l arge thu mb poi nts n e arly b ackw ard th e posterior

p art of th e body rests upo n the grou n d an d th e hi n d li mbs,

are fold ed up re ady fo r a spri n g No preli mi n ary move

.

me n ts are required in order to get th e ani m al in re adi n ess

fo r e sc ape B y a su dde n exte n sio n of th e hi n d legs the
.

body is pro pelled through th e air I n le api ng th e fore.

,

li mbs are u sed more to hold u p the an terior p art of th e

body an d to poi n t th e ani m al in the desired direc t io n o f
m ove me n t th an as actu al org an s for propulsio n I f o ne .

c auses a frog to le ap in v arious d irectio n s it will be observed ,

th a t the body is adj usted before e ac h le ap in a n ew direc

tio n by th e movem e n ts of th e fore limbs A n ordi n ary .

sp ecim e n of R a n a p zpz ens m ay le ap fro m two to three fe et

'

The movem e n ts of t h e hi n d legs in swi m mi n g are very

much like those perfo rmed in j u mpi n g I n both operati o n s
.

th e hi n d legs are alter n a tely dr aw n u p in t h e fo r m of a Z an d

q uickly exte n d ed As they are pushed b ack t h e toe s are
.
,
28 TH E BIOLO G Y OF TH E F RO G ca n » .

spre ad ap art an d as th e web betwee n them aflOrds a c o n

,

sidera b le r esi st an ce to p assi n g through the w a ter th is m o ,

t io n gives the body a forw ard i mpulse The fore li mbs are .

h eld b a ck agai n st t h e body a ft er th e strok e an d i f t h e frog

, ,

does no t m ak e sev er al s t rokes in quick successio n th e hi n d ,

li mbs are held exte n d ed behi n d th e body so th at t h e an i m al ,

affords as littl e resis t an c e as possible to glidi n g through the

w ater The fore li mbs are also u sed in swi m mi n g t aki n g

.
,

st r okes so me ti mes togeth er an d som eti m es alter n ately To .

a cer t ai n exte n t th ey aid in propelli n g the an i m al forw ard ,

but they are also employed as in loco motio n o n l an d to

, ,

guide th e directio n o f move me n t Whe n th e an i m al st arts

.

to s w i m dow nw ard the fore legs be at b ackw ard an d upw ard

, ,

th e h an d bei n g twis ted so as to press i ts bro a d surface

ag ai n st the w ater This n atur ally pu sh es th e an terior p art
.

of the body dow n I n s t arti ng to swi m u pw ard the fore

.
,

legs b e at dow n w ard elev ati n g t h e an terior p art of the body

, ,

which i s th e n pushe d upw ard by th e strokes of th e h i nd legs .

The fore legs are also used in c ausi n g th e body to m o ve

from S id e to sid e and u n equ al move m e nts of the hi n d legs
,

are employe d for the s a m e purpose B e n di ngs o f th e body

.

are also u sed to h elp steer th e course o f t h e an i m al Th e .

hi nd legs u su ally m ake a stroke at the s am e i n st an t but th e ,

frog n o t i nfreque n tly u ses the m alter n ately esp eci ally wh e n ,

struggli n g n e ar an obst a cle .

At t itu de w h en F loat in g on t h e S u rf ace — Wh e n frogs

are kept in w at er b eyo n d their dep t h they sp e n d a co n sider
,

able portio n o f th eir ti me at t h e surface with j ust th e tip o f

the n ose exposed for the purpos e of bre at hi n g air The

,
.

dist an ce wh ich t h e h e ad proj ects fro m th e w ater m ay be

v ari ed at will as i t d epe nds u po n th e am ou n t of air t ake n
,

i n to t h e lu n gs The m o re t h e lu ngs are i n fl ated th e less th e

.
,

s p ecific gr avi t y of t h e an i m al bec o m es a n d t h e higher there

, ,
II HA BI T S AND N AT U R A L H I ST ORY OF TH E F ROG 29

fore , i t rises in the w at er Wh e n at the surface the frog

.

u su ally li es quiet h an gi n g obliquely wi t h the hi n d legs in a

,

s t ate of m oder ate exte n sio n The fore legs ge n er ally are
.

h eld ou t fro m th e body I n su ch a positi o n th e frog m ay

.

r est fo r a lo n g ti m e wi t hou t p erformi n g any o t he r move

m e n ts besides those i n volved in respir a tio n The exte n ded .
,

sprawl ed ou t atti t ude o f the frog wh e n resti n g at the sur

-

face co n tr asts m arkedly with its r esti n g posi t io n o n l an d ,

whe n its hi n d legs are closely doubled up and alre ady se t

for a spri n g O n e prob able re aso n for th e ex t e n sio n of th e
.

hi n d legs is th at there is n othi n g to suppor t t hem fro m

below an d they would n atur ally h an g do w n wh e n rel axed
, , ,

from their o w n weight H owever this m ay be t h e exte n ded

.
,

co n ditio n of th e h i n d limbs i s of service in e n abli n g th e

an im a l to sudde n ly dr aw i t self dow n w ard whe n eve r d an ger

th re ate n s fro m abo ve .

D ivin g . I f a frog is appro ached wh e n it is resti n g at the

su rface of th e w ater it will dive do w nw ard w ith gre at cele r ity
,

an d m ake sever al strokes c arryi ng it some dist an ce aw ay

,

fro m i ts resti n g pl a ce Th e actio n is performed so qu ickly

.

th a t i t i s no t e asy at first to see how it is accomplish ed At .

o n e mom e n t th e frog i s resti n g in perfec t quie t an d at the

n ext i n s t an t we perceive h i m m aki n g vigorou s kicks an d

r apidly swi m mi ng aw ay B y experi me n t i ng wi t h frogs kep t

.

in a gl a ss dish an d co n ce n tr ati n g our atte n tio n o n o n e

fe ature o f th eir beh avior at a tim e we m ay g ai n an id e a ,

of the w ay this fe at is accomplished To s w i m dow n w ard .

through th e w ate r the an i m al h as to r everse i ts posi t io n as ,

a n ex t e n sio n o f th e hi n d legs in i ts n orm al r esti n g a tti t u de

w ould t e n d to thro w it ou t o f t h e w a ter Th e fi rs t move .

m e nt is th at of wi t h dr aw al from th e surface which is ac co m ,

p lish e d by sudde n ly bri n gi n g t h e h i n d legs forw a rd t hus ,

givi ng th e body a b ackw ard i mpulse This bri n gs th e hi n d .

3 0 TH E BIOLOGY OF TH E FROG CH A P .

li mbs up i n to a positio n for m aki ng th e ordi n ary s w i m mi ng

s troke Alo ng wi t h t h e withdr aw al o f the body fro m th e
.

surface th e fo re legs m ake a sudde n stroke b ackw ard an d

u pward t hus t hrowi n g th e an terio r e nd o f the body dow n
,
.

The n the hi n d legs ex t e n d an d shoo t the an im al farther

dow n ward through th e w a ter The a t t itude of the body as .
,

t h e frog rests at the su r face i s o n e of prep ar atio n fo r t h e ac t

,

o f divi n g j ust as its at ti t ude o n the grou nd is o n e o f readi

,

n ess for a spri n g At th e m om e n t th e frog le aves t h e su r

.

face bubble s o f air m ay ge n e r ally be s ee n to esc ape fro m the

,

n os t rils .

Right ing Movement s L ike most an i m als th e frog whe n

.
,

pl ace d upo n i ts b ack will reg ai n it s n orm al posi t io n I t .

does so too w ith re m ark able qui ck n ess c ert ai n ly in less

, , ,

th an h al f a s eco n d The op eratio n i nvolves the co Ordinat ed

.

actio n o f sever al muscles The posi ti o n of equilibriu m m ay

.

be att ai n ed by rolli n g ove r ei t h er to th e right or to t h e l eft ,

a n d a frog will do n o w th e o n e a nd n o w t h e o t h er so m e ,

t i mes hesi t ati n g a mo m e n t betwee n th e two cours es A frog .

wil l right i tself a gre a t m an y ti m es in qu ick successio n a nd ,

in course of ti me will beco m e so fatigue d th a t i t will ac t

slowly e n ough to give the observer a ch an ce of followi ng its
move me n t s These mov eme n t s v ary a good de al in differe n t
.

a cts but they com mo n ly occur in abou t the followi n g w ay

,

I f the frog rolls over tow ard it s le ft sid e th e right h i n d leg is ,

brought dors ally by a co n t r ac t io n o f t h e muscles o f th e dors al

side of the thigh ; th e muscles o f t h e ve n t r al sid e o f th e left
thigh also co n t r act bo t h th ese move me nt s te n d to roll t h e
body ove r to t h e left The righ t hi n d leg is oft e n brou gh t
.

for w ard so th at t h e thigh li es at a co n sid erable angle fro m

t h e body an d this gives t h e li mb a gre a ter purch ase in roll
,

ing t h e body over The l eft fo re leg is brough t d ow n alo ng

.

side of the body and the opposite m e mbe r is th r ow n ove r t o

,
II HA BI T S A ND N AT UR A L H IS T O RY OF TH E FRO G 3 1

the left side thus assisti n g the hi n d legs in th e act o f

,

r otatio n .

The V oice The cro aki n g of Ra na pzpz ens m ay be r ep

’ ’

r ese nted al t hou gh r at h er i nadequ ately by th e syll able s au

, ,

au au au au k
- - -

The voice o f th e m ale is loude r and de ep e r

.
”

th an th at o f th e fe m ale an d is more o fte n h e ar d I n l arge .

frogs th e n otes are deeper th an in sm all o n es Th e n otes .

o f frogs are m ore o ft e n he ard du r i n g th e breedi n g se aso n ,

whe n th ey are supposed to s erve th e purpose o f a sex c all .

I n the sum m er however i t i s n o t u n u su al to he ar th e

, ,

cro aki n g of frogs esp eci ally in th e eve n i n g A d am p at

, .

m o sph ere is co n ducive to their so n g an d for this re aso n th e ,

voices of th ese ani m als are ofte n h e ard upo n th e appro ach of
a shower Th e tre e frogs see m to be especi ally s e n sitive
.

to atmospheric ch an ges an d the popul ar rep u t atio n wh ich,

these cre atures e nj oy as prog n ostic ators of the we ath e r is

n o t e n tirely u n meri t ed .

Th e cro aki n g o f frogs is re adily produ ced by r ubbi n g t he

b ack o r side of th e body After e ach strok e th e frog u su .

ally respo n ds by a cro ak an d th e n l apses i n to sile n ce .

C ro aki n g is oft e n c au sed through accide n tal co n t act with

oth er i n dividu als Two frogs which wer e kep t in a dish o n
.

my t able were in the h abi t o f cro aki ng at freque n t i n t erv als ,

an d I o bserved th a t e ach t ime th e b a ck o r side o f o n e frog

w as touched by the o t her th e i n dividu al woul d r espo n d by ,

a cro ak I f n o t distu r bed th e frogs would r em ai n sile n t

.
,

i n defi n i tely .
'

F rogs cro ak as w ell u n der w ate r as o n l an d As th e air .

is forc ed out o f th e lu ngs p ast th e voc al cords i n to th e , ,

m outh the ex t er n al n ares ar e closed so as to preve n t i ts

,

esc ape The n t h e bucc al c avi t y co n t r acts fo r ci n g the air

.
,

b ack i n to t h e lu n gs ag ai n ; an d t h e s am e process is rep e ated .

I f th e he ad of t h e frog is h eld u n der w ate r while th e an i m al

32 TH E BIOLOGY OF TH E FROG CH AP .

is cro aki ng i t m ay be see n th at th e air is forced b ack an d

,

for t h betwee n the mou t h an d th e lu n gs while o nly a li t tl e i f , ,

a n y is allowe d to esc a pe through t h e n ares

,
.

U n d er co n ditio ns which are p ar t i cul ar ly agree able frogs ,

oft e n give ou t a low gru n ti n g sou n d as i f o f co n t e n t me n t .

O n th e other h an d whe n frogs are severely i nj ured t hey

, ,

so me t i mes u tter a sort o f cry which is c alled the p ai n

scre a m Whe n seized by a s nake or o t h er e n e my m an y
.
,

speci es o f frogs m ay respo n d by m aki n g thi s pi t eous cry .

I n st in ct s f or P r ot ect ion Wh e n a frog i s seized in the

.

h an d s it usu ally m akes vi ole n t efforts to esc ape I f i t is

,
.

held by t h e an terior p art o f the body th e hi n d l egs a re u sed

,

to push ag ai ns t o n e s h an d with co n sid er able forc e At th e

’
.

s a m e ti m e t h e b o dy is ge n er ally i n fl at ed wi t h air which ,

e n ab les i t to slip aw ay m ore re ad ily fro m o n e s gr asp The ’

.

sudd e n ej ectio n o f flui d fro m t h e bl add er which t akes pl ace ,

whe n the frog is c aught m ay also b e o f o cc asio nal service in

,

it s at t e mp t s t o ge t free .

F rogs som e t i mes s w ell t h e body before bei n g seized as i f

in ant ici p at io n o f th eir c ap tu re an d they are especi ally ap t
,

t o do this a ft er bei n g lightly touched Touch a frog th at is

.

resti n g quie t ly an d i f th e cre at ure does n o t hop aw ay o n e

, ,

m ay se e the body p u ff u p ; an d i f t h e body is t ouched two

or thre e tim es th e swelli n g w ill co n t i n ue u n t il t h e lu n gs co n
,

t ai n their m axi mu m amou n t of air A n an i m al such as a s n ake

.

which w as a t t e mp t i ng to s w allow a frog would fi n d t he

o p erat io n some w h at more di fficult i f t h e b ody of it s vi cti m
w ere s t ro n gly i n fl a ted F rogs o ft e n avoid c a pture be t t er by
.

re m ai n i ng perfec t ly quie t t h an by att emp t i n g to get aw ay

by j umpi ng F e ar prom p t s t h e cre at ures n o w to th e o n e
.

an d n o w to th e o t h er me t hod o f esc ap e S afe ty is also .

sought occ asio n ally by crouchi n g close t o th e grou n d an d ,

m ore ofte n by cr aw l i n g u nd er some obj ect th at p r omises to

affo r d she lt er .
34 TH E BIOLOGY OF TH E FROG CH A P .

In th e fall o f the ye ar t he body is richly stored wi th

n u t rime n t accumul a ted duri n g the summ er whil e food is
ab u n d an t D uri ng the wi n ter this m ate r i al is e mployed n o t
.

o nly in m ai n t ai ni n g t h e te mp er atu r e of t h e body an d fu r nish

ing t h e e n ergy n ec ess ar y to c arry o n th e v arious a ctivities of
t h e org an s bu t it is dr awn u po n to co n t r ibute t o th e growth
,

of the reproductive c ells A p art o f this m ateri al is stored .

in the muscl es which duri n g th e wi n t er decre ase in we igh t

,

in rel atio n to th e res t of the body G aule fou n d th at in

1
.

fe m ale frogs killed in July the gastroc n e mius muscle weigh ed

o n t h e ave r age mg for every gr a m o f body weight I n
. .

August the r a tio rose t o I n D ece mber it s ank to

I n J anu ary it w as an d in J u n e th e l ayi ng period , ,

I n th e m ale t h e decre ase in rel ativ e weight o f the m uscles

is n o t n e arly so gre at as there is much l ess m ateri al to be
,

employed in th e develop m e n t of the sexu al p r oduc ts .

The liver u n dergoes m arked se aso n al ch an ges wh ic h will

b e m ore fully d escribed in co nn ec t io n w ith the accou n t of
th at org an I n th e wi n te r it co n t a i n s a l arge a mou n t o f
.

glycoge n which almost e n tirely dis app e ar s by th e en d o f th e

,

breedi n g se aso n U n ti l e arly spri ng however the glycoge n

.
, ,

suffers comp ar atively li t tle loss The color o f th e liver a lso .

v ar ies betwee n wi n ter an d su m mer owi ng p r ob ably to differ ,

e n c e s o f n u t ri t io n I n wi n ter th ere is a n a ccu mul at io n o f

.

pigm e n t which gives th e liver a d ark appe ar ance I n su m .

m er t his pigme n t in most frogs l argely dis appe ars and th e

liver beco m es lighter in color Th e size o f t h e cells v ari es .
,

i n cre asi ng through th e su m mer re achi n g its m axi mu m in ,

Ra n a temp o r a ma in Nove mber the n d ecre asi n g through t h e

'

w i nt e r an d e arly spri n g re achi n g th e mi n i mu m in April

,

( L eo n ard )
2
or M ay ( F u n ke
)
3
The size o f th e liver in .

1 Ga ul A r e/z gr ey P /zy 5
e, B d 8 1 , 1 90 0
. . .
, . .

2 L eo n ar d
A r c/z A n a t n P /zy r p
, . Ab t h S p p , 1 8 8 1
. . .
,
hys . . u l . .

3 Fu k
n e , D en R S C/
Zh
,
/ z n A é ad
l V e at n a t Cl B d 6 8 1 9 00 . m h . . . .
, .
II HA B I T S A ND N AT UR AL H IST O RY OF T H E F RO G 35

r el atio n to th e rest of th e body acco r di n g to L angendo rfi

,
'

P loetz an d F u n ke eve n i n c r e ases du ri n g the wi n ter mo n ths

, ,
.

After the breedi ng se aso n the mi n i mu m size is re ached afte r ,

which th ere i s a gr adu al i n cre ase duri n g th e su mm e r .

App ar e ntly therefore t here is eith er a growth of th e live r

, ,

duri ng th e W in t er at th e e xpe nse of th e rest of the body or ,

th e v ar ious oth e r org an s decre ase m ore r apidly th an the

liver in size .

The blood of the frog u n d ergo es in th e spri n g a ft e r the ,

an i m al h as begu n to t ake food a r apid rege n er at io n a pro

, ,

cess which in high er an i m als t akes pl ac e at all ti mes of t he

ye ar There is a gre at i ncre ase in th e n u mber o f blood
.

corpuscles bo t h red an d white Th e m arrow of th e bo n es

,
.
,

where th e n ew blood cells are m ai n ly produced shows in th e ,

spri n g a lymphoid struc ture beco mi ng more an d more fatty

,

toward fall after the productio n o f n ew blood cells has

,

m ai n ly ce ased .

Th e ch an ges in th e fat body at di ffe r e n t ti mes of ye ar h ave

bee n studied by P loetz and F u n ke both o f who m fou n d ,

in the t w o species studied (Ra n a temp o r a r z a a n d R a na

'

es en / en fa
) t h a t t his org an ch a nged but li t t le duri n g th e

wi n t e r mo n ths bu t suffered a m arked dimi nuti o n in size j ust

,

be fore an d duri n g the bre edi n g period in the l ate spri ng .

Aft er this the r e i s a gradu al i n cre ase in t he size of the fat

body u n til fall whe n it re ac hes its m axi mu m
, .

Th e adve n t of the breedi n g se aso n i s m ar ked by gre at

ch an ges in th e reproduc t ive syste m both in the go n ads or , ,

org an s which pro duce the sex cells an d t he v arious accessory ,

org an s Th e v ari a tio n in th e size of th e ov ary b efore and

.

a fter the disch arge o f t h e ripe o va i s e n ormous After t h e .

eggs are l aid in th e spri n g th e ov ary shrivels to a sm all

,

fr ac tio n of i ts previous di me n sio n s D uri ng the summ er it .

i n c r e ases in si z e an d in th e fal l i t m ay fill most of the bod y

,
36 TH E B IOLO G Y OF TH E F RO G CH A P .

c avity The oviduct i s als o e n l arged be fore an d du r i ng the

.

breedi n g se aso n Th e gl an ds in i ts w all r e ach a high d egre e

.

of d evelopm e nt an d secre te an e n ormous a m ou n t of a mucus

like subst an ce arou n d t he eggs as they p ass dow n th e lu m e n .

Aft e r the eggs a re disch arged th e gl ands di mi nish in size an d

,

activi ty an d the size of th e whole duc t i s much red uc ed

, .

Th ere is a di mi n u tio n in th e size o f the testes a fte r th e

esc ap e of the sperm atozo a and th e n a gradu al i n cre ase in
si z e duri ng t h e su m mer u n til fall .

C orrel ated w ith t h e rip e n i n g of t he sp erm a to z o a an d the

appe ar an ce o f sexu al i n sti n cts o f th e m ale frog th ere i s an

i n cre ased d evelop me n t o f t h e b as e of t h e i nn er digi t of th e

fore arm an d an e nl arge me n t o f cert ai n m uscles which are
c o n cer n ed in th e cl aspi ng reflex B oth the i nn er digi t and
.

cl aspi n g muscles are l arge r in th e breedi ng period t h an at

other tim es an d it is prob able th at th eir i n cre a sed develop
,

m e n t is dep e n de n t u po n ch an ges t aki n g pl ace in t h e sexu al

gl an ds S ome t im es th ere are cert ai n exter n al ch aracters
.

develope d in th e fe m ale also duri ng th e breedi n g se aso n I n .

th e fem ales o f R a n a femp o r a r z a H uber h as described der

'

m al p apill ae which o ccur especi ally u po n the b ack an d sid es

of th e bo dy an d the u pper surface of the legs O n the b ack .

th ey are usu ally co n fi n ed to the posterio r h alf of th e body ,

but o n the sides th ey exte n d fo r w ard n e arly to th e tip o f the

n os e. I n the m al e th e ski n i s e n tirely smooth or possesses
in a few c ases o n ly ve r y sm all p apill ae The color of these
.

p apill ae is a whi tish o r ligh t rose an d they are rou n ded

,

or co n e sh aped in ou tli n e and fo u r to five millimeters in

-

di a meter Th ey are richly supplied wi t h blood bu t are

.
,

e n tirely d evoid of d ark pi gme n t Wh e n sec t io n ed th ey

.

are show n to be due m ai nly to a t hicke n i ng o f t h e ou t er

por t io n of th e cu tis and t o be m ade u p l argely of c o n

n ec t ive tissu e The ove rlyi ng epid ermi s i s no t n o tice ably
.
11 HA B I T S A ND N AT U RA L H I S T ORY OF TH E F RO G 37

thicke r th an it i s elsewhere S i n ce th ese org an s app e ar

.

duri n g th e breedi ng se aso n it i s prob able th at th ey h ave

,

some fu n ctio n in rel atio n to reproductio n I f th ey do no t .

direc tly serve to e nable th e m al e to ret ai n hi s hold o f the

fe m ale th ey m ay ac t as s t i muli c ausi n g hi m to cl asp m ore
tightly whe n h e feels th e fem ale slippi n g fro mhis grasp
, ,

Color Ch an ges O n e of th e most rem ark able ad apt atio n s

.

of m any ki nds of frogs for co n c e al m e n t fro m t heir e n emies ,

i s th e power of ch angi n g th eir colo r in h armo n y with t hei r

surrou ndi ngs Th e tree frogs possess t his property in th e
.

h igh est degree Wh e n th e se an im als are a mo n g the g r ee n

.

le aves o f a tree t hey assum e a bright gree n colo r Whe n

, .

o n the b ark their ski n tur n s t o a gr ay or bro w n

,
I n bo th .

c ases the color of t h e frog closely rese mbles th a t o f the

surrou ndi n gs an d serves to m ak e i ts p ossesso r di fficult to
disti n guish The v alu e of such a powe r as a m ean s of
.

pro tec t io n fro m e n emies i s obvious No frog however .

,

rem ark abl e m av be th e ch an ges in color i t m ay u n d ergo is ,

able to assu me all sh ades an d hu es F rogs possess t h e .

property o f ad ap ti n g th e mselves o nly to th e predomi nan t

colors of thei r e n viro n m e n t whic h are gree n th e color o f
, ,

veget atio n an d so m e sh ade of gr ay or brown th e usu al

, ,

color o f th e soil an d the b ark of tre es They c ann ot tur n .

re d or bl ue or viole t an d in fact th e power to d o so would

, , ,

be o f lit t l e v alue to th em i f they possessed i t .

Ra n a p zp z en s like m ost o f t h e m embers of its ge n us

’ ’

, ,

possesses a mu ch less r ange of color v ari atio n s th an th e

tree frogs ; n everth eless it can ch an ge i ts colo r to qui t e a
m arked d egree I f in a d ark e n viro n m e n t its ski n beco m es
.
,

muc h d arker ; the bl ack spo ts co n t ai n so much pigm e n t

th at t h ey rem ain u n ch an ged u n der all co ndi t io n s but th e
‘

lighter regio n s betwe e n the m are subj ec t to m arked ch an ges .

E xposu r e to b r i ght light giv es th e ski n a much lig h te r co lo r,

3 8 TH E BIO L OG Y O F TH E F RO G CH A P .

th e gree n an d gol de n c olors com e out to a much gre ate r

exte n t an d the bl ack pigm e n t cells beco m e less co n spicuous
,
.

There i s little doubt th at pow er of color ch ange in Ra n a

i iens is of s ervice to the an im a l as a m e an s o f co n ce a l
pp
me n t T h e frog 1 5 less co n spicuous in a d ark e nviro n m e n t
.
,

wh e n its ski n assu mes a d arker h ue and whe n in t he gr ass ,

o r weeds its gre e n color a tio n serves the s ame purpos e Th e .

mech an is m of color ch anges an d the v arious stimuli by ,

m e an s o f whic h they are set u p will be tre ated of in th e ,

descrip tio n of t h e S ki n .

E nemies — As frogs are amo ng th e mos t d efe nseless of

an i m als t hey fal l an e asy prey t o a v ariety of c a r n ivorous
,

cre atures w ho devour th em in gre a t n u mbers F irst o f

, .

these e n e mi es in o rd er of destr u c t ive n ess is doubtless to be

cou n ted m an who o n accou n t o f his fo nd n ess for frogs
, ,
’

legs t o say n othi n g o f hi s sci e n tific curi o si ty h as al most

, ,

extermi n ated so me speci es in m any loc ali ties I t i s in t h e .

breedi n g period in th e e arly spri n g t h a t t h e d es t ructio n o f

frogs is gre ates t si n c e the an i m als th e n a ppe ar most abu n
,

d au tly an d are most e asily c augh t W ater r ats an d sku nks .

c atch m any frogs t h e l at t er in E urop e accordi ng to F isc h er

, ,

S ig w art hu n t i n g ou t the frogs from t h e hollows in w hich

,

they ofte n co n gregate duri n g t h e wi n ter There are a n u m .

ber of bird s which prey upo n frogs such as cran es h ero n s , , ,

an d crows ; b u t their gre a t est e n e mies n ext t o m an are th e , ,

v ariou s speci es o f s n akes of wh ic h accordi ng t o F isch er

, ,

S ig w art they h ave an i n t e n se i n sti n c t ive fe ar

,
W he n in the .

w a t er t hey m ay also fall a prey to the l arger sp eci es o f

tur t les .

I n E urope seve ral fishes such as th e l arger h erri n g an d

,

tr out p rey u p o n frogs ; and s m aller fishes are very dest ru c

, ,

t iYe t o the t a dpol es .

To a c ert ai n exte n t fr ogs are p r eyed u po n by oth e r m em

11 HA B I T S A ND N AT U R A L H IS T ORY OF TH E FRO G “3 9

bers of their o w n cl ass Th e l arge C ryptobr anchus devours

.

frogs an d eve n to ads I h ave sever al tim es fou n d l arge

,
.

bullfrogs with R a n a p zf z ezz s in th eir s t om achs an d it fre

'

que n tly h appe n s t h a t sm all i n dividu als fall vic tim s to l ar ge r

m embers of t hei r o w n speci es .

A mo n g the i nver t ebr at es there are few species th a t

actively prey u po n the frog i f we exclud e th ose for m s whic h

a re p ar a sitic M any aqu atic bugs such as B elostom a B e na

.
, ,

cu s Z aith a Ran atr a and eve n t h e sm all b ack s w i m m ers

, , ,
-

No to n ect a c atch th e you ng t adpoles an d suck o u t t h ei r

,

blood W ater be etles such as D y t iscus an d th e ste althy

.
, ,

l arv ae of t h e dr ago n flies m ake u se o f the s am e source of

food M ort ali t y a m o n g the t ad pole s is n atur ally high as
.
,

th ey are p r eyed u po n by m an y forms w hich are u nable to

cope with the adul t frog Water fo w l fish es an d aqu atic
.
, ,

i n sects preve n t the gre at m aj ority from re achi ng m a turi t y

an d the you n g frog is exposed to m an y d an gers fro m which

older an d l arger i n dividu als are exe mp t I t i s very proba .

b le th at but a sm all p art o f the favored few who re ach

m at urity an d p erp e tu ate th eir ki nd di e of old age The .

sto m ach of so m e l arge r an i m al forms the i n evi t able d esti n a

tio n o f all but a sm all p er ce n t o f th e p r oduct o f any
brood .

The crayfish is ofte n fou n d d evouri ng the d e ad bodies o f

frogs and it is no t i mprob able th at occ asio n ally it m ay s ap
,

ture an u n w ary speci men alive ; b u t for th e m ost p art i t , ,

p rob ably m akes use of frogs killed by so m e oth er m e an s .

C ert ai n speci es of G lossipho ni a ( C lepsi n e) amo ng the leeches , ,

live upo n frogs and turtl es bu t they d o no t r equ ire a very

l u ge qu an ti t y o f food si n ce o n e m e al m ay su ffice to ke ep
,

t hem alive for over a ye ar L ike highe r an i m als frogs are

.
,

a t t acked by mosquitoes but it is u n ce r t ai n h o w much in c o n

,

ven ien c e arises fro m this source .

40 TH E BIOLO G Y OF TH E F ROG CH A P .

P ar asit es — Th e fr og like m ost o f th e highe r an im als

, ,

serves as the host of a l arge n u mber o f p arasi tic forms b e ,

lo ngi n g both to th e an i m al an d the vege t able ki ngdom s .

Th e leech es m e n tio n ed in th e previous sectio n might almost

be s aid to be p arasitic si nc e they rem ai n a tt ach ed to the
,

frog for a lo n g period Th e l arv ae of b lo w flies ( C alliphor a

.
,

Lu cili a) someti m es i n fest the i n testi n e o f frogs ; bu t they

usu ally prove a gre ater p est to to ads Th e fem ale l ays it s .

eggs in th e n ost r ils of the to ad an d the l arv ae th at h atch o u t

,

feed u po n th e m embr an es o f the n as al c avity an d m ay ,

work th ei r w ay i n to the br ai n and so m eti me s the eyes of

thei r host I h ave fou n d n o record of their occurre n ce in
.

the n as al c avi t ies o f frogs al though i t is n o t improb able th at

,

they are occ asio n ally fou n d t he r e .

O f th e sever al speci es of Ne m atod es fou n d in th e frog ,

Rna é a o n em a n zgr o z/en osa wh ich occurs in sever al E urop e an

’
,

species is p erh aps th e best k n ow n si n ce its li fe h istory

, , , ,

p r ese n ts sever al exceptio n al an d i n teresti n g fe atures A .

ki n d of alter na tio n o f ge n er atio n s occur s in this sp ecies ,

t he r e bei n g a free form livi n g outside th e body an d a p ar a ,

sit ic form w hich is usu ally fou n d in the lu n gs Th e l atte r is .

herm aphrodi t ic an d produc es eggs w hich give rise to rh ab

,

dit ifo r m embryos which p ass i n to th e a lim e n t ary c an al an d

t h e n ce ou tside th e body Th ese e mbryos d evelop i n to the
.

free form which co n sists of bo th m ales an d fem ales Th e

, .

eggs produced by th e fe m ale are fe r tilized an d develop

wi t hi n h e r body H ere the e mbryos live an d gro w by
.

d evouri n g th e i n ter n al org an s o f th eir m o t h er aft er which ,

th e you n g m atricid es m ake thei r esc ape i n to th e w ater .

Whe n o pportu n i t y offers th ey crawl i n to t h e lu ngs of a frog

,

an d th ere d evelop i n to t h e p ar asi t i c h erm a ph rodi t ic form .

An allied s p ecies A sea r z s en fo m e/a s L eidy occu r s in thi s

’

, ,

c ou n t r y in th e lu n gs of R a n a p zjmen s
'

.
42 TH E BIOLOGY OF TH E F RO G CH A P .

bee n re ce n tly fou n d in so m e Am eric an sp ecies by Nic k erso n .

The peculi ar ge nus Po lysto m u m ch aracterized by h avi ng a ,

circle of several disti n ct suckers at t h e posterio r end o f th e

body i s o ft e n fou n d in t h e uri nary bl adder o f frogs M o n o
,
.

s to m n m or n a ra m L eidy occurs in th e body c avity an d ,

H o/os fo m z/m n z t z a n m L eidy an d s /oa z sen s s n é ela va tn s

' ' '
’ ’

D i es in the i n testi n e of Ra na p zp z en s M o n as zom a m

’ ’
‘
. .

z iz en m i n fests th e lu n gs of R a n a es en len la an d Ty lo ael

’
'

el/
'

p ,

h as be e n fou n d in the n eur al c a n al o f th e

'

p /
z z s

s a m e speci es Coao n oeep na la s m a la é zlz s D i es T elr a eozy /e

’ ’
’ ‘
. .
,

lz n a Ru d and sever al sp eci es of D zs zo m n m occu r in

’

ery s z a /
’

‘ ‘
.
,

an e n cys t ed st ate in v arious p arts o f the body o f sever al

E u r ope an frogs .

O f P rotozo an p ar asites th ere are seve ral n o teworth y forms

a mo n g the I n fusori a Th e l arge mul t i nucle ate O p ali na co m .

m o n ly occurs in th e in t est in e e sp ec ially the rectu m 0

‘
. .
,

r a na r u m an d
,
a re fou n d in Ran a ,

while other species i n fest o ther ge n er a o f A mph ibi a Thre e .

species of B al an tidiu m viz en foz oo n E hr e/o ng a f a m S tei n ,

.
, ,

an d a n oa en z S t ei n i n h abi t the ali m e n t ar y c anal of Rana th e

’
’
’

, ,

first speci es livi ng c hie fly in th e rectu m Ny efo l/z er ns eo a r zl .

’

f o r m z s S tei n an d B a r sa m a z n fes zz n a lzlr E h r are i n t esti n al

' ' ’ ’
‘
.

p ar asites ; the l atte r w as fou n d by L eidy in the i n t esti n e of

R a n a p ip z en s
’

O f the protozo a th ere are two sp eci es o f B odo

flagellat e
which i n fest the i n te sti n e T ryp a n oso ma s a ng z/z n z} G ruby
'

occurs in the blood o f several E urope an species an d I h ave ,

fo u n d a closely alli ed if n o t ide n tic al species in th e blood o f

R a n a p ip z en s
’

Th e l argest n u mber o f protozo an p ar asites of th e frog fall

wi t hi n th e exclusively p ar asi t ic cl ass of S porozo a The S poro .

z o a i n festi n g t h e Am eric an species o f Amph ibi a h ave received

little atte n tio n b u t i t is prob able th at a l arge proportio n of

,
11 HA BI T S A ND N AT UR A L H IS T O RY OF TH E F RO G 43

the m will prove ide n tic al with sp ecies fou n d in E urope .

D escrip t io n s o f most of these forms m ay be fou n d in L abb é s ’

S porozo a an d in th e excelle n t ch apte r o n th e S porozo a by

”
,

M i n chi n in L anke st er s Tre at ise o n Z o Olo gy fro m whic h

’ ”
,

th e followi n g lis t o f species has m ai n ly bee n compiled Th e .

order C occidiide a is represe n ted by H y a/ok/oss z a

'

fou n d in the e n velope o f th e kid n ey and th e i n tes t i n al p ar a ,

si t es rece n tly d escribed by L aver an

an d M es n il a n d Co eez a z n m ra na r u m L abb e M oly é az s
' '

’
’
’
.
,

E n fz z which occurs like th e two precedi n g speci es in R a n a

'

, , ,

is doub t fully u n ited wi t h Corez a z n m r a na r u m by

' '

’
es cu len t a ,

M i n chi n w h o reg ards th e K a ry ap /z ag a s r a n a r u m of L abb e as

,

also bu t an o t her form o f this s a m e species .

The blood p ar asites belo ngi ng to th e H aemosporidi a are

r eprese n ted by at le ast two speci es whic h i n fest frogs .

L a nkes fr e/Za r a na r u m L an k ) an d a n alli ed

( .

species L m on zlz s occur in the red and white blood

’ ’

, .
,

corpuscles of R a n a esen/en fa The forms d escribed as .

H aemog r eg a r z n a m ag n a an d Cy ta m ceé a ea r ter gf er a L abb eare

' '

co n sidered by M i n chi n as an om alous forms o f o n e o f th e

speci es o f L ank est rella D a ezj lo so m a [Z a ner a n ia ] r a n a .
f

rum D sp len nen s Labb e) is c o n sidered by H i n tze as bu t

.
’

o n e form o f L a n kes lr e/ [a r a n a r n m .

O f t h e M yxosporidi a L ep lo lner a O lz lma e/z er z and L r a n a ,

'

occur in th e u ri niferous tubules of R a n a temp o r a r z a an d

'

R es en /en fa and P /ez s z op no r a D a n z lew séy z in the muscles of

' ' ’
‘
.
,

K a fj Tos which is st ated to occu r I n G i glio

'

a mc
’
eé a r en z s ,

th e re nal epi theliu m of bo t h t h e frog an d the m ouse is a .

,

sporozo an o f u n cer t ai n system atic posi t io n .

O f pl an t p arasites there is a species of S aprolegni a which

,

sometim es at t acks t h e ski n of Ra n a p rpz en r an d prob ably of

’ ’

oth e r speci es of frogs I t forms l arge li gh t colored blotches .

,
-
44 TH E B IO LO G Y OF TH E F RO G CH A P .

which m ay spre ad over a co nsiderable p art o f the body .

Ne c t u ru s and oth e r amphibi an s are li able to a tt acks fro m

th e s a m e fu ngus E i d am has d escribed a sp eci es of fu n gus
1
.
,

B ar z a z oé elar r a n a r u m whi ch i n h abits th e ali me n t ary c an al

' '
’
,

of Ra n a esczz /en fa a n d R oxy r fi z n a

’

. .

Times and P laces of B reeding — Th e b r e ed i n g period

of frogs i s in the e arly spri ng soo n a ft er the an i m als h ave ,

eme rged from their wi n ter qu arters As in most amphibi an s .

,

th e eggs are l aid in the w ater usu ally a mo n g the v ege ,

t at io n n e ar th e sh ore a n d receive n o a tte n tio n fro m the

,

p are n ts after th ey h ave bee n deposi ted Whe n the breedi ng .

se aso n is over t h e frogs sc at t e r and resu m e an active pred a

,

tory life Kil n exceptio nal c ases however th ey h ave bee n

.
, ,

k n ow n to cr awl b ack i n to the mu d aft er th e bre edi n g s e aso n

is over and resu me for a ti me thei r wi n ter sleep At D orp at
,
.
,

M arquis observed th at in o n e ye ar R a n a f n s ea i mmedi at ely ,

a fte r the breedi n g se aso n cr aw led b ack i n to the mu d a n d,

re m ai n ed two weeks be fore ag ai n e mergi n g .

The co m me n ce m e n t of the breedi n g d epe n ds in a gre at

me asure upo n the temperature The breedi n g of Ra n a .

m usu lly o ccu rs e r Arbor th e l ter h alf of

'

pj z en s a n a A nn in ,
at ,

April o r th e first p art o f M ay I n ge n eral a l ate spri ng .

,

d el ays th e ti me of bre edi ng an d w ar m we ather o n th e other , ,

h an d h aste n s i t
, F isch er S ig w art w ho for fourtee n ye ars
.
-

h as c arri ed o n an exte n sive series o f observ atio n s u po n Ra n a

fu sea fou n d th at th e differe n ce be twe e n t h e e arliest and
,

t h e l atest p e r iod w as o n e mo n th ( fro m F ebru ary 2 6 to

M arch 2 6 ) in B arm o o sw eih er an d o n e an d a h alf mo n ths ,

( fro m F ebru ary 2 6 to April 1

5) in H o ld e n w e ih e r Th e .

breedi ng peri od d ep e nds to a c ert ai n exte n t upo n loc al

co nditio n s I n sh allo w po n ds which are exposed to th e
.

su n an d wh ere co n sequ e n tly the w ater

, beco m es w armed
1 E i dam , Co h sB i g
n
'
e t ra e z ur B i l gi
o o e der Pflan z e n , Ed 4 p
.
, . 1 81 .
11 HA BI T S A ND N AT U RA L H IST O RY OF TH E FROG 45

e arly in th e se aso n frogs bre ed much e arlier th an in w ater

,

w hich o n a ccou n t o f i ts dep th or t h e l a ck of su n shi n e is

h e ated o n ly very slowly I n m asses of w at er fe d by cool

.

spri ngs th e breedi n g of frogs an d of oth er forms of life is

muc h del ayed .

Th e differe n t specie s of frogs breed at differe n t tim es .

I n G erm any R a n a f ns ea breeds ge n er ally in M arch In .

Ra na a r e a /is th e associ at io n of th e sexes co m me n ces two

or t hree weeks l ater but t h e ti m e o f t h e egg l ayi n g n e arly
,

coi n cides acco rdi n g to B or n w i th t h at o f t h e precedi n g

, ,

species but i ts duratio n i s som ewh at lo n ger R a n a esen/en z a

,
.
‘

l ays eggs o n ly in M ay or J u n e .

The breedi n g period o f a species n atur ally v aries with th e

l ati tude co mi n g o n l ater as we p ass n orth w ard an d e arlier
,

as w e p ass sou th Thu s Ra n a lemp or a r z a whose breedi n g

’

.
,

se aso n in E ngl an d an d m iddle E urop e i s in M a rch does n o t ,

bre ed u n til M ay in Norw ay but in souther n cou n tries i t m ay ,

b r e ed eve n as e arly as J anu ary .

D r M org an h as recorded the results of four ye ars observ a

.
l ’

tio n s o f sever al A meric an species of fr ogs in t h e vici n ity o f

B altimore M aryl an d ,
.

“The first frogs to la an d a mo n g th e very first A er z s

(
’

y ,

r Z/n s
g y exc epted ) to a pp e a r ar e th e wood frogs (Ra n a ,

A few w arm days in e arly spri n g su ffi ce to bri n g

th em out The followi n g records give a ge n er al id e a of th e
.

tim e F ebru ary 2 3 1 8 9 1 an d M arch 8 t h 9 t h an d 1 0 t h

.
, , , , ,

1 880 . The eggs o f these h ad bee n l aid several days Th e .

egg bu n ch es are fou n d in s m all pools o n t h e edges of weeds ,

ge n e r ally amo n g th e low hills an d are ofte n s tu ck t o t wigs ,

o f bush es Th e bu n ches are ge n erally large fo ur to six

.
,

i n ch es in di ameter an d co n t ai n very m any good sized eggs

,
-

I n the s a m e pools it i s qui te usu al to fi n d the firmer egg

1 Mor g an , Am . Nat .
, V ol
. 2
5 .
46 TH E B IOLOGY OF TH E FROG CH A P .

bu n ches of Amblys t o m a for this U rodele also l ays i ts eggs

,

very e arly .

S o m ewh at l a ter two species o f tree frogs appe ar in th e

sm all p o ols in th e woods ge n er ally in quite sm all and there
, , ,

fore duri n g t h e day ofte n quite w arm puddles ; so me ti mes

, ,

in t h e s a me pools as the wood frogs o ft e n er in t h e di t ch es ,

b y t h e side o f the ro ad These two frogs are [Jy /a p ie/leer

.

z n z z an d Ch o mp / 21721 5 l r z r er z a f n s The eggs o f thes e speci es

’ ' ' ' '

g .

a r e very si mil ar a n d I k n o w o f n o cer t ai n m e t hod of disti n

,

g u is hin
g th e o n e fro m th e o t her Th e bu n ch es a r e sm.all ,

a t t ached t o bits o f gr a ss or li e sim ply o n t h e bo tto m a n d

, ,

e ach bu n ch co n t ai n s fro m five o r six t o fift ee n or t we n ty

eggs I h ave the followi n g record of the t i mes at which t h e
.

eggs were fou n d : H yl a M arch 9 1 0 1 3 ; April 5 1 8 9 0

'

.
, , , ,

Cho ro p hilus F ebru ary 2 3 1 8 9 1

,
M arc h 1 3 a nd 24 1 8 9 0
, ,
.

“T he eggs o f R a n a Iz a /eez n a are fou n d still l ater som e

'

ti mes in th e s ame loc al iti es as th e wood frogs o fte n er in pools ,

in the ope n grou n d quite aw ay fro m t h e woods Th e follo w .

in g are th e records : M arch 2 5 April 1 5 1 8 9 0 ,

E ggs of ,
.

a o [en tzgz n os n s w ere repor t e d from th e s am e r egio n o n

'

April 1 4 1 8 9 0 a n d April 5 an d 6 1 8 91
, , ,
.

D urat ion of t h e B reeding P er iod The dur at io n o f .

the br eedi n g period like the t i me o f i ts first appe ar an c e is

, ,

very d epe n de n t upo n exter n al co n di t io n s especi ally temper a ,

ture The peri od of copul a tio n or th e t i m e duri n g w hich

.
,

m al es m ay be fou n d cl aspi n g t h e fem al es o fte n co n sider ably ,

exceeds t h e period betwee n the firs t an d l ast depositio n o f

eggs This is due to th e fa ct th a t th e m ales seize th e
.

fe m ales oft e n sever al d ays before th e fi rs t eggs are l aid .

Accordi ng t o F isc her S igw art s obser vatio n s of R a n a f nsea

-
’
,

t h e fe m ales m ay lay eggs i f t h e we a ther is w arm t hree days

, ,

aft er t h e begi nn i n g of copul a t io n ; b u t in cold we a th er the

l ayi ng m ay be de fe rr ed fo r t we n ty or t hir ty d ays The d ura .

11 HAB I T S A ND N AT U RA L H IST O RY OF TH E FROG 47

tio n o f c o pul atio n in frogs kep t in th e terrariu m w as fou n d

t o v ary be t wee n six an d twe n ty th ree d ays -
Th e less v ari a
.

b ility o f the breedi n g period in this c as e is doub t less du e t o

th e more u n i for m co n di t io n s u nder which th e an i m als are
kep t The period be twee n th e appe ar an ce of the first egg
.

m ass an d the tim e wh e n all th e fe m ales h ave ex t ruded their

eggs w as fou n d to v ary in differe n t loc alities bet w e e n four ,

a n d twe n ty seve n d ays

-
accordi n g to th e temp er a ture ; in
,

t h e terr a riu m a v ari at io n be t w e e n t en an d t we n ty eight d ays -

w as obs erved Accordi ng to P fltiger th e l ayi n g p eriod in

.
,

Ra n a esrn len za l as t s o n ly two d ays ; I t i s qu i t e prob abl e

‘

t h at a si n gle copul at io n ex te n ds throu gh the whole breedi n g

s e aso n I n R a n a f n sea Pfltiger h as observed th at a m al e

.
‘

m ay cl asp a fem ale for seve ral w eeks an d in R a n a es cu len ta

,

t h e s a me observer h as record ed an e mbr ac e which l a ste d a

mo n th S t ei n ach records a still lo n ge r period for R a n a

.

f a sea ,
th e p a irs rem ai n i n g u n ited for a s much a s s eve n

weeks I n these c ases t he we at h er w as cool o t her w ise th e

.
,

disch arge o f t h e sexu al produc t s would h ave occurred much

more qu ickly
Copu lat ion —I n copul atio n th e m ale cl asps t h e fe m al e
.

just behi n d h er fore legs w here h e h an gs tightly an d m ai n

,

t a i n s h is hold persiste nt ly agai n st all efforts to dislodge hi m .

O ft e n t he t hu mbs ar e i n terlocked to i ncre ase th e firm n es s

o f his hold .Th e body of the fem ale is very much co m
pressed in co n sequ e n ce o f this but it app are n t ly c auses h er
,

li t tl e i n co nve n ie n ce Th e l abor of locomo t io n o f t h e p air

.

falls m ai n ly upo n the fe m ale th e m ale exer t i ng hi msel f o n ly

,

occ asio n ally for t he m ai n te nan ce of equilibriu m Wi t h t h e .

exceptio n o f t he effort n ecess ary to e n able hi m to ret a i n his

hold the m ale duri n g copul at io n is si n gul a rly i nactive an d
, ,

will e n dure u n favor able co n di tio n s r at h er t h an m ake an y effort

to seek a be t ter situ a tio n S om e t i m es sever al m ales will be
.
48 TH E BIOLOGY O F TH E FROG CH AP
.

fou n d cl aspi ng d iffere n t p arts o f th e body o f o n e fem al e ,

formi ng m asses B ega t t u n gsklu m p e n

,
in th e mids t of ,
”

which th e fem al e c an sc arcely be see n M ales will o fte n .

cl asp fe m ales which are d e ad o r fem ales of an other species, ,

or eve n to ads The m ales o f Ra n a f nsea h ave bee n observe d

.

cl aspi ng c arp an d bei ng c arried arou n d by th e fi sh which ,

were u n able to divest th e mselves o f their burde n Th e eyes .

o f m an y of these fish es are destroyed by th e thu mbs of th e

cl aspi ng frogs M ales will u su ally cl asp o n e s fi n ger o r al most
.
’
,

an y obj ec t t h ey c an seize although th ey w ill n o t as a rul e

,

re t ai n hold of obj ects o ther th an fem ale fr o gs for a v ery

lo ng ti me The cl aspi n g i n sti n ct to a l arge exte n t over
.

co mes fe ar A frog which will m ake viole n t e fforts to esc ap e

.

fro m th e h an ds will o ft e n hold to o n e s fi nger an d e n tirely ’

desist fro m i ts efforts to esc ape duri ng the p eriod of copul a

t io n The cl aspi n g i n sti n ct i s so stro ng th at th e an i m al m ay
.

be severely i nj ured without showi ng any dimi nu tio n o f i ts

a rdor . The body o f a m ale frog m ay eve n be cut in two in
the middl e an d t h e fore p art w ill still c li n g te naciou sly to the
fem ale for hours o n ly desis t i ng wh e n th e cre ature becom es
,

so we ak fro m loss o f blo od th at i t c an n o lo nger re t ai n its

posi t io n .

E gg L ayin g an d F er t iliz at ion — Th e extrusio n of eggs .

usu ally occurs o nly aft er t h e m ale h as cl aspe d t he fem ale

fo r sever al d ays T h e eggs whic h are disch arge d fro m the
.
,

o v aries r a t h er slo w ly a r e c o n veyed fro m the body c avity i n to

,

t h e oviduct th rough which t h ey p ass i n to t he u t eri wh ere

, ,

t hey fi n ally all coll ect di s t e n di n g th e thi n w alls o f th ese

,

org an s t o an e n orm o us d egre e As S p all an z an i discovered .

lo n g ago fe m al es killed in th e firs t p ar t o f the breedi ng sea

,

so n h ave most of the eggs s t ill in t h e o v ary Later m an y .

eggs ar e fo u n d in thei r p ass age d o w n t h e oviduc t a l though ,

they are n o t ofte n m et with in the bod y c avi ty F em ales .

5 0 TH E BIOLOGY OF TH E F ROG CH A P .

o ver m asses o f eggs Th e super n u mer ary m al es th erefore

.
, ,

pl ay a p ar t in fer t ilizi n g th e eggs as well as those which

,

h ave succe eded in obt ai n i n g a m ate .

Wh at th e sti mul u s is t h at pro mpts the m al e to disch arge

his sper m at the s a me t i m e the eggs are extrud ed from t h e
fem ale is n o t al t ogethe r evi de n t t h e s a me p r obl em prese n ts
i tself in the c ase o f fish es .

Con g r egat ing at B reedin g G r oun ds D u r i ng th e bre ed

.

in g se aso n frogs u su ally co ngreg at e at c ert ai n poi n t s in sh al

low w ate r in co n sider abl e n u mbers At this ti m e t h e r e
.

seems to be m an ifested a greg ariou s ness which d oes n o t .

a ppe ar u n der ordi n ary circu ms tan ces and whi ch is n o t

,

e n tirely accou n ted for by th e te n de n cy o f t h e an i m als to

seek a si mil ar h abi t at for breedi n g Accordi n g to F isc h er
.

S igw ar t s obse rv a tio n s o n Ra n a f n sea i f o n ly a few p a irs

’

occur in any loc ali t y th ey ge t as clo sely togeth er as possibl e

, ,

a nd t heir egg m ass es for m a l m ost a co n ti n uous sh eet The .

l ayi n g grou n ds are the sce n es of li vely ac tivi t y Alles h astet,

u n d dr an gt . The super n u merary m al es c rawl over an d

work through the m asses of eggs an d accordi ng to F isc h er
,

S igw art effect t h e fertiliz a tio n of th e o va w hich m ay n o t

,

h ave bee n re ached by sperm atozo a at t he time o f th eir

disch arge .

Wh e n th e sexu al products are disch arged th e frogs go ,

b ack u po n th e l an d an d sc at ter in all di rec t i o n s M ost .

frogs le ave th e breedi ng grou n ds a t n e arly t h e s am e ti m e .

At o n e day a pl ace m ay be t ee mi n g wi th th ese cre atures

while o n th e followi ng day n o t a si ngle i n dividu al c an be
fou n d H e n ce forth the frog is a solit ary an i m al h avi n g lost
.
,

all its sexu al i n s t i n cts a n d soci al proclivi t ies I t le ads i ts

.

li fe as i f n o o t he r m ember of its species were in exis t e n ce .

E g g L ay ing w it h ou t t h e P r esence of t h e M a e l — Th e
ques t i o n w h e t h er o r no t th e fem ale frog W ill lay eggs with o ut
11 HA B I T S A ND N AT U R AL H IS T O RY O F TH E F RO G 5 1

the embr ace of the m ale h as be e n i n vestig ated by Nu ssb au m l

in Ra n a f nsea . F em ales w ere isol at ed soo n a ft er they c a m e

o u t fro m th eir w i n ter qu ar t ers w h e n t h e eggs were s t ill in ,

the ov ary Th e eggs were fou n d t o le ave th e ov ary p ass

.
,

dow n the oviduct an d collect in t he uterus as they do u n der

,

n orm al co n di t io n s They w ere n o t extru ded from t h e u terus

.

all a t o n ce as they are in copul at io n but were p assed o u t ,

sl o w ly a few at a t i m e so m e eggs oft e n bei n g ret ai n ed u n t il

, ,

co n sider ably a ft er the breedi ng s e aso n I n a fem ale killed .

l a te in t h e su mmer o n e u t erus w as fou n d to co n t ai n a l arge

a mou n t of j elly but th e eggs th em selves h ad broke n dow n
,

an d dis appe ared .

E gg l ayi ng in R a n a esr n len fa is app are n tly more depe n de n t

upo n exter n al co n ditio n s While R f nsea re adily l ays it s . .

eggs whe n in c ap t i vi t y esen /en /a do es so o n ly if th e eggs

,

h ave accu mul ated in the uteri at th e ti m e of c ap ture .

Whe n the fe m ales are t ake n e arlier t h e eggs t h at are in t h e ,

ov a r y are n o t extruded bu t are ret ai n ed th ere u n t il th ey are

,

fi nally resorbed .

P r op ort ions of t h e S exes — A m al e frog as a rule p airs , ,

wi t h o nly o n e fem ale duri ng t h e breedi ng se aso n and it i s ,

adv an t ageous to the species t h a t th e s exes should be n e arly

equ al in n u mbers This rel atio n is in fac t u su ally fo u n d t o

.
, ,

ob t ai n amo n g adult frogs B or n in s t udyi n g t h e proportio n

2
.
,

o f th e sexes in you ng frogs c a m e t o t he co n clus io n th at t he ,

fem al es oft e n gre at ly prepo n der at e o ver the m ales I n o n e .

bro od the fem ales w ere fo u nd t o c o n s t i t u t e n i n ety fi ve per -

ce n t of th e tot al nu mber Pfliiger an d G riesh ei m w h o in

3
.
4
,

vest igat ed the subj ect wi t h co n sider able thorough n ess fou n d ,

1 A r c/z m z é V o l 46 1 8 9 5
’

. . .
, .

2 B o r n B r es l a u er ar ts ] Z ez t sek 1 8 8 1
. .
, ,

3 Bd 8 82
Pfliig e r ,
. 29 , 1 .

4 Gr i h im
es e , Bd . 26 , 1 88 1 .
52 TH E BIO L OG Y O F TH E F RO G CH A P .

th at in you ng frogs it i s o fte n v ery di fficul t to disti nguish

th e s exes Th e go n ads o f the m ales frequ e n tly co n t ai n
.

n u m erou s egg fo llicles an d h ave th e ge n er al appe ar an ce o f

O v a ries an d m an y frogs which would n atur ally be t ake n fo r

,

fem ales S how o nly m al e ch aracters at a l ater peri od o f devel

o p m en t E s t i m ates b ased o n th e a ppe ar an c e of th e sex
.

org an s of frogs duri ng the fi rst ye ar of thei r li fe g ave a very

l arge perce n t age o f fe m ales This p erce n t age w as fou n d to
.

v ary gre atly in lots o f frogs t ake n at differe n t loc ali t ies .

The degree of develop me n t of the se x org an s does n o t

closely correspo n d wi th t he develop me n t of the other p arts
of th e body and t he r e is co n sequ e n t ly a gre at v ari atio n in
,

the app e ar an ce o f th ese org an s in speci me n s of th e s a me

age . A s there is n o evide n ce t h at th e m ort ali t y of the
you ng fe m al es gre at ly exce eds t h a t o f th e you n g m ales an d ,

as the proportio n of t h e ad u l ts of th e t w o sexes is n e arly th e

s am e i t i s prob able th at m an y of the you n g frogs which

,

would ordi n arily be di ag nosed as fe m al es are j uve n ile “

h erm aphrodites in whic h th e fe m ale ch ar acters p redo mi
n ate for a tim e bu t whic h l ater develop i n to m ales This .

co n clusio n is su ppor t ed b y t h e fact th at t here are m an y

_

c ases in w hich th e sex gl ands are i n term edi ate in app e aran c e
be t w ee n ov ari es and testes E gg follicles are o ft e n fou n d in
.

the t estes of frogs at a l ater p eriod of d evelopme n t .

Copu lat ion in L at e S u mmer F isc h er S igw ar t h as o b

.
-

served t h a t in July an d Augus t th e sexu al i nsti n c t of t h e

m ales k ept in terr ari a ofte n ass er t s i tself a seco nd time ,

esp eci ally if they h ave bee n w ell fed an d are in good co ndi
tio n No sexu al products are extruded a t this ti me fro m
.

either sex al t hough t h e m ales show a stro ng proclivi t y t o

,

cl asp t h e fe m ale an d will ev e n cl asp oth er m ales A c ase i s .

rel at ed of a m ale who observed a fem ale which had seized

an e arthworm an d in attem pti ng to sh are th e mo r sel cl am
,
II HA BI T S AND N AT UR A L H IS T O R Y OF TH E F RO G 53

bered over h er b ac k w h e n the cl aspi n g i nsti n c t sudde n ly took

,

possessio n o f hi m an d h e rem ai n ed cl aspi n g h er bo dy

, .

L ater in th e s e aso n this c laspi ng i n sti n ct dis appe ars .

H ib ern at ion I n th e l a te fall frogs bet ak e themselves

.

to w ater an d bury th emselves in t he mu d ou t o f r e ach of

frost H ere th ey li e in a dorm an t co n ditio n u n til th e n ex t
.

spri ng The ge n er al vi t al activi t ies of t h e an i m al ru n dow n

.

so low th at li tt le expe n diture o f e n ergy is required t o m ai nt ai n

life There is n eed therefore for o n ly a sm all a mou n t of
.
, ,

o xyge n an d s k i n respir atio n t he n su ffi ces

,
D uri ng t h e .

w hole wi n ter th e frog does n o t bre a t h e air wi t h t h e lu n gs .

Th e temper ature o f th e body si n ks u n til it is o n ly a few

degrees above t h at of t h e surrou n di ng m edium A s th e .

frog t akes n o food duri n g this ti m e i t mu st keep u p its vi t al

,

activi ty at the expe n se o f m ateri al stored in it s tissues I ts .

temper ature eve n i f o n ly a little above t h at of its surrou n d

,

i n gs requi r es the u se of a cert ai n amou n t o f combus t ible

,

m at eri al fo r i ts support D uri n g the su mmer t h e frog feed s

.

vor aciously an d whe n cold we a th er e nsues its system i s

, ,

stored w i t h a rich supply o f food m ateri al which is gr adu ally

expe n ded through t h e wi n ter mo n ths This food mus t keep .

the h e ar t be a ti n g and su ppor t the v ariou s ac tivi t ies o f th e

physiologic al m achi nery of the body An d in addi t io n to .

supplyi n g e n ergy for this purpos e i t must afford th e sub ,

st an ce for the grow t h o f th e sex ual products which i n cre ase ,

duri n g the wi n ter at th e exp e n se o f the other p arts of the

org an ism ..

L if e in S ummer — Aft e r the eggs are l aid in t h e e arly

.

spri n g t h e frog le ads an active pred at ory and solit ary life
,
.

Aft e r i ts lo n g wi n ter sl eep an d the expe n diture of subst an ce

a n d e n ergy duri n g th e breedi n g se aso n which closely follows

the aw ake n i ng in t h e spri n g the frog i s n at u r ally in gre a t

,

n eed of food and i t becom es a ve r y vo r aciou s fe e der

, D ur .
54 TH E B IO L O GY OF TH E FRO G CH AP .

ing th e e arly p ar t o f th e su m m er it is bu sily e ng aged in t h e

a tte mp t t o s a tisfy its h u n ger I n midsu mm er whe n th e
.
,

body has co mp e n s ated fo r it s losses t h e frog o fte n be t akes ,

itsel f to a pl ace of co n ce al me n t co mi n g out o n ly a t i n terv als

,

t o ob t ai n foo d This pe r iod of co mp arative i nactivi ty has

.

bee n spoke n of as a su m mer sleep b u t accordi n g to F isc h er

,

S ig w art a tru e su m mer slee p does n o t occur either in frogs

o r to ads There is o nly a perio d of comp ar ative rest aft er
.

the n eed for a l arge a m o u n t of food has ce ased .

I n sum mer th e frog has food in a bu n d an ce an d it i s l ess

, ,

li able to fall a prey to e n emi es th an duri n g the breedi n g

period I t c an b ask in t h e su n seek th e sh ade w he n it is
.
,

t o o w arm or plu n ge i n to t h e w ater to m ois t e n its ski n or

,

cool its body I t i s usu ally o n it s gu ard however ag ai n st

.
, ,

wh a tever e n emi es m ay prey u po n it bu t from m ost of th ese ,

i t freque n tly h as n e ar at h an d a re ady me an s o f esc ape I t .

is fre e to e nj oy life as best an a mphibi an can an d to store ,

u p food fo r p assi n g t h e wi nt er and for t h e rip e ni ng o f t h e

sexu al produc t s for the follo w i ng spri n g To ke ep alive it s .

s t ock a process whi ch i n cide n t ally i mpli es t h e pres erva tio n

,

of it s o w n body is the gre at e n d of t h e frog s exis t e n ce

,
’
.

I n j ur ies P ow er of Regen erat ion — As th e frog i s preyed

u po n by sever al e n e mies specim e n s are o ft e n fou n d in which
,

fi ngers or t oes or som e times t h e e n t ire h an d or foo t are

, ,

missi n g I n most c ases t hey are doub t less i n dividu als which
.

h ave h ad these p arts bi t t e n off a n d were fo r t u nat e t h a t o n ly

a por t io n of t heir body w as left in th e possessio n of t h e

e n emy E ve n severe wou n ds in the frog h e al very re adily

. .

F isch er S ig w art h as observed th at i f a frog i s wou n ded i t

-

be t akes i t self t o th e w ater an d if an i n d ividu al ke eps in t h e

,

w ater duri n g t h e su m m er o n e m ay be pre t ty sure th at it h as

,

received some i nj ury .

T h e power o f rege n er atio n possessed by so m e o f t h e

II HA BIT S A ND N AT UR A L H IS T O RY OF TH E FROG 55

lo w er m p hibi a is very m ark ed Trito n is able to rege n

a .

e rat e its t a il or li mbs or eve n its eye if a sm all portio n

, ,

of t h at o rgan i s le ft B ut in th e frog an d so far as is

.
, ,

k n ow n in th e other A n ur a the power o f rege n er at io n is

, ,

al most e n tire ly los t E ve n in t h e t adpole st age i t is mu ch

.

reduced Th e t adpoles of t h e higher a mphibi a are able t o

.

rege n er ate the t ail i f it is cu t o ff but their power o f rege n

,

cr a ti n g t h e li mbs i s very li mi t ed T he rege n era tio n of the

.

limbs o f t adpoles w as first reco rd ed by S p all an z an i F rai sse .

,

howeve r who cut off t h e li mbs fro m bo t h you n g an d old

,

t adp o les arrived at e n tirely n eg ative results L ater B ar fu rt h

,
.

c arried o n s ever al experi me n ts o n bo th O ld an d you n g t ad

poles and fou n d t h at i f t he legs were c u t off fro m you ng
,

speci me n s th es e org an s would be freque n t ly rege n e r ated ,

al t hough slo w ly .Th e power o f t h e t adpol e to rege n er at e

missi ng li mbs w as fo u n d t o decre as e r apidly with ag e .

E f f ect s of H eat an d Co d l — The frog belo n gs amo n g

.

t hose an i m als w hich are co m mo n ly spoke n of as cold

blooded This expressio n d oubtless t akes it s origi n from

.

t h e fact t h at the temp er a ture of such forms is u su ally low .

The h igher vertebr ates such as t h e m am m als an d birds

, ,

h ave a high bodily t emper atu re an d wh at is most re m arka

, ,

ble the temper ature in most o f th ese keeps n e a rly co n st an t

,

u n der m ost diverse co n di t io n s A bird or a m am m al m ay

.

live in extrem es o f clim ate belo w 4 0 an d 5 0 below z ero ° °

a n d co n si der a bly over 1 0 0 F a nd ye t t h e te m °

F . .
,

p e rat u re of t h e blood will n o t v ary more th an a sm all

fr actio n of a degre e These an im als h ave an almost perfect

.

m ech an ism for t h e regul atio n of the bodily he at Wi th .

a rise o f t e mper a tu re t here is in the m a mm als an i n cre a

,
se ,

of persp ir atio n an d an i n cre ased ev apor at io n fro m the

surface of th e body which te n ds to cool t h e blood Whe n . . .

the te mper ature si nks there is less ev apor atio n fro m the
,
5 6 TH E BIOLO G Y OF TH E FRO G CH A P .

su rface an d th e cold a cts i ndire ct ly as a s t i mulus to i n cre ase

,

met abolism an d c au ses th e combustio n o f t he bodily fuel

t o proce ed at a m ore r api d r ate an d t h ereby to co mpe n s a te

for the loss o f he at by r adi atio n B y vir t ue o f this m e ch an .

i sm the higher an i m als are able to ke ep in an a c t ive c o n

di t i o n in bo th wi n t er an d su m mer Wi t h t h e cold bloode d .
-

a n i m als it is differe n t Th eir temp er a ture rises an d falls

.

in correspo n de n ce wi t h th e te mper at ure of t heir e n viro n

me n t I n the cold th eir me t abolism is slow thei r te m
.
,

e ra t u re ru n s dow n an d co n s eque n t ly th ey beco me slu ggish

p ,

an d i n activ e As it beco mes w arm er thei r te mperat ure

.
,

rises their m e t abolism i n cre ases an d t h ey become more

, ,

a ctiv e an d alert A liz ard whi ch is m ad e al mos t sti ff wh e n

.

the we at her appro ache s th e fre e z i n g poi n t beco mes t h e

most agile o f cre atu res in th e su n sh i n e o f a hot day .

Th e t e mper ature o f the cold blooded an i m als i s n o t how -

ever e n tirely at th e m ercy of th e e n viro n m e n t E v aporatio n

,
.

fro m the sur fa ce o f th e body t e n ds to ke ep th e te mper at ure

o f th e an i m al in w arm we ather belo w t h at of the surrou n d
in g at mosph ere An d as th e we a ther appro aches the
.

fre ezi n g poi n t th e sm all a mo u n t of m et abolis m in th e an i

,
.

m al serves to keep i ts te mper at ure som e w h a t above th at o f

it s e n v iro n me n t .

The effect of high te mp er atures o n th e frog h as be e n

stu di ed by M aurel an d L agriffe At 2 6 t o 3 0 C t he frogs .
l ° °
.

become ac tive a n d res t less A t 3 1 t o 3 3 C th ey show .

° °
.

evide n t sig n s of discom fort F ro m 3 4 t o 3 6 C t h ey j u mp .

° °
.

abou t wildly wi th ou t an y app are n t se n se of directio n At .

a tem per at u re o f 3 7 to 3 9 C t hey lose their se n se o f equ i

° °
.

librium an d i f expose d to a te mper ature o f 3 9 to 4 0 C

,
° °
.

t hey di e I f ho w ever th ey are exposed o nly for a short

.
, ,

t i me at t h e l atter t emper ature t h ey m ay subseque n tly re ,

1 M u l a d L g iff
re an C mp a Sr B l P is t m 5
e, o 9. r en a
’
. o c. aa . ar ,
o . 2, 1 00 .
5 8 TH E BIOLOGY OF TH E FRO G CH A P .

u n til i t froze As a fil m o f ic e beg an to form th e frog

.
,

a tte mpt ed to keep u p a t the su r face of th e w at er bu t i t w a s ,

push ed dow n belo w an d forced to rem ai n th ere u n til froze n

in a soli d c ake o f ice H ere i t w as kept fo r five h ours
while th e su rrou n di n g temper atu re r an ged fr o m —6 to
.
,

C Whe n th e ice w as the n th aw ed the frog w as sti ff

.
,

a n d S ho w e d n o sig n s of life but after an hour and a h alf i t

,

h ad revived .

F rogs fro m w arm cou n tries c ann ot e n du r e so low a te m

p e rat u r e as those fro m higher l atitu d es B u t th e frogs in .

n orther n regio n s ar e oft e n kill ed by cold especi ally duri n g ,

severe wi n ters I f th ey are preve n ted fro m buryi n g de e p

.

e n ough in the mu d the frost m ay overt ake t h e m I n som e

,
.

loc ali t i es the frogs m ay be l argely extermi n at ed duri n g a

p eriod of severe cold so t h at few are fo u n d t h e n ex t spri n g
, .

I n regio n s of high l a ti t ude wh ere the grou n d is p erm an e n t ly

,

froze n b elo w the surface th awi n g ou t o nly for a few fee t

,

duri n g the su m mer frogs do n o t occur si n ce n o m e an s are

, ,

afforded to e sc ape fro m bei n g solidly froze n duri ng t he

wi n ter
.

I t w as observed by Kn au t h e th at th e color o f frogs

exposed t o th e cold bec ame very d ark eve n w he n th ey ,

were pl aced in su n ligh t w hich u nd er n orm al co n di tio n s

,

c auses t h e ski n to assum e a ligh ter hu e .

Ab sorpt ion of W at er — F rogs do n o t dri n k like t h e

.

higher a n i m als but absorb th e w at er they require t h rough

,

t h e ski n. The m e ager an d shriveled co n di t io n o f a fr o g

which has bee n kep t some ti me in dry a ir co n t r as ts m arkedly
wi t h its plu mp appe ar a n ce a ft er it h as j ust bee n t ake n fro m
th e w ater T h e ski n is loosely att ach ed t o t he body an d
.
,

a co n sider able qu an t i t y o f w at er m ay collec t in t h e l a rge

subcu t an eous lym p h sp a c es D o n alds o n fo u n d t h at a grou p

.

of frogs a ft er bei ng kep t in dry air for sever al hours lost -

II HA BI T S A ND N AT U R A L H IST O RY OF TH E FROG 59

1 4 per ce n t o f th eir w eight Whe n pl a ced

. b a ck in th e
w ater ag ai n they reg ai n ed very n e arly their previous weigh t
in twe n ty four ho urs -
B o th t h e loss an d absorp t io n o f
.

w a te r were fou n d to t ake pl ace more r apidly in th e summ er

th an in th e wi n t er An experi me n t by Tow n so n o n a species
.

o f tree frog showed th at a specim e n weighi n g n i n ety fi ve -

gr ai n s i n cre ased in weight by seve nt y six gr ai n s after b ei n g -

kept in the w ater for an h our .

S h eddin g of t h e S k in A t cert ai n periods th e frog c asts

.

off it s cu t i cle or ou t er l ayer of ski n The p art sh ed co n sis t s .

m erely of a very t hi n tr an sp a re n t mem bran e o n ly o n e or

t w o cells t hick This co m es off in l arg e p at ches which m ay
.

be se e n adh eri ng to t h e an i m al here an d t here th e ski n ,

coveri ng th e t o es usu ally co mi ng off l ast Th e fi r st m ol t .

t akes pl ace in th e spri ng at abou t t he t i m e of t h e bre edi n g

se aso n I n R a n a f n sea F isch er S ig w art fou n d th at after the
.
-

first m olt whi ch occurs fro m l ate in F ebru ary to e arly in

,

April a seco n d molt follo w s in t he l atter p ar t o f M ay or th e

,

first p art of Ju n e a third in July and usu ally a four t h in

, ,

August I n colder se aso ns the p eriod o f the m ol t co mes

.

l ater an d the fourth mol t m ay t h e n n o t occur F ive molts

,
.

were n o t observed eve n duri n g the w arm es t su m mers F rogs .

oft e n eat t h eir shed ski n a fte r t h ey h ave rubb ed it off wi t h

th e aid o f their feet Th e s a m e h abi t has bee n observed in
.

to ads an d inthe l arge s al am an d er C ryptobr an chu s .

H ypn ot ism — A frog m ay be throw n i n to th e so c alle d

,

hyp n o t ic st at e in sever al w ays I f i t is s eized in th e h and s

.
,

l aid upo n its b ack a n d held a few mo me n ts u n t il it h as

,

ce ased its struggles it will usu ally rem ai n m o t io n less for a

,

co n si der able ti me som etim es fo r hours The posi tio n t ake n

,
.

is a v ari able o n e There is a te nde n cy to assu me an at titud e

.

such as would be produced i f the moveme n t s of t h e frog

we r e ch eck ed so meti me duri ng its efforts to re gai n an u p
60 THE BIOLO G Y OF TH E FRO G CH A P .

right positio n Accordi n g to V er w o rn t h e muscles i n volved

.
,
1

in t h e righti ng m ove m e n ts are in a s t at e o f t o n ic c o n t rac

tio n a s i f t h es e move me n t s w ere sudd e nly i n hibi t ed Th e .

bre athi n g move m e n t s an d t h e h e ar t b ea t s ar e at first accel

e ra t e d bu t a t a l a ter s t age th eir r a t e falls bel o w t h e n orm al
,

a n d th ere is a d ecre a s ed respo n sive n ess t o ex

( H eu b el
) ,

ter na l sti muli Dif .

fe r e n t frogs v ary
gre at ly as reg ards
bo t h the e ase w i t h
which th ey m ay be
hyp n oti z ed an d t h e ,

dur a tio n of th e hyp

u o t i e st a te I n so m e .

c ases i f a frog i s si m
ply pl aced o n it s b ack
wi t hout bei ng h eld ,

i t m ay becom e hyp
n o t iz e d a ft er it has
righ t ed i tself a few
FI G 7 R
. . t-
in t h
an a c ll den
m e s a n
.
d h
r za
e’
fo r a e so -
a e
“ ;

hy p t ic s t t T t pp figu sh s lo n g tim e in 8 0 11 1 6
,

re ow
‘

no a e . u u er

th p si i
e ossum d h t h b ck is ph ase of t h e pro
t on a e w en e a

ubb d i t h t h fi g T h s m t t i t ud
r e w e n er . e a e a e

is m i t i d h t h f g is p l c d
a n a ne w c ess o f turn i ng ov er
en e ro a e on

b ck is sh l figu
,

it s as ow n in t h e ow er re
f £61 724
a
S p e m m en s
.

( M difi d f m V
,
O
o e ro e rw o r n .
)
ccordi n g es ezz /en fa , a

to V erw o rn w he n l aid o n th eir b acks some t i mes quickly

, ,

draw th e hi n d l egs close to th e body close t h eir eyes an d , ,

lie wi t h t h ei r muscles in a st te o f o i c c tr c io
a t n o n a t n — a ,

co n di t io n w hich suggests the d e ath feig ni ng of cer t ai n

i n sects .

To n ic co n t r ac t i o n s o f di ffere n t p arts of t h e body m ay

1 V e rw o rn , Die sg
o e n an n t e H yp n o se de r Th i e re , 1 8 98 .
II HA BI T S A ND N AT U R A L H I ST O RY OF TH E FROG 61

ofte n be i n duced by rubbi ng th e b ack an d sid es I f th e .

frogs are in a n orm al resti n g p osi t i o n t hey frequ e n tly r aise ,

themselves up o n thei r legs an d re m ai n mo tio n less a nd rigid

for som e tim e I f whe n in this st at e th ey are l aid o n th eir
.

b acks the l egs s t ill re t ai n the s am e at t itude as be fore

, .

F rogs m ay be aw ake n ed fro m their st ate o f hyp n osis by

an y sudde n sti mulus an d th eir recovery i s ofte n i mmedi ate
,
.

Th e durat io n o f this st ate m ay be muc h prolo nged i f all

se n sory impressio n s are so far as possible re moved .

RE F E RE NC E S

S ee e s p ci lly th
e a e w or ks bb t t A ll
of A o ,
en, h m D mé
B o ule n g e r , B re , u

r il et B ib ro n , Diir ig e n , F isch S ig t H
er -
w ar , ay, Ga d L ydig RO l
ow , e , se

vo n Ro s h f
en o , an d S pall z i ci t d i t h
an an e n e re fe re n c s t t h p c di g
e o e re e n

ch ap t er.
62 TH E BIOLOGY OF TH E F ROG CH A P .

C H A PT E R I I I

EX T E RN A L CHA R A CT E RS OF TH E F RO G
IT will be co n ve n i e n t to begi n our s tu dy o f th e frog by a
d escriptio n of th e pri n ci p al ex t er nal fe atures of it s structure .

The fl at te n ed more or less tri angul ar lz ea a is bro adly u n ited ’

to the t r u nk t here bei n g n o regio n th at c an b e p rope r ly

,

c alled a neck The l arge ey es com mo n ly protrude c o n sid

.

e rably bu t can be wi t hdr a w n i n t o th e orbi ts

,
P ress upo n .

o n e of t h e eyes with the fi n gers an d it will be fou n d th a t i t

,

c an be forced i n w ard eve n beyo n d t h e ge n er al surfa ce o f th e

h e ad I f n o w t h e mouth o f t he frog be op e n ed it w ill be

.
,

see n th at the r e is a m arked pro mi n e n ce in th e roo f du e to

the fac t th at th e eye is presse d ag ai n st th e m embr an e li n i n g
t h at portio n of th e c avi ty Th e o r az i o r ey e so elzez o f the
'
‘
.

frog therefore i s n o t s ep ara ted fro m th e m o uth by an y of

, ,

the bo n es o f the S kull w hich is a very d iffere n t co n di t io n

,

fro m wh at w e fi nd for i n st an ce in ou rselves I n the ce n t er

, ,
.

o f t h e eye is a d ark ov al ope n i ng t he p up il which is sur

, ,

rou n ded by a brightly colored ri ng o r z r z s The eye as in

' '
-

.
, ,

ourselves c an be covered by a p air o f ey elz as The npp er ey elz zl

’ '
’
.
, ,

however i s c ap abl e of bu t li t t le move m e nt bu t t h e lo w er lid

, , ,

which is thi n an d more or less tr ansp are n t c an be dr aw n u p ,

so as to cover n e arly t h e whole eye I t will be n o ticed t h at

.

e ach t ime the frog closes i ts eyelids th e eye i s pressed i n t o

t h e he ad a fac t w hich gives t h e w i n ki n g of t h e frog so
,

pecu li ar a n appe ara nce Th e l o w er lid of the frog is n o t

.

q ui t e the s a me org an as t h e lower eyelid of mo st anim als .

I II EX T E RN A L C HA R A CT E RS OF TH E FROG 63

It correspo n d s r at h er to the lower eyeli d proper plus a ,

n z elz l a t z ng m emor a n e The l atter struc t u re in most an i m als

' ' '

in which it occurs is very disti n ct fro m bo t h th e o ther eye

lids I n a bird for i ns tan c e it app e ars as a thi n m embr an e
.
, ,

w hich c an be dr aw n over t h e eye fro m the i nn er an gle of th e

orbi t I n th e frog ho w ever it is situ at ed j ust above th e

.
, ,

lo w er li d of w h ich i t appe ars to form a co n ti nu atio n I t i s

, .

thi n n er an d more tran sp are n t th an the lower lid and sep a

r a ted from i t by a sh allo w groove .

B ehi n d t he eye i s a n e arly circul ar are a cov ered by a

te n s e m em b r an e ,k n ow n as th e ty mp a n ic m em br a ne w hich ,

fo rms t h e coveri n g of the dr u m o f th e ear Ne ar t h e ce nt er .

of thi s membr an e m ay be see n a sm all promi n e n ce c aused

by t he end o f the colu m ella or bo n e which co n n e cts at it s
,

i n n er end wi th a sm all ope n i n g in th e skull which c o m m u

n ic a t e s wi t h th e i n n er e ar When th e ty mp an ic m e mbran e
.

is se t in mo t io n by t h e w aves o f sou n d which strike it the ,

vibr a tio n s t hus c aused are com mu n ic at ed to the i n t er n al ear ,

a n d thu s give ris e to th e se n s at io n of h e ari n g as w ill be ,

t re at ed m ore in d et a il in a l at er ch apter O n t h e i nn er sid e .

O f t h e tymp a n ic me mbr a n e lies a c avit y t h e E u s la e/

'

z z a n lu be , ,

w hich o pe n s i n ter n ally i n to the mou t h I f a bris t le be .

p assed t h rough this m embr an e i t w ill be see n t o emerge ,

t h rough a r a t her l arge rou n ded op e n i n g n e a r th e an gl e of

t h e j aw The exter nal fe at ures of t h e au di t ory org an o f t h e

.

fro gdi ffer m arkedly fro m t hose of m an in t h at all tr aces of '

a n exter n al e ar are abse n t a n d t h e t m an ic m e mbr an e lies

y p ,

exposed at the surface o f th e body i n s t e ad of lyi n g at t h e

i n n er en d o f a lo n g p ass age .

Ab o ve an d behi n d t h e blu n t tip o f the s n ou t li e t h e n os

z r z ls or ex ter n a l n a res These ope ni n gs are gu arded by
'

‘
.

va lues w hich ope n a n d close in co nn ec t io n w i th t h e m ove

m e n ts co ncern ed in r espi r atio n The ti p o f the u pp e r j aw .

64 TH E BIOLOGY OF TH E F ROG CH A P .

is slightly mov able an d i f i t be pressed u pw ard th e v alves

, ,

of t h e n o strils beco me closed an d preve n t the p ass age o f air ,

throu gh th e n ares P ass a bristle through th e n ostril an d

.

it w ill be fou n d to e merge i n to the mou t h t h rough o n e of a

p air o f rou nd ed ope n i ngs th e in fer n a l n a r es situ ated som e , ,

wh at beh i n d th e correspo n di n g exter n al op e n i ngs Th e .

a n u s or ope n i n g o f t h e clo a ca
,
lies so m ewh at do rs al in posi ,

t io n at th e posterior e n d o f th e body .

O n th e u pper sid e o f t h e h e ad in fro n t o f the eyes there , ,

u su ally occurs a sm all ligh t colored m ark th e l r o zo sp el ,

~
,
.

I n so m e speci m e n s t his spo t m ay be e n ti re ly co n ce al ed by

pigme n t ; b u t in most c as es i t m ay be d e tected al t hough i t ,

is oft e n quite i n co n spi cuous Th e bro w spot i s a fe at ure o f .

co n sider able i n t erest fro m th e fact th a t in the e mbryo n ic

,

d evelo p me n t o f th e frog i t is in co nn ectio n wi t h a peculi ar

ou t g rowth of the br ai n kn o w n as th e ep ip /iy sis or p in ea l
g l an a Whe
’
. n th e ou t er or dis t a l p ortio n o f t his structure
beco mes co n stric t ed o ff aft er th e bo n es of t h e skull h ave ,

d e veloped it is quite widely s ep ar a ted fro m th e b as al

,

por t io n w hich p ersis t s in c o n n ec t i o n with the br ai n eve n in

,

t h e ad ult frog The pi n e al gl an d i s fou n d in al most all ver

.

t eb ra t e s i n cludi n g m an in w h o m it w as give n by cert ai n

, ,

an cie n t philoso ph ers t h e i m por t an t fu n ctio n of bei n g th e

se a t of th e soul I t h as b ee n a scer t ai n ed t h at this structure
.

is a rudi m e n t o f a st al k w h ich formerly co n n ec ted wi t h a

m edi an eye ; in fact th ere are cert ai n reptiles ( H atteri a) in
,

w hic h t his eye is fai rly well developed co n t ai n i n g a cor n e a , ,

le n s reti n a co at s of pigm e n t an d o t he r structures ch arac

, , ,

t e rist ic o f a well develo ped visu al org an ; bu t in most v erte

-

br ates th e eye n o lo nger ap p e a rs or is repre se n ted by t h e ,

m erest rudi m e n t Th e co n n ecti o n o f t h e pi n e al gl an d wi th

.

t h e surface in th e d evel o pm e n t of th e fr o g an d t h e p ersis ,

t e n ce of th e brow spo t m arki n g th e p oi n t o f thi s forme r

66 TH E BIOLO G Y OF TH E FROG CH A P .

t o es i ncre ase successively in le n gth fro m the first 01 i n n er ,

o ne ,
to th e four t h t h e fift h to e bei ng co mm o nly a little
,

shor t er t h an t h e third Th e first t w o t oes co nt a i n three

.

j oi n ts e ach th e third an d fi fth four e a ch and t h e elo n g at ed

, , ,

fourth to e five j o i n t s O n th e u n der sid e of t h e articul atio n s

,
.

betwee n th e bo n es o f th e toes are s m all cushio ns t ermed

s u ba r lieu l ar p a rls The toes are co n n ected t o ge t h er by w eb
.
,

whi ch serves to m ake th e fo o t an efficie n t p addle as th e

an i m al swi ms through the w a t er The a mou n t o f web b e
.

twe e n the to es v ari es gre a tly in d iffere n t speci es of frogs ,

an d is a ch a ra cter which is th erefore m ad e u se of for pu r

poses o f cl a ssi fi c a tio n I n Ra n a p ip iens t h e web exte n ds

.

from the l ast j oi n t o f t h e first to e to th e n ext t o th e l ast

articul atio n of t h e seco n d to e th e n fro m th e l a st j oi n t o f th e
,

s eco n d to th e — n ext to the l as t ar t icul atio n o f the third ; fro m

th e l ast j oi n t of the third to a li t tl e beyo n d the seco n d j oi n t

o f the fourth an d fr o m abou t t h e s a me level o n the other
,

sid e of t his to e t o t h e l ast j oi n t of th e fift h D iffere n t i n di .

vidu als v a ry co n si der a bly a s reg a rds th e ex t e n sio n o f the

web e speci ally i f they com e fro m differe n t l oc ali t ies a n d
, , ,

t h e above st at em e n ts th erefore ,
mu s t n o t be expected to
,

a pply to all c a ses .

The ordi n ary resti n g posi tio n o f th e frog i s a squ atti ng

p os t ure wi
,
th th e an terior p a rt o f th e body elev a ted o n t h e

fo re li mbs which are b e n t at th e elbo w s a nd tur n ed i nw ard

,
.

Ne ar the middle of th e b ack is a sort o f hu m p du e to a

be n d at thi s pl ace in the vertebr al col u m n The h i n d limbs .

are folded toge t her t h e k n ees poi n ti n g outw ard a n d forw ard
, ,

an d th e e n ds of th e a n kl es lyi n g n e ar e a ch oth er a t th e hi n d

en d of body I n this positio n the frog is in re adi n ess t o

.

le ap whe n al arm ed by th e sudd e n exte n sio n of the hi n d

, ,

legs .

The skin of the frog is al most everywhere s moo th w i t h ,

III EX TE RNA L HA RACT E RS
C O F TH E F RO G 67

th e exc eptio n o f sm all sc a t t ered promi n e n ces occurri n g

m ai n ly o n t h e b ack an d o n t h e dors al side of th e hi n d legs .

No t hi ng correspo n di n g to h air or sc ales is to be fou n d in

the frog ; in fact W it h t h e r are exceptio n of rudi me n t ary
,

sc ales in so m e forms such struc t ures are e n tirely abse n t fro m

,

all of t h e rece n t Am phibi a The ge n e ral loose n ess of th e

.

a tt ach me n t o f the ski n is a fe ature which c ann ot fail to be

n oticed .

B ehi nd the eyes there exte n d t w o usu ally light colore d

,
-

ridges form ed by a t hicke n i n g of th e ski n an d k nown as ,

th e a orso la ler a l ner m a l p liefe or fola s Th ere are usu ally

’
-
’ ’
.

sever al sm aller an d somewh a t irregul ar lo n gi tudi n al folds of

ski n bet w ee n th ese Th e color of the ski n is much d arke r
.

o n th e upper or dors al sid e t h an below where i t i s e n t irely

,

whi t e
. The l a rge bl ack pigm e n t spo ts w hich occur o n th e
do r s al side of t h e b o dy an d legs ar e subj ect to much v ari a
tio n in size an d sh ape Th ere is Usu ally a p air of l arge
.

spots be twee n th e eyes an d a si ngle medi an sp o t in fro nt of

thes e Th e spots be t wee n t he dorso l ateral folds show a
.
-

te n de n cy to arrange t hemselves in two rows The spots o n .

th e h i n d legs are frequ e n tly elo n g at ed so as to form tr an s

verse b an ds I n additi o n to the bl ack pigme n t t here are
.

gree n an d golde n colors which are prese n t in v aryi n g

,

proportio n s .Th e ch an ge s in color w h ich th e ski n m ay

u n d ergo u n de r cert ai n co n ditio n s wil l be discussed in a l a t e r
ch apte r .
68 TH E BI O L O GY OF TH E F RO G CH A P
.

C H A PT E R I V

P R E LI M IN A RY A CCOUN T O F TH E IN T E RN A L ST RU CT UR E

M ou t h Cavit y the mouth o f th e frog is h eld widely

. If
ope n ,
th e followi n g p ar ts will app e ar I ii t h e r oof o f th e .

mou t h th ere i s a p ai r o f
rou n ded promi n e n c es c aused
by t he eyes a s h as alre ady ,

bee n m e n ti o n ed Arou n d .

the m argi n of the u pp er j aw

is a row of fi n e sh arp closely , ,

set leelb which are co n ic al in

sh ape an d curve d i nw ard
more or less a t th e tip E x .

ter nal to th e t eeth is a fl eshy

fold or upp er lip a nd o n th e , ,

i nn er sid e i s a groove t h e ,

s u lcus m a z in a lis hich re

g w ,

c eives the lo w er aw wh e n i t
j
is closed A n t eriorly this .

groove is c rossed o n e ac h
side by a lo w elev at io n t h e
F I G 8 —M u h f h f g id ly u lmn a
,

o t o t e ro w e
m
’

p os t a le im d i
d E E us chi t ub s ;
. ,

e r r r
op e ne ta an e

G gl is ; J l
.
,

a t ely behi n d t h e tip of t h e

o tt j ; L l t o w er aw a

l sub s l f ss ; M m di j aw t h e sulcu s is deepe n ed

, , ,

e ra ro t ra o a e an

sub s t l f ss ; N p s i
,

ro ra o a o ter o r
s ; O s p h gus ; P p ul
,

n are ag ai n
oe o formi ng the m edia n
a v1 ,

s t l ; S p i g f h u b os t a l fos a ; o n e ach
, ,

nar ro ra e o en n o t e
s r r s
c l ; T t gu ; lp t ub
,

vo a s ac on e er
side o f th e pulvi nar s are the
, ,

l m p li g
cu u l re n ua e.
iv TH E I NT E RN AL S T R UCT U RE 69

la ler a l f
s u b r os ir a l hichoss m, w
ar e mere de ep e n in g s of the
sulcus m argi n alis Th e lower j aw i s e n tirely devoid o f t e e t h
.

a n d is held tigh t ly pressed ag a i n st th e upper j aw t h e m an dib ,

ul ar muscles bei ng n orm ally in a st ate o f to n ic co n t r actio n ;

th e tip of t he lo w er j aw i s flexible an d is c ap able of bei ng ele
vat e d or depressed i n d epe n de n tly o f th e rest of th at struc ture ,

the j o i n ts of th e m ov able p art p a rs m en ia lis l yi ng u n der

, ,

t h e pulvi n ars o f t he u pper j aw Th e elev atio n a t t h e extrem e

.

ti p o f the lower j aw (t u ber cu lu m p r eling u a le) fits i n to th e

m edi an subros t r al foss a and there i s a sligh t d epressio n o n
,

either side of this t ubercle correspo n di ng to the pulvi n ars .

Th e two j aws t h erefore fi t toge t h er p art for p art w ith gre at

, , , ,

n ice t y. I n fact they form an air tight j oi n t w hich as w e

,
-

sh all se e l at er is a n ecess ary fe a ture in rel atio n to the p ec ul

,

iar mode of bre athi n g w hic h t h e frog i s fo rced t o employ .

Th e tip of th e upper j aw is like w ise mov able i ts fre e portio n ,

correspo n di ng in exte n t to th at of th e lower j aw so th at t h e ,

two p arts c an be r aised an d lowered togeth er I t m ay be .

n o ted th a t th e elev atio n o f t h e t ip of the j a w effects th e

closure of the n ar es a poi n t o f co nsid erable i mport an ce in
,

rel atio n to t h e process o f re spi r atio n .

I f a bristle i s p assed i n to o n e of t h e an terior n ares i t m ay ,

be see n to e merge i n to t h e an terior portio n of th e mou t h

c avi t y by o n e o f a p air o f rou n ded or oval ope n i ngs th e ,

eb oa n re or p os ter ior n a r es
,
B etwee n th e cho an ae is a p air
.

of pro mi n e n ces which be ar the o o mer ine feetb A t the .

posterior en d o f the bucc al c avi t y n e ar the angle o f the j aw

ar e th e l arge ope ni n gs E u s laebia n t u bes wh ich le a d outw ard
, ,

to t he ty mp a nie m embr a n e as m ay e asily be d emo n str at ed

.

by m e an s of a bris tle I n t h e m ale frog another an d a

.

sm aller p ai r of ope n i n gs m ay be see n o n t he lower sid e of

the bucc al c avity a li t tle in fro n t of th e E ust achi an tubes
,

th ese are the ope n i ngs of th e voca l sacs and th ei r co n ti ,

70 TH E B I O L OG Y OF TH E F RO G CH AP
.

un i ty wi th thes e org an s m ay b e s how n by p assi ng a bristle

i n t o th e m or by i n fl ati ng them by m e ans o f a blowpip e .

P osteriorly the bucc al c avity p r ese n ts two op e ni n gs in th e

middle li n e The ve n tral op e n i ng or g lof/is i s a n ar r ow
.
, ,

lo n gitudi nal slit in the middle o f a pro mi n e n ce c aus ed by

t h e c artil ages an d muscles of th e l ary n x ; it is ke pt closed

except du ri n g the p ass age o f air i n to o r ou t of th e lu ngs .

The dors al ope n i n g m arks th e begi nn i n g o f t h e esop /z ag u s ,

whic h le ads to t h e sto m ach Although c ap able o f gre a t

.

diste n tio n th e esoph agus is kept closed excep t whe n food

,

is bei n g sw allowed .

The flo or of t he mouth i s v ery diste n sibl e an d u n d ergoes

co n ti n uou s m ove me n t in respi ra tio n I n t h e middle p art .

m ay be see n th e by o ia ca r tilage which gives at t achm e n t for

’
,

th e t o n gu e and sever al muscl es th at move the floor of th e

mouth . The lo ng u e o f t h e frog is att ach ed in fro n t to
t h e lowe r j aw an d below to the hyoid c artil age I t s sh ape i s .

subj ect to gre a t v ari atio n accordi n g to the d egree o f c o n

trac t io n of it s v arious muscl es bu t in i ts n or mal rel axed
,

co n d itio n it is oblo ng fl atte n ed som ewh at n arr o w ed in

, ,

fro n t an d p r oduced at its posterior a n gles i n to t w o lobes

,

which exte n d b ackw ard o n eith er side of th e glo ttis th e

posterior m argi n is co n c ave an d the S ides which proj ect over
, ,

t h e at t ach m e n t at th e b ase l e ave a co n sider able sp ace o f th e

,

floor o f th e mouth u n covered I n th e m ucous me mbran e .

coveri ng t h e to n gue th ere are n u m erous gl an ds which secre t e

t h e mucus by w hich this org an is alw ays covered Th ere is .

also a l arge n umber of ill f which th er e e t w o ki n ds

p p
a ae o ,
_
a r

t h e fi lifo rm which are co n ic al o r t hre adlike in sh ape an d

, ,

the fu ngi fo rm which are l arge r a nd less n um e r ou s t h an t h e

,

former The l at t er are narro w at the b ase an d exp an d ed a t

.

the dis t al e n d I n a n aver age speci me n of R f usca F ixsen

. .

fou nd two hu n dred and thi r ty eight of th ese p apill ae I n a

-
.
 TH E IN T E RN A L S T RU CT U R E 71

speci m e n o f R p ipiens I h ave fou n d as m any as six h u n dred

.

an d forty .

Th e to n gu e of th e frog c an be re adily thro w n out o f the

mou th as i t is in c ap turi ng an i n sect an d wi t hdr aw n ag ai n
, ,

wi th gre at quick n ess Th e s ticky secre tio n with which i t

.

is covered c auses i t to adhere to i nsec t s or othe r prey wi t h

which i t comes in co n t act The victi ms are the n draw n
.

b ack i n to th e mouth wh e r e th e to n gu e m ay assi st in pushi n g

,

t he m b ack i n to th e thro at wh ere t h ey c an be sw allowed

,
.

The sticky subst anc e o n the frog s to ngu e is n o t produced

’

by the mucou s gl an ds but is derived in p art at le ast i f n o t

, , ,

e n ti rely fro m the in fer m a x illa fy gla n d w hich lies above th e

, ,

an terior p a rt of the roof of th e m o n t h Thi s gl an d is p artly.

i n closed by th e pre m axill ary bo n es j ust in fro n t of th e n as al

c aviti es I t re ally co n sis t s of an aggregate o f se ver al sm all
.

gl an ds ( twe nt y to twe n ty fi ve in R a n a escu len fa ) with

-

as m an y i n depe n d e n t d ucts le adi n g i n to th e c avity of th e

m outh W iedersheim has show n th at the secretio n of these

.

gl an ds is re m ark ably adh esive .

The mou t h c avity in ge n er al is li n ed by a mucous m e m

b r an e which v aries co n siderably in structu re in differe n t
-

regio n s P osteriorly i t is th row n i n to folds which co nve r ge

.

to w ard the esoph agus Th e epi t heliu m whi ch fo r ms the

.

su perfici al portio n o f this m embran e is cili ated over a l arge

p ar t o f the mou th an d t h ere are n um erou s goblet cells
,

sc attered abou t amo ng th e o thers Th e a c t io n o f th e cili a

.

m ay be d emo n s t r at ed in a live or r ece n tly killed frog by

sc atte r i ng powdered c armi n e over the roof o f the m outh .

Th e c ar m i n e grai n s will be see n to be c ar ried very slo w ly

b ackw ard and eve n tu ally they will be dr aw n i n to th e eso p h
,

a gus. Th e whole me mbr an e o f the m ou th t akes p art to a

gre a ter or less ex t e n t in th e produc t io n o f mucus .

The Teet h — The teeth of th e frog are ve ry n um erous

.
,
72 TH E BIOLOGY OF TH E FRO G CH A P
.

but of sm all size an d u n i form stru ctu re With th e exc ep .

t io n of the t w o p a t ches of vo m eri n e te e t h they a re co n fi n ed ,

t o t h e u pper j aw The j aw te eth r es t ag ai nst the d e n t al

.

proc e ses o f t h e m axill ar y an d prem axill ary bo n es to which

s
,

t h ey are att ached by ce me nt subst a n ce They are embedd ed .

in th e mu cou s m e mbr an e o f t h e mou t h ,

beyo n d w hi ch th ey proj ect o nly for a sh ort

dis t an c e E ac h t o o t h is a ppr o xi m a tely c y
.

lin dric al in form t aperi n g sligh tly t o w ard t he

,

u pper en d w hich is so m ewh a t i n curved

,
.

Th e b as al porti o n o f t h e tooth which i s fas

te n ed t o the j aw is c alled th e r oo f U po n .

t h is rests the cr ow n w h ic h is sep ar ated from ,

t h e roo t by a tr a n sverse furrow I n th e .

c e n ter i s a c avi t y filled with the p u lp w hich ,

i s a very v a scul ar tissu e in w hich the c ells

F I G 9 — T th f
- ee o ar e si tu ated th a t produce
fm
fizsvstf fi;
u
n ew m a teri a l for the growth of th e tooth .

f c ; B l t l Th e gre ater portio n o f t h e crow n is co m

a e ,
a e ra

i fé f j g osed o f subs an ce c lled which

n

i
ac cf o
p t a flen f in e

“ f t
o ’
ec

on forms a h ard c alc areous w all traverse d by

o o w er ,

i t)
l i
m P ul p
,
n u m erous fi n e br a n chi n g c a n als w hic h l e a d

fro m th e pulp c avi ty The u p pe r h alf o f th e .

cro w n is co ated w ith a very h ard resiste n t l ayer o f en a m el ,

w hich is co n sider ably thicke n e d over the ti p Th e e nam el .

sho w s a str atified s t ruc t u re b u t i t do es n o t co n t ai n th e ver t i

,

c al prisms fou n d in high er fo rms O u t side of t h e e n am el

.

t h ere is a thi n resiste n t m embr an e th e cu licu /a a en l is

,
Th e,
’
.

roo t of the too th is composed o f a subst an ce resembli ng

bo n e .

Th e teeth of the frog are n o t used for m as tic atio n bu t o nly ,

for h oldi n g prey w hich i s t h e pri mi t ive fu n c t io n o f tee t h

,

a mo n g vertebr a t e a n i m als Th ere is a co n t i n u al repl ace

.
74 TH E B IOLOGY OF TH E FROG CH A P I V .

i n t o t h e body c avi ty fro m in fro n t are t w o thi n very diste n ,

sible s acs w i t h a re t icul ated appe aran ce t h e lu ng s With a

, ,
.

blowpipe they m ay be i n fl ated fro m th e glot t is an d swell

e n ormously Whe n th e air is expelled from them t h ey c o n
.
,

tr act t o a very s m all size .

The di ffere n t p arts of the ali m e n t ary c anal v ary co n sider

a bly in size an d t ex t ure Above an d pr o j ecti n g behi n d t h e
.

liver i s the s to m a clz a thic k w alled muscul ar org an which

,
-

t ap ers tow ard th e posteri o r e nd wh ere the py lor ic co n s tr ic

tio n m arks its poi n t o f sep a r a tio n fro m th e sm all i n testi n e .

A n teriorly t h e sto m ach i s co nn ected with t h e short esop lz a

g u s le a di n g fro m t h e posterior e n d of th e bucc a l c avi t y .

Th e s m all intes tine which proc e e ds from th e pyloric e nd o f

,

th e sto m ach at first be n ds forw ard an d ru n s n e arly p arallel

,

w i t h t h e sto m ach thi s portio n is c alled th e du o den u m ; th e

portio n behi n d this or th e ileu m cu rves abru p t ly b ackw ard
, , ,

an d a ft e r formi n g sever al coils

,
sudd e n ly wid e n s out i n to
,

th e la rg e in testine The an terior portio n of the l arge i n te s

.

ti n e i s c alled th e r ectu m ; poste r iorly i t n arrows i n to th e

cloa ca ,
which p asses above th e ve n t r al p art of t h e p elvic
girdle an d termi n ates in th e a n u s o r ven t Alo n g i ts whole ,
.

exte n t the ali m e n t ary c an al is suspe n d ed fro m the mid dor -

sal portio n of the body c avity by a thi n tr a n sp are n t sheet ,

of me mbr an e t h e m esen tery

,
.

I n the U sh aped loop b et w e e n th e s t o m ach an d i n testi n e

-

li es an elo n g ated light colored org an o f irregul ar sh ape th e

,
-

p a n cr ea s. Th ere is a d ark red rou n ded body th e s leen

,p , ,

a t t a ched t o t h e m ese n tery n e ar th e an terior e n d o f the l arge

i n testi n e.

The r ep r odu ctive o rg a ns o r go nads lie o n eith er side of

, ,

t h e alime n t ary c an al an d ar e supported fro m t h e d ors al body

w all by s p eci al sheets o f m e mbr an e like the m ese n t ery The .

go n ads of t h e fem ale or ova r ies duri ng the breedi n g se aso n

, ,
FI G . 10 .
—O g s
f m l f g T h lim t y c l h b
r an of a t ff
e a e ro . e a en ar an a as een cu o

a t h gull t ( g l )
t de c tum e d m s t f h li
u . h b an re an o o t e ve r as e e n re

m d T h igh o ve y d f t b dy
. ls m d d th
e r t o va r t icl an a o a re a o re o ve an e ve n r e

o f th h t h b u df e e ar ds bd bd mi l i t d
as ee n t rn e o rw ar . a . v, a o na ve n c u an

tu d b ck ; l m
rn e li a m i y ; p d c p us di
coe . es , c oe a co -
e se n te i c ar t e r c . a , or a

p m f t b dy ; d
o su d s l or ; g t gull t ; 1
a o l ft u icl ; . ao, or a ao r t a u e . an , e a r e

1 l g l ft lu g ; l d l ft iduc ; l d i p i g i t th b dy
, ,

’
. u , e n . ou e ov t ov ts o en n n o e o

c i y ; l d i p s t i dil i u t us ; l y l ft y; l
.
, ,
”
av t . ov ts o er o r atat o n o r er . av e o va r r ,

f li p s c l i ; igh t u icl ; t ct um ;
, ,

p ti or ; pt on o ve r . cv , o t a va ve n r . on, r a r e rc re

b d igh t kid y ; l g igh lu g ; p t lp tl i ;

,

r .
,
r ne r d u r t n ru . re n a or a ve n r . ou ,

igh t i duc t ; i g i b dy c i
.
, ,

r d i ov
p t h r t y ;
. ov d i
’
ts o en n n o t e o av r . ov ts

p s i dil t i ; y t t sys t mic t u ks ; bl u i y bl dd r ;

,

o ter o r a at o n s s . r e r n u .
,
r n ar a e

u t ; t icl ( A ft P k
,

ur ,
re er d P k
v , ve n r ) e. er ar er a n ar e r.

75
 TH E B IOLO G Y OF TH E F RO G CH A P .

a re gre atly e nl arged an d m ay be recog nized by th e sm all , ,

globu l ar d ark colored eggs which for m t h e gre a ter p ar t o f

,
-

t heir m ass E x t er n al to the ov aries are th e l arge w hit e c o n

.
,

se m
CL

6 11 3

FI G I L —R iss c i f m h l f sid ; t h isc

an a t emp or a r i a . D e t on ro t e e t e e v e ra

s m h t dis p l c d us b d bil duc ; b by b dy f hy id ;

.

o ew a a e . a n , an ; .
,
e t . o o o

bl u i y bl dd ; b l i g i cl c ; t c us t i sus ;
,
'
r n ar p a er ,
it s o en n n to oa a c . ar , on ar e r o

blm c b llum ; l cl c ; 3 f hi d b ; p d
,

c e re e t c oa a cn . c e n ru in o t r v e rt e ra c . a ,

c p us di p sum ; b l c b l h mis p h ; d ly d s l lym p h

, , ,

or a o cr . z e re ra e e re . . s, or a

si us ; d du d um ; p p ic c id ; t E us t chi t ub ;
,

n v, o en e . cor e o ra o eu s . a an e

F R PA f l ; g l gl tt is ; g l gull ; il i um ; isch um ;
, ,

t p i ro n o ar e t a o u et IL IS, i
-

b d kid y; l l f u i cl ; l l g l ft lu g ; l li ; M M K m
.
, , ,

ne an, e t a r e u e n r ver C en to

b ; lf l lf c t y l b ;
. . .

m ck li ; u l ch f fi s
, , , ,

e e an n . a . 1 ne ra ar o r t ve r t e ra a .
,
o a or o e

p t l p ic l b ; d pi s um ; p d p ic di um ; P M X
,

o o m t o e 0 S T, o o an e -
t er n e er ar ,

p m xill ; p c s; p p st i is ; p p ubis ; t
. .
, ,

re p a a n, an re a . na , o e r o r n ar u, rc ,
re c

t m ;
u l g igh t lu g i t sm ll i s ti ; p
r u r pi l c d; n ; s . n a n te ne s . ca , S
’
na or S PH .

s p h t hm id ; p l s p l ; t s m ch ; si us s s; t g
. ,
,

ETH ene o s e en s to a s. v , n ve n o u u .

gu ; t st is ; u p tu i t th cl c ; U
, , ,

i
’
t te re t er ts a SI
’ ‘

on e ; s, ur ur er re n o e oa a ,

s yl ; icl ; ly l lym p h si us ; m i
, ,

u ro t e v , ve n t r t e v . . s, v e n t ra n vo . vo er n e

h sicul s mi lis
,

t ee t m vs . se ,
ve a e na .

voluted t u bes the ovidu cts Th ese are also su spe n ded to th e ,
.

dors al b o dy w all by thi n s h ee t s o f m embr an e ; th ey h ave n o

co n n ecti o n wi t h the ov ari es ; an teriorly t hey o p e n i nto the
1v TH E INT E RN A L S T RU CT U R E 77

body c avity n e ar the b ase o f the lu n g by a wid e fu nn el

sh aped mou th i n to w hich th e eggs fi n d their w ay after t hey
h ave be e n disch arged i n to the coelo m Th e w alls of t h e .

oviduc t s are t hick a n d gl an dul ar except tow ard th e posteri or

e n d w h ere th ey becom e exp an ded i n to th i n v ery diste n
, ,

sible s acs the u ter i in which t h e eggs coll ec t before bei n g

, ,

l aid The u teri op e n sep ar a tely i n t o t h e d o rs al side o f th e

.

clo ac a .

Th e go nads o f th e m ale or testes are very differe n t in

, ,

siz e an d appe aran ce fro m t h e ov ari es E ach tes t is i s an .

ovoid whi t i sh org an o ccupyi n g a si mil ar posi t io n to t h e

, ,

O v ary in th e body c avi t y an d suspe n ded in a si mil ar w ay by

a m e mbr an e the m es or clz iu m t o th e d o rs a l body w all

, ,
At .

th e an t erior en d of both the ov aries an d t es t es ar e att ached

th e f a t bodies corp o r a a dip osa w hic h are e asily recog n iz able
, ,

o n a ccou n t o f th eir yellow color a n d their divisi o n i n t o a

n u mber of fi ng er like lobes -

.

The kidn ey s are reddish fl at te n ed oblo ng org ans lyi n g

, ,

a g a i n st th e dors al body w all o n ei t h er sid e of t h e vertebr al

colum n Their posi tio n is so m e w h at be hi n d th e m iddle

.

o f th e body c avi ty an d they are brought i n to vi ew whe n

,

th e other abdo mi nal viscer a a re removed or pushed aside .

E ach kid n ey is co n n ected wi t h a tube th e u r eter which , ,

ru n s alo n g i ts outer ed ge an d e mpties n e ar th e ope n i n g of,

it s fellow i n to th e dors al sid e o f th e clo ac a

,
I n t h e m ale .

th e u re t ers are exp an ded dis t ally to form sem in a l vesicles ,

which in some species of frogs re ach a co n si der able size .

Th e u r in a ry bla dder i s a l arge bilobed sac lyi n g in th e pos

,

t erio r en d o f th e body c av i ty an d ope n i n g i n t o th e v e n tr al

,

sid e of t h e clo ac a ju st below th e ope n i n gs o f th e ureters .

I f t h e bl a dder i s i n fl ated by me an s o f a blo w pip e i n s erted

i n t o th e clo a c a it will swell t o a large size an d an accu rat e
, ,

i de a m ay be g ai n ed o f it s for m an d co nn ectio n s .
78 TH E BIOLO G Y OF TH E FRO G CH A P .

Th e w alls of the body c avi ty an d the v ariou s org an s

co n t ai n ed in it ar e cover ed by a t hi n m oist glis t e n in g , ,

membr an e th e p er ito n eu m This m e mbr an e is perfec tly

,
.

co n ti n uous throughou t and is simply refl ected ove r th e ,

v arious org an s I f we i m agi n e th at th e body c avi ty were

.

origi n ally empty an d th at th e org an s i t co n t ai n s were

,

pushed i nto it fro m the outside c arryi ng th e p erito n eu m in ,

FI G . 12 .
—D i g m
c ss s c t i a ra f th b dy f of a ro e on o e o o a f g sh i g th
ro ow n e

c u s f t h p i t um by d t d li
o r e o e bd
er o ne a ot e ne, a . v, a bd mi l i ;o na ve n

d d s l
. a o, t ; l il i um ; m t i t s t i
or a ao r a ; b d kid z n e ne ne y ; m m muscl s ’
e

f b ck d bd m s p c t i ly ; m m s y; p p p p
, , , , ,
'
o a an a o en re e ve es , e e n te r . er , . er

p i l l y f p i t um ; p t p s c l i ki ; py
,

ar e t a a er o er o ne cv , o t ava ve n ; é s S n s

sp m y s tis ; subcu us l y m p h t u s tyl ;

.
, ,

er ar ly or te s . cu . . s, ta neo sa c ; u . s ro e

isc l p i t um ( A f P k d P k
,
'

v p .
p er , v . er ,
v e ra er on e . te r ar er an )
ar er .

fro nt o f t he m it will hel p us to u n d erst and th e rel atio n o f

,

thi s membran e to the structures i t su rrou n ds I t is no t to .

be i n fe r red ho w ever th a t these rel atio n s were brough t

, ,

about in j ust this w a bu t th e d evic e will b e useful in e na

y ,

bli n g u s to ge t t h e co ndi t io n s cl e arly in mi n d The alim en .

t ary c an al the liver t h e lu n gs the go n ads ovid u cts bl adde r

, , , , , ,

fat bodies an d oth er org an s are c o vered w i t h perito n eu m

, ,

w hich u su ally adh eres closely t o th e surfa ce Th e m ese n .

Iv TH E IN T E RN A L S T RU CT UR E 79

t eries an dth e si mil ar shee t s of m embr an e s u ppo r ti n g th e

ov aries o viduc t s a n d tes t es are d o u ble as w o uld n a tur ally
, , ,

be the c ase if th es e organ s were pu sh ed in in th e w ay m en

t io n ed Th e peri t o n eu m p asses dow n from t h e bod y w all
.
,

covers thes e org an s th e n p asses b ack to t h e body w all agai n

, ,

th e t w o shee t s of me mbr an e comi n g close together except

where they are sep ar ated by th e org an they surrou n d Th e .

arteri es an d v ei n s supplyi n g th e org an s ge n er ally ru n b e

t w e e n th e two l ayers o f the su ppor t i n g membr an es Th e .

por t io n of th e p erito n eu m surrou n di n g t h e alim e n tary c an al

a n d it s appe n d age s is c alled th e viscer a l t h e p art

appli ed to the body w all t h e p a r ieta l lay er t he m ost

-

.
,

p art th e p ari et al l ayer is grow n fa st to the dy muscles ,

but o n th e dors al side of the body it i s sep ar ated from the

w all fo rmi n g a l arge ly mp lz sp a ce t h e cister n a m ag n a or
, , ,

s u bver teb r a l l m lz s in uTfi Th e kid n eys li e in this sp ace ;

y p
he n ce t h ey are covered w ith p eri to n eu m o nly o n the ve n tr al
sid e an d n o t co mpl etely i nvested by i t like t h e other
,

viscer a Th e m embr an e prev iously m e n t io n ed which ex t e nd s

.
,

be t we e n th e ve n tral bod y w all an d th e peri c ardiu m an d

liver is a portio n o f th e peri t o n eu m formi n g a sor t of ve n tr al
, ,

m ese n tery ; it is r efl ected upo n the ve n t ral body w all o n

th e o n e h an d and spre ad ou t over th e peric ardiu m an d liver
o n t h e oth er .

Th e c oelom i s filled with a tr an sp a re n t fluid th e coelom ic ,

or p er itonea lflu id w hich i s esse n t i ally t h e s am e as t h e lymph

,

fou n d in oth er portio n s of t h e body O wi ng to th e fluid in

.

which th ey li e an d th e sm ooth n ess o f their peri t o n e al co a t

in g th e o r g an s in th e body c avity ar e e n abled to glid e over
,

e ach o t her wi t h little fric t io n

O r gans out side of t h e B ody Cavit y —
.

Ab o ve th e c oelo m
.

th ere i s a seco n d c avi t y surrou n ded by th e bo n es o f t h e v er

t eb ral colu m n an d skull and co n t ai n i ng the cen tr a l n er vo u s
80 TH E BIOLOGY O F TH E FROG CH A P.

sy s tem This n eu r a l ca vity as we m ay well c all it exte nds

.
, ,

far th er forw ard t h an th e coel o m a n d is sep ar at ed fro m th e ,

l atter by the b ases o r ce n tr a o f the v e r t ebrae Th e an teri or .

portio n o f th e ce n tr al n erv
ous sys t e m o r br a in li es , ,

in th e skull an d is co n t i n ,

u ed pos t eriorly as th e sp i
na l co r d which i s i n closed ,

wi t hi n t he vertebral col
u mn I f w e m ake a cross .

sec t io n through th e frog

some w here n e ar the m id dle ,

we sh all fi n d th at the body

co n t ai n s two lo n gitudi nal
c avities sep ar ated by the
ce n tr a o f th e v ertebr ae ,

LL S C
th e c oel o m b elow a n d t h e . ,

FI G 3 O g s i th
. 1 u l c i t y n eu r al t u be above Arou nd
r an n e ne ra av .

i p t f th i s c i t y
.

Th t
e an e r o r
bo th of th ese is a l ayer
ar o av co n

t i s th
a n b i hich is c m p s d f
e ra n , w o o e o

th elf c y l b s f l ; th c
t o l
a or
o fo
mu sclese
which io
s mu ch e e re

b l h mis p h s b l ; t h di thicker dors ally al t hough i t

.
,

ra e e re cr i e

c p h l d ; t h O p t ic l b s
.
,

en e a on, z en e o e

o pt l ; c
. b llum blm ; d m
‘

e re
co
e
mpletely surrou n
c
ds the an e
,

dull bl g t m d bl
,

a o on c oelo m below A n d ou t sid e

a a, e o n c, n eu

l c l ; p d s p i l c d di g of the muscles fro m which

. . .
.

ra an a s c na or en n

fi l m t mi l f t;
.
,

i th
n e u y er na e , e, e e ,

( A ft P k d P k
. .

er )
ar i t i s sep ar at ed by l arge
er a n ar er .

lymph sp aces crossed by a few b ands of co n n ective tissu e ,

is t h e ski n .

Wi th the exceptio n of th e loose at t at ih m en t of the ski n ,

all th e fe a tures o f s t ru c ture m e n tio n ed in th e l ast p ar agr a ph

belo ng t o t h e ver t ebr at e a ni m als in ge n e ral We h ave n ex t .

t o se e h o w t h ese fu n d a me n t a l fe a tures o f structure c a me to

be es t ablished .
82 TH E BIOLOGY OF TH E FROG CH A P .

M an y of th e old er wri t ers w ho specul ated o n th e probl em

o f de ve lo p m en t were o f th e opi n io n th a t the egg co n t ai n s in

mi n i at ure all o f th e org an s o f th e body in th e s a m e form and

rel at io n in which t h ey occur in the adul t an d th at the p arts ,

simply exp an d as n u trim e n t i s absorbed u n t il th e emb ryo

a t t ai n s it s m aximu m siz e .The process of d evelopm e n t w as
comp ared t o t he u n foldi n g o f a flo w er fro m a bud in w hi ch the
’

p ar t s occur in th e s ame rel ative posi t io n a s in the exp an ded

blosso m a nd si mply u n fold through the absorp t io n o f sap u n til
th e flo w er re ach es it s fi nal form This vi ew which is k n o w n
.
,

as evolu tio n or p r e fo r m a tio n w as ch am pio n e d by such m e n

,

as H a ller ,
L eib nitz B o n n et an d S pall an z an i
, ,
i t re ally
a mou n te d t o a de n i al o f develop me n t Wh at appe ars to be
.

such accor di n g to t hi s i n terpre t atio n is si mply grow t h the

, , ,

exp an sio n of some thi n g preform ed The Abb é S pall an z an i

.
,

w h o m a de exte n sive a n d most excelle n t studi es o n th e b r ee d

ing h abits of Am ph i b i an s co n sidered the eggs o f t h ese ani

,

m als t o be s m all embryos of t adpoles whose p arts through

the i n flu e nce of t h e sperm atic flui d were sti mul ated to gro w t h
a n d a c t ivi t y
. O the r wri t ers am o n g who m C F Wolff o c c u
,
. .

pi es t h e most p r o mi n e n t pl ac e reg arded th e egg as a m ass

,

of si mple u n org an ized m at eri al which beco mes more an d

more co mplex as d evelo p me n t proc eeds This doctri n e .
,

which is ge n e r ally k n ow n as th e th eory of ep igenes is h as

o
,

becom e m ore widely accep te d in rece n t times although n o ,

o n e at prese n t espouses ei the r e p ige n sis or evolutio n in the

form s advoc ate d in th e eightee n t h ce n tury .

The c ell t h eory or th e doctri n e th at ani m als an d pl an ts

,

are com posed of livi n g u n its or cells w h ich w as pro mulg ated
, ,

by S chleid e n an d S chw an n in 1 8 3 9 did much to correct t h e

,

extrav agan t forms o f t h e old er t heori es o f evolu tio n an d ,

st ar t ed i nvestig ators o n the ro ad to a tru er co n ceptio n of

developme n t A farther step in adv an ce w as m ade wh e n
.
 T H E DE V E L O PM E NT OF TH E FROG 83

, it w as a scert ai n ed th at th e egg is a si n gle c ell No t m an y.

ye ars aft erw ard it w as discovered th at the sp erm at o z o On is

likewise a si ngle cell an d th at in fer t iliz at io n there is a u n io n
,

o f the n uclei o f the ovu m an d sp erm at o z o On e a ch c o n t rib u t

,

ing an equ al sh are of chrom ati n to t h e n ucleus of t h e fer

t iliz e d egg an d th e n ce t o all th e c ells o f th e body o f the
,

embryo The whole aspec t of t h e p ro b le in o f d evelopm e n t

.
‘

is ve r y di ffere n t fro m wh at i t appe ared to the olde r n at u r al

ist s . H owever w e m ay reg ard th e mod er n forms of th e
doctri n es o f evol utio n an d epige n esis — fo r both poi n ts of
,

view are still h eld ,

it is c ert ai n th at dev elop m e n t ac tu ally
p r oc eeds from a si n gle cell to a body co n sisti ng of a multi
tu d e o f cells o f gre at v ariety of form an d fu nc t io n Thi s .

c ell m ay be e n ormou sly complex co n t ai n i n g so m eh ow fea

,

tu r es whi ch represe n t all th e di ffere n t org an s of t h e body or ,

i t m ay be comp ar a tive ly sim pl e in structure an d th e differ ,

e n t iat io n s a ppe a ri n g in t h e e mbryo m ay be the results o f the

in ter ac t io n s of its p arts an d t h e i n flu e n ce o f exter n al co n di

tio n s E volu t io n an d epige n esis both h ave t h ei r advoc at es
.
,

but th ei r differe n ces h ave beco m e l ess wide as k n owl edge o f

e mbr yology h as adv an ced .

The J elly an d it s Uses — Th e egg of the frog wh e n it

i s l aid in th e w ater i s surrou n ded by a sph eric al m ass o f
tr an sp are n t j elly At first th e co at of j elly i s less t h an the
.

di ame t er o f th e egg in th ick n ess but through th e absorp t io n

,

o f w ater it gr adu ally swells u n til it be co mes two or thre e

ti mes this di ame ter The j elly co n sists o f three l ayers a
.
,

thi n in n er lay er clos ely a ppli ed t o t h e egg a thi ck middle

,

lay er o f more fluid co n sis t e n cy an d a t hick o u ter lay er

,
Th e .

co ats u n d er th e microscop e sh ow a co n ce n tric se r ies of fi n e

/
li n es i n dic ati n g t h e strat ific atio n o f th e m at erial T h e fu n c
,
.

tio n o f this j elly is prim arily th e protectio n of th e eggs I t ,

k e eps the m fr ee fr om di r t b acte r i a an d th e spo r es of fu ngi

, , ,
84 TH E B I O L O GY O F TH E F RO G CH A P .

an d a lso fr om the att acks o f aqu atic i n sects an d s n ails and ,

v ariou s m ech an i c al i nj u r ies wh ich would other w ise affect

the m B er n ard an d B rat u sch ek co n sid er th at the j elly also
.
1

serves to ke ep th e eggs w ar m er th an the su rr ou n di n g w at er ,

which in the sp r i ng wh e n t h e eggs are laid is o fte n very col d

, , ,

a n d frequ e n tly covered with i ce Th e j elly is supposed to .

ac t upo n th e pri n c i ple o f a hotbed allowi n g free e n tr an ce ,

to th e su n s r ays but ch ecki n g r adi atio n from th e egg The

’
,
.

FI G . I 4
—E gg i j lly
.
'

n e .
( A ft er S chul z
t e .
)

h e at w aves th at r ad i a te fro m th e eggs are lo n ger less re fran ,

g ib le an
,
d p ass through t h e j elly less re a dily th an t h e shorter
w aves whi ch are abu n d an t in t h e di re ct h e a t o f the su n
,
.

B er n ar d an d B rat u sc h ek fo u n d in experi m e n t i ng u po n th e
2

j elly th at the gre at er t h e w ave le ng t h t h e less h e at p assed ,

t hrough in comp ariso n wi th an equ al a mou n t of w a ter u n d er

th e s a me co n di t io n s Th e j elly w as thus sh ow n to h ave t h e

.

1 B er n a r d un d B rat u sc h e k , Der Nu t z en de r S ch le im h u lle n fu r die

F rosch eier , B i o l. Cen tr lb .
, Bd . 1 1 ,
1 89 1 .
2 I bid .
 T H E DE V E LO P M E N T OF TH E F RO G 85

p r op e r ty r equ i r e d to keep the eggs w arm er th an th e su r

r ou ndi ng m ediu m Th e bl ack pigm e n t o f the eggs which
.

r e adily absorbs the he at r ays also fu n ctio n s in th e s am e

m anner
Resist ance of t he S exu al P r odu ct s t o Cold —The egg
.

m asse s o f t h e frog are l aid so e arly in th e spri ng th at the

w ate r co n t ai n i n g the m is fr eque n tly fr oze n F isc h er S i gw art
.
-

reco r ds fi n di ng egg m asses which were froze n solid for two

d ays duri n g which t h e te m per atu re s an k to 8 C Wh e n
,
°
.

th e eggs we r e gr adu ally th awed out they u n d erwe n t a n orm al

,

process o f d evelop me n t al t hough som e w h at slower th an

,

u s u al an d g ave ris e to l arvae wh ich left t h e j elly two d ays

,

afterw ard H o w lo n g eggs m ay be froze n an d how low a

.

temp e rature t hey can e n dure and still r et ai n their powe r of

develop me n t is no t determi ned .

The sperm atozo a of th e frog m ay al so be fro z e n without

fat al results Th e Abb é S p all anz an i showed th a t sperm ato
.

z o a fr oze n in ice for h alf an h our are able to c au se eggs to

develop but i f kep t in ice for sever al hours this power is lost
, , .

Th e sperm atozo a app are n tly h ave less power of resist an c e to

cold th an th e eggs c areful co m p ariso n s howeve r h ave n o t
, ,

yet be e n m ade
—
.

S t ructure of t he U n di id
v e d E gg The e ggs o f th e
.

frog as th ey occur in th e body of the fe m al e a fter th ei r

disch ar ge fr om the ov ary are su rrou n ded by a very thi n vitel
lin e membr a ne whi ch represe n ts its cell w all
,
O n o n e side.
,

represe n t i n g th e a n im a l p ole the egg is colored by bl ack

,

pgi m en t which
,
as is show n in sectio n s is m ai n ly co n fi n ed to
, ,

th e pe r iphery i m medi ately u n der the vitelli n e me mbran e .

This pigme n t is in th e form of mi nute gr an ules embedded in

the p r otopl asm The n u cleu s o f the egg lies excent rically
.

ne ar th e mi ddle o f t h e d ark pole Whe n th e egg is in the

.

o v ar y t h e n u cleus is l arge an d co n t ai n s a l arge a mou n t of

,
86 TH E B I OLO G Y OF TH E FRO G CH A P .

fluid k n ow n as n u clea r sap bu t j ust precedi n g t h e esc ape of

,

t h e egg i n to t h e body c avity an d duri n g it s p ass age dow n

t h e ovid u ct th e n ucleus beco m es s hru n ke n through t h e exu

d atio n o f i ts fluid an d u n d ergo es a process o f divisio n ih

,

volved in the form atio n o f th e fir s t p ola r body As S chul tze .

h as show n alo n g wi t h th e shri n k age o f the n u cleus th ere

,

a ppe ars a m a ss o f fluid be n e a th t h e an i m al pole which h e

co nsiders to be th e n ucle ar sap The pl a ce i s m arked by a .

light colored spot n e ar t h e ce n ter o f th e d ark c a p The

-
.

gre at m ass o f th e frog s egg is m ade up o f y olk wh ic h

’

occurs in th e form o f gr an ules e mbedded in th e cytopl asm .

Th e yolk is a s emi flu id albu mi n ou s subst an c e which i s e m

-

ployed fo r th e n u t ri t io n o f th e dev elopi n g embryo I t is .

m ore abu n d an t tow ard the light — colored or veget al pole of

t h e egg th e regio n arou n d th e an i m al pole co n t ai n i n g rel a
,

t ively more cy t opl asm The yolk gr a nules are of v ariou s

.

sizes an d are u su ally spheric al or ov al in form B y t h e

,
.

a ctio n of cert a i n re age n ts th ey m a be broke n up i n to flat

y
t e n ed pl a tes wh ich accordi n g to S chultze do n o t exis t
, ,

as such in t h e livi n g egg .

M at u r at ion — Th e proc ess of m atur atio n co n sists in t w o

.

s u c c essw e d ivisio n s o f th e ovu m resul t i n g in th e form atio n o f

,

t h e t w o pol ar bodi es These bodies are m i n ut e globules

.

extruded at the an i m al pole o f t h e egg M orphologic ally th ey .

a re cells produced by very u n equ al divisio n s of t h e egg

, ,

w hich i s o n ly a cell o f very l arge size The first pol ar .

globule is give n off whil e t h e egg i s wi t hi n th e body th e pre ,

limi n ary steps of th e process o ccurri ng j ust before th e egg

le aves th e ov ary The l arge w a t ery n ucleus shri nks th e
.
,

chro m at i n becomes aggr egated i n t o de fin i t e bodies o r c h ro

m o so m e s a spi n dle forms a t right an gl es t o th e surface of t h e
,

egg an d h al f th e chrom ati n wi t h a sm all a mou nt of cyto

, ,

is extruded as the fi r s t pol ar body .

 TH E D E V E LO PM E N T OF TH E FROG 87

I n Ra n a f usca seco n d pol ar body is give n off after

t he

the egg h as bee n l aid an d wi t hi n h alf an h our aft er fertiliz a

,

tio n n o res t i n g st age o f th e n ucleu s i n terve n es betwee n t h e

two m at ur a tio n divisio n s an d in th e seco n d divi sio n as in
, ,

th e first h alf th e chrom at i n m ateri al o f th e egg is extrud ed

,
.

The two pol ar bodies in Ra n a fus ca are o f about equ al size ,

a n d are ei the r a tt ached to t h e an i m al pole o r flo a t in t h e

fluid w hich accu mul ates betwee n the egg and t he vi t elli n e
m embr an e I f t h e j elly is removed fro m a freshly l aid egg
.
,

a n d the light spo t or fove a at t h e an i m al pole observed

, ,

wi th a h an d le n s th e ex trusio n o f th e seco n d pol ar body

,

m ay be wit n essed .

Whe n th e m at ur atio n d ivisi o n s ar e completed the a mou n t ,

o f chro m at i n in th e egg h as become mu ch reduced an d th e

n umber o f chr o m oso m es di mi n i sh ed by o n e h al f Whe t her .

t h e c hro moso mes in th e m atured egg are double so th at th e ,

reduc t io n in n umber is o nly app are n t o r wh eth er there is an ,

a ctu al loss of h alf o f the chro moso mes i s a ques t io n s till

u n der discu ssio n .

F er t iliz at ion —Th e ac t of fer t iliz atio n co n sists in the u n io n

.

o f two cells th e ovu m fro m t h e fem ale an d the sp er m a ta z oo n

’ ’

fr o m the m al e As be arers of heredit ary qu alities th ese

.

two cells are equ al but in size an d form t hey ar e as dissimil ar

,

as they c an well be The sperm cell i s a mi nute elo n

.
,

ga ted body co n sisti n g o f a narrow poi n ted lz ea d form e d ,

m ai n ly of the n ucleus a sh o rt m iddle p iece j u st behi n d t h e

,

he ad an d a lo n g very sle n de r ta il As before described

, ,
.
,

t h e semi n al fluid of the frog is sh ed over th e egg m asses as

t h ey ar e extruded i n to the w ater from th e body o f th e fem ale .

Th e sp erm atozo a which sw arm in th e se mi n al fluid are active ,

a n d s w i m abou t by th e movem e n ts of the t ail worki ng th eir ,

w ay through the j elly o f th e egg m ass u nt il o n e co m es in

c o n t act wi t h an egg Th e n t h e sp erm at o z o On slowly pe n e
.
88 TH E BIOLO G Y OF TH E FRO G CH A P .

tr ates the egg subst anc e Th e e n tr an ce o f o ne sperm ato

.

z o On se ems t o c ause so me ch an ge in th e subst an c e o f th e

egg wh ereby o t her sper ma tozo a are preve n ted from e n teri ng
it as n orm ally a n egg is fe r t iliz ed by o n ly o n e
,

sp erm cell although ther e m ay be thous and s

,

of o th ers in th e i m me di at e V ici n ity Whe n .

th e h e ad o f th e sp erm at o z o On h as e n tered
th e egg i t begi n s to e nl arge an d it s n ucleus
, , ,

which is n o w k n ow n as t h e m a le p r on ucleu s ,

assum es a sp heric al form I t migr a tes slowly

.

to w ard the ce ntr al p ar t o f th e egg d r aggi n g ,

in beh i n d i t a m ass of pigme n t gr an ules fro m

th e periphe r y so t h at it s course co mes to be
,

m arked by a d ark stre ak B e fore th e m ale .

pro n u cleus h as pe n etr ated very far th e pro

c ess of m a tur atio n i s brought to co mpletio n
by th e form atio n of th e seco n d pol ar body .

Th e n ucle ar m ateri al re m ai n i ng in the egg

aft e r this seco n d m at u r at io n d ivisio n go es
i n to a r esti ng st ag e formi n g th e fem a le p r o
,

n u cleu s o r t h e n ucleus of th e m atured ovu m

,
.

FI G 5 —
. 1 Sp
. The m ale an d fem ale pro n uclei a ppro ach
er

252 2532 ,e
2
an d fi n ally fuse i n to o n e which i s c all ed the ,

t o (
. A f t L c o
p u la t io n n u cleu s
er a Th e n u mbe .r of chromo
V l t t 5 1 so m es co n tributed by bo t h p are n ts to t he
a e e °

G g )
eo r e .

n u cleus of the fert il ized egg is t h e s a m e an d ,

this h as doubtless a fu n d a m e n t al rel at io n t o th e fact th at ,

o n the aver age offspri ng i n herit qu ali t i es fro m bo th th ei r

,

p are n t fo r ms in an equ al d egree Thi s correl atio n of

.

e qu ality of i n herit anc e fro m t h e two p are n ts with the

equ ali t y of t heir co n tributio n s o f chrom ati n m at eri al to th e
fer t ilized eggs l e n ds stro n g suppor t to the vie w th at i t i s to
th e chrom at i n o f the n ucleu s th at we m us t look for the
 TH E BIOLOGY OF TH E F ROG CH A P .

H
FIG 6 —S gm i f chul tz ) c ll s g A
A S to o ta e ,
e n tat o n f gg (
e t er e e
-

1 e o . ,

i h h b gi i g f th s c d fu igh t c ll s g sh i g
.
. .

; B ta e ,
e on rrow e e ow n
t t e
-

w e e nn n o ,

t h c ss fu
e ro rro w a t t h im
e lan p l ;aE igh
o et c ll,
set g ; C
-
D eF G a e , , , ,

six t c ll see n -
e ta e g s sh i g i t i s i t h p l f cl g ; H "
ow n var a on n e an o hi y ea va e ,
t rt

c ll s t g m g s l m g s
,

F “D f h F E )
’

M t t
’

t w o -
e a e .
( ro or an p e ve o en g
o g e ro
 TH E D E V E LO PM E N T OF TH E FROG 91

r arely itli es at righ t an gles to i t although it m ay occur in

,

al most an y i n term edi a te posi tio n .

The Seco n d cle av age appe ars abou t t hree qu arte r s o f an

hour a ft er th e first ; th e furrow ex t e n ds gradu ally fro m the
a n im al to t h e veget al pole at right a ngles t o the first fur r ow
, ,

an d divid es th e e gg i n to four cells The first an d seco n d

.

cle av age pl an es st an d in a toler ab ly co n st an t rel atio n to t h e

axes o f the body o f the e mbryo the first cl e av age pl an e
,

m arki ng t h e m edi an or s agitt al pl an e of th e future ani m al

b u t th is is a rule n o t without excep tio n s .

Th e third cle av age furrow com es in a littl e above th e

equ ator o f th e egg an d a t righ t angl es to the o t h er t w o the
fou r upper cells cut off by t his divisio n are a li t tle sm alle r
th an th e lowe r fou r At th e n ext cle av age the furrows ru n
.

n e arly vertic ally an d he n c e at right a ngles to the t hi rd c leav

age pl an e S om eti m es t h e furro w s meet at th e an i m al pole

.
,

but more freque n t ly they cut through th e firs t or seco n d

cle av age furrows prod uci n g thus a bil ater al arr an geme n t o f
,

th e cells The fourth cle av age furrows are subj ect to more
.

v ari a tio n at th e veget al pole o f t h e egg an d th e subseque n t

,

divisio n s soo n become so irregul ar t h at i t is i mpossible t o

tr ac e ou t an y pl an of procedure A fi ft h cl e av age occur s
.

typic ally p ar allel to the third appe ari ng first in the u pp e r

,

h emisphere an d the n in th e lower The cle av ages thus far

.

the rul e th at e ac h cle av age pl an e com es in at

to the previous cle av age pl an e D evi atio n s .

fr o m t h e t ypic al m ethod of cle av age are app are n tly o f li t tle

mome n t eve n in th e first few divisio n s as su ch ab n orm ally
, ,

dividi n g eggs m ay n everth eless produce pe rfec t embryos .

C le av age t akes pl ac e more r apidly in the d ark o r an i m al

pole si n ce at th at pl ace t h e p r o t opl as m is most d e n se Y olk
, .
,

which is most abu n d an t in t h e w hi t e o r veget ative side of th e

egg del ays cell divisio n an d w e fi n d in the l ater st ages o f
, ,
 TH E BIOLO G Y OF TH E F RO G CH A P .

G I
FI G . 17 .
—S gm e f m ti i
f t h bl s t p
en t a t o n o f A th e egg a n d or a on o e a o o re .
,

eigh t c ll s g s f m
-
e sid ; 8 b gi i g f six t c ll s t g ;
ta e ee n ro o ne e e nn n o ee n -
e a e

C t hi t t c ll s g ; D f y igh t c ll s g ; E F d G succ s
,

r y -
w o -
e ta e o rt -
e -
e ta e ,
an e

si l t s t g s f cl g ; H b g i i g f h bl s t p i h f m
, , , ,

ve a er a e o e a va e e nn n o t e a o o re n t e or

sm ll c sc i cul bl s p i id
,

o f a I
a re thent
g f h
C r ar a to o re o n e ve e t a t ve s e o t e
f
,

e
g g ( A . M g ) ter or an .
94 TH E B IOLOGY OF TH E FROG CH A P .

s m aller as the proc ess o f in pushi ng is co mpleted Th e pro -

cess of g astrul atio n which is exe mplifi ed in i ts typi c al fo rm in

,

the d evelopm e n t o f a st arfish or sea u rchi n bec o m es very -

much modifi ed in differe n t an im als S uch i s the c ase in t h e .

d evelopme n t o f t h e frog The l arge accu mul atio n o f yolk at

.

th e vege t al side of t h e bl astul a preve n t s the i n v agi n at io n o f

this regio n fro m t aki n g pl ace in th e typic al w ay Th e s am e .

FI G 19 .
—S gi tt l s c t i
a a e on t h ugh f g mb y B bl s t c l
ro a ro e
gm r o . , a o oe o r se en

c i y; BP f bl s p ut p id mic l y
.

t a t io n av t ,
lip o a ; E E
to o re f,
o er or e er a er o

e ct d m ; E N i
o er , n n er l y f c t d m ; Y y lk c lls
o r n e i vo u s a er o e o er o e

( A ft M sh ll )
.
,

er ar a .

e nd is re ached p artly by a proce ss o f in pushi ng and p ar t ly -

by the overgrow t h o f the whi t e pole by t he d ark Th e in .

pushi n g an d overgrowth t ake pl ace m ore o n o n e S id e o f the

egg t h an the other an d thes e pro cesses are first i n dic ated
,

by the a ppe ar an ce of a cresce n t ic groove a lit t le below t h e

equ ator of th e egg The cresce n t repres e n t s t h e begi n n i ng
.

of the bl astopore The groove is d eepest at the ce n t er and

.
 TH E D E V E LO PM E N T OF TH E FRO G 95

thi n s ou t tow ard the e dges w hich gr adu ally exte n d arou n d ,

th e lower pol e of th e egg I n this w ay the cresce n t becomes .

co n verted i n t o a circle an d the circl e gr adu ally beco mes ,

sm aller a nd sm aller u n til o n ly a s m all p art of t h e light colore d

'

yolk k n ow n as the yolk plug appe ars in th e midst of th e d ark

, ,

are a. Th e white pol e is thus overg r ow n by the d ark but ,

n o t wi th equ al r apidity from all sides the closi n g in t aki n g ,

-

pl ace much more r apidly o n th e side wh ere th e cresce n tic

fold origi n ally appe ared an d w hich subseque n t eve n ts prove
,

to be the an terior en d o f the e mbryo .

I f we m ake a ver t ic al sectio n t hrough th e embry o at right

an gles to th e cresce n tic bl astopore w e sh all fi n d the l at ter ,

is t h e m ou t h o f a c avity whi ch exte n d s so me dist an ce i n to

th e egg Above this c avity w hich is c alled th e a r chen ter o n
.
, ,

is a comp ar at ively thi n r oof closely ap p lied t o th e upper ,

w all o f the embryo an d at th e floo r o f t he c avity is a l arge

,

m ass o f yolk cells Th e a rch e n tero n r eprese n ts th e c avity

.

produced by th e process o f gastrul at io n I t is due in gre a t .

,

m e asure at le ast to th e overgro w th o f th e dors al lips of t h e

,

bl astopore the c ells formi ng th e floor bei n g form erly at th e

,

su rface of the egg Accordi n g to M arsh all the c avity

.
,

arises in gre a t p ar t through th e splitti n g ap art of t h e yolk

cells but while th is m ay be a factor in the c as e i t cert ai nly

, ,

c a nno t be t h e predomi nan t o n e ( S ee Robi n so n an d Assh e .

to n Assh et o n M org an As the arche n tero n

i n cre ases in size th e bl astoc oel or segm e n t at io n c avi ty n ec es
,

sarily becom es sm aller Accordi n g to M arsh all th e form er

.

bre aks thr o ugh i n to th e l at t er an d t h e two form o n e c avity ,

Th e G erm L ayer s — Th e form a ti o n of th e g astrul a pro

duces a two l ayered e mbryo e ac h l ayer bei n g several cells
-

1 Ro bi s
n o n an d A ssh et o n , Q u ar t f o u r M ic
. . . S ea V o l 3 2
, .
,
1 891 .

2 A ssh e t o n [ bid V ol 37 . 1 8 94
, .
,

3 M or g an , T h e D e ve l pmo e nt o f th e F gs ro
’
E gg , 1 8 97 .
96 TH E B IOLOGY OF TH E F ROG CH A P .

thick The ou ter o f th ese l ayers is th e ectoder m th e i nn e r

.
,

the en toder m Th e cells o f th e form er are sm all an d pig

.

m en t ed ; thos e o f the l atte r for the mo st p art are com p ar a

t ively l arge ligh ter in colo r an d co n t ai n a l arge a mou n t of
, ,

yolk The two l ay ers are co n ti n uous wi t h e ach oth e r at t h e

.

lips of th e bl astopore B efore th e proc ess of i n v agi n a tio n

.

FI G 2o .
— S agi tta l s ci
e t on t h ro it g h a f g mb y B bl s t c l ; B P
ro e r o . a o oe ,

d s l lip f bl s p ; B P l l p f bl s t p ; E E p i d mic
.
,
’
or a o a to o re v e n t ra i o a o o re ,
e er

l y f c d m; EN i us l y f c t d m ; H hy p
,

a er o e to er , nn e r o r n er vo a er o e o er o

bl s t d m; T m s g s ul c i y ; Y y lk p lug
,

a or en t o er ,
e e n t e ro n or a tr a av t ,
o .

(Af M sh ll )
t er ar a .

is co mpl e t ed t here a ppe ars a t hird germ l ayer th e m esoder m ,

o r m eso bla s t bet w ee n th e o t h er two The m esoderm ap

, .

p e a rs a ll arou n d t h e bl as t op o re a n d a s this o pe n i n g closes ,

m ai nly from in fro n t b a ckw ard the t w o m asses of m es o ,

der m o n ei t her si de are brough t n e ar e ac h o t her in the m id

d o r s al li n e The fre e ve n t r al e dges of th e m asses o r sh eets
.
98 THE BIOLOGY OF TH E FROG CH A P .

ver t ebr a l colum n an d o ccu rri n g in t h e e mbryo w he n

, ,
n ot

a lso prese n t in the adults of a ll V C l t e b l Lie a n i m a ls ,

’

- . It

FI G 22T s s s c t i t h ugh ra n ver e e on ro a f g mb y b f th cl su f

ro e r o e o re e o re o

m dull y u l f lds C c l m b dy c i y ; CH
. .

th e e ar or ne ra o oe o or o av t ,
n o to

ch d ; E E p id m i c l y f c d m; E N us l y f c t
.
,

e er a er o e to er n e rv o a er o e o
or ,

d m M m s d m; ME u s m t ic m s d m M H i
,

er e o er o ter o r o a e o er n n er o r
;
.
,

lp ch ic m s d m ; NC u l g ; ND d s l t f sp i l
, ,

e o ne ra ro o ve or a ro o o na
S an n er ,

sp i l c d ; T h ; W li di t iculum ; Y
,

; NS
n erve na or a rc t
e n e ro n ,
ve r ver ,
, ,

y lk (A f M sh ll )
o . t er ar a .

is lw ays t h e fi rst p art o f th e skel e t o n t o m ake i ts appe ar

a

a n ce in t h e e mbryo as i t w a s t h e firs t p ar t t o a ppe a r in t h e

,

evolu t io n o f t h e r ace Whe t h er in t h e frog it i s e n t o dermic

.
 TH E D E V E L O PM E NT OF TH E FROG 99

in origi n as i t cert ai nly is in som e of the A mph ibi a an d in

,

m an y o t her vertebr ates or wheth er as m ai n t ai n ed by M o r

, ,

an it i s developed from th e m eso derm is a m atter abou t

g , ,

wh ich th ere i s a differe n c e of opi n io n M iss H D K i n g 1

. . .

h as rece n t ly s t udied th e form a tio n of the n otochord in B ufo

len tigin os u s a n d R a n a p a lu s tr is an d has com e to the c o n ,

c lusio n th at th e n o t o chord in the an t erior en d o f th e

e mbryo arises fro m th e m esoderm w here as in the poste r ior ,

p ar t of th e e mbryo it is developed fro m both m esoderm an d

e nt oderm .

E xt ern al Ch an ges — A t th e ti m e wh e n the bl astopore

i s n e arly close d th e egg i s s t ill in a spheric al form excep t ‘

th at alo n g w h a t i s to be the dors al sid e o f th e body of t h e

e mbryo t here is th e begi n ni ng of a bro ad depressio n k n ow n
as t h e p r im itive g r o ove O n ei t h er sid e of this are two
.

folds th e i n n er an d t h e ou te r m edulla ry folds which are

, ,

co n t i n u ed as an elev atio n arou n d th e an terior en d of th e

pri mitive groove an d are produc ed b ackw ard o n ei ther side
o f t he bl as t opore Th e ou t er m edull ary folds gr adu ally
.

fad e aw ay bu t th e i n n er o n es b eco me elev ated an d arch

,

over t h e groove betwee n t hem F i n ally th e two i n n er folds .

m ee t an d fus e al o n g t h e m edi an li n e c o n verti n g th e groove ,

i n t o a tube The poi n t w here th ey first fuse correspo n ds

.

t o the n eck regio n of the e mbryo an d the closure of th e tube

proceeds bo t h forw ard an d b ack w ard fro m this poi n t Th e .

fusio n ex t e n ds b ack w ard so t h at folds o n ei t h er side o f the

bl as t opore close in abo ve th at ope n i n g in suc h a w ay t h a t it
becomes n o lo n ger visible fro m th e outside As th e m edu l .

l ary tube i s co m pleted i t is co n stric t ed off fro m the ecto ~

derm above an d t h e l at t er becom es co n ti n uous over the

,

mid dors al li n e S ubseque n t ly it d evelop s i n to th e br ai n

-

an d spi n al cord of the embryo .

1 K i g
n ,
B io l . B u ll V ol 4 .
,
1 9 03 .
 TH E B I O L O G Y OF TH E F RO G C H AP .

As th e above ch anges are t aki n g pl ace th e e mbryo

elo n g a tes in the directio n of the n eu r al tube w hich m arks ,

th e lo ngitudi n al axis of t h e fu t ure an i m al O n ei t her sid e .

of the an terior e n d of the n eu ral tube there appe ars a p ai r

FI G 3. 2 . l pm D e ve f th
o mb y A lk p lug s g ; B sh i g
ent o e e r o . , yo -
ta e ,
ow n

th m dull y f lds h bl s p
e e ar o t ly cl s d d b l t h l t
e a to o re n e ar o e an e ow e at e r

i gi t i h i ch is t f m t h us ; C D l t s t g s ; E h
, ,

th e n va na on w o or e an a er a e ,
t e

m dull y f lds h g g h dc d th bl s p Ab
, ,

e ar o ave ro w n to et er an o ve re e a to o re . o ve

th us i s th udim
e an f t h t il
e r (F m M g
en t o e ft Z i gl )
a . ro or an , a er e er .

of thicke n i n gs of t h e ectod erm Th e an terio r m embers of .

e ach p air th e sense p la tes grow forw ard an d meet in fro n t

, ,

of t h e en d of th e n eur al tube a d epressi o n appe ars in e ach

pl ate and m ark s the b egi nni ng of the ven tr a l s ucker of the
1 02 TH E B I OLOG Y OF TH E F RO G CH A P .

bl astopore w as closed over L ate r this i nv agi natio n m eets

.

a n d fuses wi t h a diverticulu m fro m t h e pos t erior p a rt of th e

arche n tero n thu s es t abli shi n g an ope n i n g bet w e e n th e l a t t er

,

a n d the exterio r The t ail arises as an elev atio n of th e

.

regio n in fro n t o f th e bl astopore which grows b ackw ard an d

,

pushes the an us to a more ve n tr al positio n L ater i t b e .

co mes fl atte n ed fro m sid e to side an d its upper an d lower ,

e dges beco m e produced i n to a th i n exp an sio n or t a il ,

fin .

The n os tr ils app e ar as a p air o f exter n al d epressio n s or

pits a li t tle above th e rudi me n t of t h e m o u th Th ese pits .

deepe n an d fi n ally com m u ni c at e w i t h t h e bucc al c avity

, .

Above an d to the side s of t h e n as al pits th e begi n ni n g of

th e ey es i s i n dic ated as a p air o f t hicke n i n gs o f th e ec t o
derm Th e ou tli n e of t h e e n l arged an t erior portio n o f t h e
.

m edull ary tube m ay be observe d fro m th e surfac e I t is .

be n t dow n w ard in fro n t an d sh ows a divi sio n i n to t h ree

,

regio n s w hic h becom e th e th ree prim ary vesicl es o f the

,

br ai n Ne ar th e pos t eri o r o f t h ese vesicles th ere is d evel

.

oped o n ei t h er sid e an i n v agi n a t io n or p it of t h e ec t od erm ,

whic h fi n ally si n ks in an d becom es cu t off fro m th e surface

an d fo r ms th e vesicle o f th e in n er ea r .

At th e t i me th e n eur al t ub e is forme d t h e superfici al c ells ,

o f t he ectoderm becom e fur n ish ed in m any pl ac es wi t h cili a

by me an s o f which th e e mbryo slowly ro t at es withi n t h e
j elly Th e ge n eral directi o n o f the s t roke of the cili a is
.

fro m before b ackw ard Th e m oveme n t i s stro ngest at

.

the an terior en d o f t h e body an d i s we aker o n t h e ve n tral

,

th an o n the dors al side “ A t adpol e of 6 or 7 mm will

. .

progress i f pl ac ed u po n i ts side in w ater alo n g the bo t to m

, ,

of a flat gl ass vessel at t h e r a te of o n e millim eter in fro m

,

four t o seve n seco n ds ( s

A sh.
”
et o n Aft er th e t adpole

is h atched from th e j elly th e cili a gr adu ally dis appe ar .

 TH E D E V E LO PM E NT O F TH E FROG 19 3

O rg an s f r om t h e E ct oderm dditi o n to formi ng

.
-
In a

the ou t er l ayer o f the ski n over t h e e n tire surfaCe o f t h e

e mbryo th e ectoderm gives rise to cert ai n oth er s t ruc t ures
which com e to lie w i t hi n the b o dy C hie f am o n g th ese i s .

t h e ce n tr al n ervous sys t e m w hos e begi n n i n g in th e m edu l

l ary groove h as alre ady bee n described The n eur al t ube i n t o .

which t h e medull ary groove d evelops loses its origi nal c o n

n ec t io n wi t h the surfac e ; an t eriorly it beco m es e n l arged
an d forms the br ai n th e rem ai n i n g por t io n s developi n g i n to
,

th e spi nal cord The t hicke n i n g o f t h e w alls o f the portio n

.

o f th e tub e which forms t h e cord di mi n ishes t h e ce n t r al

c avity u n til i t becom es reduced to a fi n e c an al k n ow n in ,

t h e a dul t as the ca n a lis cen t r a lis Th e an terior porti o n of

.

t h e t ube beco mes divide d by slight co n stri c t io n s i n t o t hree

vesicles wh ich form desig n ati n g th e m from be fore b ack

, ,

w ard the fo r e m id an d lz in d b r a in
, , ,
Th e hi n dbr ai n .

beco mes wid e n ed fro m side t o sid e especi ally in fro n t ; its ,

flo or an d sides t hicke n but the roo f excep t for a sm all fold

, ,

a t the e n d wh ich d evelops i n to t he cer ebellu m rem ai n s ,

thi n an d m embran ous an d beco m es t hrow n i n t o a series of

,

folds which support a m ass of blood vessels k n ow n as th e

clz o r o id
p lex u s Th e
. por t io n o f t h e hi n dbr a i n wh ich does

n o t form th e c erebellu m is co n verted i n to t h e m edu lla .

The ce n t r al c avi ty beco m es wide n ed ou t formi n g the f o u r tlz ,

ven t r icle which com mu n ic a tes pos t eriorly w ith th e c an ali s

,

ce n t r alis o f th e cord an d an teriorly w i th the ve n tricle of th e

midbr ai n .

The midb r ain grows out dors ally and l ater ally i n to a p ai r
o f hollow processes th e op tic lobes whose c avities or v e n tri
, ,

cles com mu n ic a te w ith t he m edi an c an al which becomes ,

n a rrowed by the thicke n i n g of its w alls an d forms the a q u e ,

du ct of S y lv iu s or iter a ter tio a d g u a r t u m ven tr icu lu m

,
The .

floo r o f th e mi dbr ai n forms the cr u r a cer ebr i .

 TH E BIOLOGY OF TH E FRO G CH A P . V

The fo r eb r a in so o n beco mes sep ar a ted i n to t w o p ar ts ,

t he tlz a ls za lo n behi n d a n d the cer ebr a l lz em is lz er es

en ce /
p , p ,

which grow out fro m t h e l atter in fro n t Th e floor an d .

w alls o f th e form er beco me thi ck e n ed to form th e op tic tb a

la m i the roo f re m ai n s th i n an d m e mbr an ous an d the c avity
, ,

beco mes the tb ir d ven tr icle F ro m th e roo f of th e th al a

.

m en c ep h alo n there a rises a medi an hollow ou tg r owth t h e ,

in ea l g la n d whic h ex t e n ds dors ally re a ch i n g th e surfa ce

p , ,

ectoderm where i t beco m es exp an ded i n to a sm all k n ob

,
.

The k nob beco m es co n s tric ted o ff w h e n th e bo n es of th e

skull d evelop an d fo r ms th e brow spo t previously described ,
.

Th e floor o f the th al a me n ceph alo n gives rise to a hollow

outgrowth t he inflz n dibu lu m which exte n ds dow n w ard I t
, ,
.

com es i n to close co n t act w ith a n o t her s tru ctur e t he p it u ita ry ,

body which i s d eveloped fro m th e ec t od erm of t he dors al

,

w all of th e sto m od eum Th e sid es o f t h e t h al a me n ceph alo n

.

give ris e to a p air of l ater al diver t icul a th e op tic ves icles , ,

which grow ou t u n til they c om e in close co n t act wi t h the

surface ectod erm Th e dist al e nd o f the vesicles wide n s
.

ou t to form the r etin a o f th e eyes th e st alk givi n g rise to ,

t h e op tic n er ve .

Th e an teri or w all o f th e forebrai n prod uces a p air o f

pouch es t h e c erebr al h em i sph ere s which fi n ally beco m e
, ,

t h e l a rgest p art of th e br ai n Their c avi t ies t h e la ter a l ven

.
,

t r icles com mu n ic ate wi t h th e third ve n tricle by an op e n i n g

, ,

t he fo r a m en of M on r o .

The n erves arise as p aired ou tgrowths bo th fro m the br ai n

an d cord pushi n g th eir w a bet w e e n t h e cells o f t h e other
, y
o r g an s dividi n g an d r ami fyi n g as they push ou tw ard
, ,

t ow ard th e v a riou s p arts they su pply Th e spi nal n erve s .

begi n as two i n de pe n d e n t O iit gro w t h s repres e n ti ng th e ,

dors al and ve n t r al roots ; th ese soo n u n ite i n to a si ngle

n erve .
1 06 TH E BIOLOGY OF TH E F ROG CH A P .

The li n i n g of the m ou th c avity i s for med from an invagin

a tio n O f ecto derm th e s tom o deu m which pushes in u n til i t
, ,

bre aks through i n to the arch e n t ero n A simil a r ectoderm al .

i nv agi n atio n t h e p r octodeu m forms th e li n i ng o f a sm all p art

, ,

o f the posterio r e nd o f th e ali m e n t ar y c an al Th e le n s an d .

FIG . 26 .
—C ss s c t i
f g mb y A R ch t ; M S m s
ro e on of a ro e r o .
, ar e n ero n ,
e o

bl s t ic s mi s ; N
a o ch d ; NS u l c s ; lll m dull y ub ;
te n o to or ne ra re t e ar t e

s ; S N sub ch d l d ; S O S P s m ic d p l h
, , ,

PR p ph ro n e ro noto or a ro o at an S an c

i m s d m (F m M g ft M sh ll )
, , , ,

n c e o er . ro or an , a er ar a .

cor n e a as w ell as th e reti n a of th e eye an d th e vesicle o f ,

t h e i nn er ear a l so t ake t h eir origi n fr o m this l ayer

,
.

O r g an s f r om t h e E nt oder m — Th e e n t o derm o r th e .
,

germ layer w hich is i n vagi n ate d w i t hi n t h e egg gives ris e t o ,

th e li ni n g o f t h e ali me n t ary c anal and of all org an s which

 TH E D E V E LO PM E N T OF TH E FRO G 1 07

a rise as out grow t hs fro m it Th e first of these to be formed

.

i s the liver which at t h e begi n n i n g appe ars as an ou tpock et

,

in g of th e ve nt r al sid e n e ar t h e an terior e n d Th e ou t .

pocketi n g becomes fo ld ed an d br an ched bei ng co n verted ,

fi nally i n to a n u mb er o f clus t ers of tubules all emp t yi n g ,

i nto the co mmo n c an al th e bile du ct which is produced

, ,

by a le ng t he n i n g of th e n eck of th e origi nal ou tgrowth A .

l at er al ou t growth o f t h e bile d uct forms th e g a ll bla dder .

The c ells li n i n g the t ermi nal br an che s of t h e hep atic dive r

ticul a beco m e th e secreti n g cells o f t he liver The co nn ec .

tive t issu e blood vessels an d outer co ati n g of the liver are

, ,

d erived fro m th e m es o bl ast .

Th e p a n cr ea s arises much in t h e s am e w ay as th e liver ,

bu t as a p air of outgrowths i n ste ad o f a si n gl e o n e They .

form however a S i n gle org an an d t h eir ducts l at er becom e

, , ,

co nn ec ted wi th the bile duc t O nly th e secreti n g portio n .

o f th e p an cr e as an d the li n i ng of its duc t s ar e of e n t od ermic

origi n th e co nn ec t ive tissu e blood vessels etc arisi n g as
, , ,
.
, ,

in t h e liver fro m th e middle germ l ayer

,
.

Th e bla dder aris es as an outgrowth o f the ve n tral sid e

of t h e ali me n t ary c an al n e ar t h e posterior e n d ; its li n i n g
, ,

th ere fore is o f e n t odermic origi n

,
.

Th e lu ng s appe ar as a p air o f pouch es from the sides o f

the e soph agus Th ey m ake li t tle grow t h u n til qui t e l ate in
.

the life of th e t adpole The regio n of th e eso p h agus from

.

which the lu ngs a rise bec o m es d epressed an d p artly sep a

r at ed off from th e p art above t o form th e la ry nx t h e m ou t h ,

o f th e d epressed portio n goi ng to form the g lo ttis of the

adult .

Th e gill slits in the frog appe ar in the form o f five solid

ou t growths o n e ach side o f th e an terior portio n o f the
a rche n tero n I n secti o n t hey a r e sh o w n t o be in th e form
.

o f a double fold such as would be produced if th e w alls of

1 10 TH E BIOLO G Y OF TH E FROG CH A P .

t ary c anal diverticul a Th e co n n ective tissu e mus

an d it s .
,

c u lar an d p erito n e a l l ayers a re derived fro m t h e m esobl a st

,
.

F o r the most p art it i s co mpose d of bu t a si n gl e l ayer of

cells The epi t h eliu m o f t h e mou t h an d a sm all portio n of
.

th e clo ac a are produc ed by t h e ectode rm these bei n g th e ,

o nly portio n s o f the li n i n g o f th e ali me n t ary c an al n o t of

e n todermic origi n .

O r gan s f r om t he M esoderm — The developme n t o f the.

m esoderm h as bee n tr aced to th e s t age in w hich i t co n sists

of two double l ayered sh ee ts o f t i ssue exte n di ng fro m th e
-

n otochord above to th e ve n t r al si de of th e body Th e two .

sh eets of m esod erm are sep a r at ed by the n otoch ord excep t

for a shor t dis t an ce in fro n t o f and behi n d this structure ,

w here th ey beco m e co n ti n uous a cross the m iddle li n e A .

divi sio n soo n occurs in t h e m esoderm sep ara ti ng a dors al ,

portio n k n ow n as the ver tebr a l p la te fro m a ve n tral p art

, , ,

c alled t h e la ter a l p la te The form er becomes divid ed tr an s

.

ve rse ly i n to a n u mber of blocks c alled m o to m es or m u scle

y ,

seg m en ts E ach o f t hese beco m es thicke n ed S O th at the c en

.

t r al c avi t y becomes reduced in size a n d fi n ally dis appe ars .

Th e divisio n o f the vertebral pl ate i n t o segm e n ts begi n s in

t h e n eck regio n of t h e e mbryo an d proceed s b ackw a rd .

Th e segme n ts soo n becom e sep ar ated fro m e ac h oth er by

sept a of co nn ective tissue which assu m e the form o f a V
wi t h its ap ex poi n t i n g tow ard the an terior e n d o f t h e body .

The myo t om es are e asily se e n at th e sid es of th e body of a

you ng t adpol e especi ally in th e regio n o f the t ail Th e cells
,
.

o f the myo t o mes elo ng ate in a directio n p ar allel with t h e

lo n g a m of th e an i m al an d become co n verted i n to muscle
fibers .

The two l ayers o f the l ater al pl ates becom e widely sep a

r ated by th e e nl argem e n t o f t h e i n terve n i ng bod y c avi t y
o r c oelom The i nn er or sp la n c/z n ic lay er beco mes closely
.
 TH E D E V E LO PM E NT OF TH E F ROG 1 1 1

a pplied to th e e n toderm o f th e a rch e n t ero n an d forms t h e

supporti ng tissu e an d muscul a ture of th e alim e n t ary c an al
an d its diverticul a Th e ou ter or som a tic lay er comes to li e
.

ag a i n st th e ou ter ecto derm an d forms th e i n n er portio n

( co n n ective tissu e muscle ,
an d p erito n eum
, ) o f the body w all .

The i nn ermost por t io n of both th e so m ati c an d spl an ch n ic

l ayers of m esobl ast becom e differe n ti ated as a sep arate
l ayer the p er itoneu m which is co n ti nuou s all arou n d th e bo dy
, ,

c avi ty As th e right an d left h alves o f t h e coelo m arise i n d e

.

pe n de n tly an d gr adu ally exte n d tow ard t h e mid v e n t ral li n e -

th ey are sep ar ated for a tim e by a medi an ve n tral p ar t i t io n .

This subseq u e n tly bre aks do w n alo n g m ost o f t h e l e n g t h of th e

ali me n t ary c an al pu tti n g th e t w o si des o f th e body c avi t y in
,

co nn ectio n with e ach othe r The m edi an p arti tio n persis t s

.
,

however for a S hort d ist an ce an teriorly formi n g t h e ve r tic al

, ,

me mbran e which exte n ds fro m the liver an d p eric ardiu m

to th e ve n tral body w all A still sm alle r portio n occurs
.

be twee n th e body w all an d th e ve n tr al sid e of th e c lo ac a .

The b eu r t an d p er ica r diu m t ake their origi n from th e

m esoder m n e ar th e an terior en d of the ve n t r al sid e of t h e
body A p ai r o f fissures appe ars in the sh eet of m esoderm
.

in this regio n ; these gr adu ally e n l arge a nd exte n d tow ard

the middle li n e Th e l ay e r roofi ng over t h ese fissures b e
.

comes raised u p o n eith er si de an d the two folds thus formed,

m e et e ach oth er above formi ng a sort of tube With i n thi s

,
.

t ube ar e i n closed som e sc at t ered cells whic h arr an ge the m

selves i n to a l ayer th a t beco mes th e e nd o t h eli al li n i n g of t h e

‘

h e art Th e c avity ou tside t h e t ube beco mes t h e c avity of

.

t h e peri c ardiu m an d th e t ube i tself thi cke n s an d becomes

,

t r an sformed m ai n ly i n to the h e ar t bu t it s outer l ayer gives

,

rise t o a t hi n shee t o f t issu e t h e viscer al portio n of th e peri

,

c ardiu m The tissue which a t first co n n ects t h e h e art wi t h

.

the ve n tral sid e o f the peric ar di um becom es broke n t hro ugh ,

 TH E BIOLOGY OF THE FROG CH A P .

a nd t h e sid es o f t h e peric ardi al c avi t y becom e co n t i n u

tw o

ous ; t h e dors al co n n ec tio n of t he h e art dis app e ars at a

l at er p eriod The vi scer al l aye r of peri c ardiu m whic h closely
.

i n vests the h e art becom es refl ected u po n th e sides o f th e

surrou n di n g c avity w h ere i t be com es co nti n uous wi th the
,

p ari e t al l aye r t h e rel atio n s of t h e t w o p arts bei ng esse n ti ally

‘

th e s am e as th at o f th e portio n of pe ri t o n eu m surrou n di n g
the ali m e n t ary c anal and t h at li n i ng the c oelo m O wi n g to .

FI G . 28 . A B C t h g s i h d l pm
re e s t a e n t e eve o ent o f th e h ea rt . E en d o

h l i um ; P E i c dium ; P H p h y x ; W ll h f
, , , ,

t e , p er ar ,
ar n , w a of eart .
( er M o r
A t

g an .
)

i ts i n cre ase in le n g t h th e h e art becom es be n t in th e form of

an S ; a n t eriorly it beco m es co n t i nu ed i n to th e t r u n cus
a r ter ios u s w hich divid es i n t o two br an c hes w h ich proceed
,

t o w ard t h e gills wh ere t h ey bre ak up in to th e a o r tic a r clz es

, ,

w hich dis t ribute br an ch es t o t h e gill fil ame n t s Th e bl o od .

vessels firs t appe ar as l acu nae o r sp aces betwee n t he cells o f

1 14 TH E BIOLOGY OF TH E FROG CH A P .

FIG 29 H or iz
h ugh
o n ta l s ct i
d c d mb y e on ch t ro an a van e e r o . ar en

b ch l ch s gill
. .

t
e ro n
; E R i Q ; H H H l ; H 8
ra n a ar e 1
,
2
,
3
,
S l tS ,

hy d sli ; H M h y m d b l cl f ; H Y hy id ch I N i f d b
,

01 t o an l u ar e t ,
o ar ; ,
n un i u

lum ; O F lf c y p it ; O S p ic s lk ; P p p h s ; S s gm t l
,

o a to r o t ta ro n e ro ,
e en a
,

duc t ( F m M g f M sh ll )
, ,

. ro or an , a te r ar a .
 TH E DE V E L O PM E N T OF TH E F RO G 1 15

l ate r op e n i n to the c oelom by fu el sh aped cili ated mou ths nn -

or n ephrostom es bu t th e l at ter soo n lose th eir co n n ectio n

,

with th e t ubules an d a cquire seco n d arily an ope n i ng i n to

the br an ch es of th e re n al vei n s in the ve n tr al p art of the
kid n ey The tubules in cre ase to a very l arge n u mber an d
.

become richly supplied with blood vessels ; th ey form wi th

the co nn ective tissue which bi n ds t hem togeth er a co mp act
m ass whic h assum es th e form of th e kid n ey o f the adul t .

T h e segme n t al or pro n ephric d uc t w hich served as the out

let of the pro n ephros is worked in t o form t h e Wolfi em a z/ef ’

or u r efer of th e adul t The M ulleri an du ct w as formerly

.

supposed to arise by a spli t ti n g of t h e segme n t al du c t bu t ,

a ccordi n g to M ac B ride M arsh all G emmill an d m ore

1 2
,
3
, ,

rece n tly H all it d evelops qui te i n depe nde n tly of th at

4
,

s tructure .

The reproductive org an s fi rst appe ar as ridges of th e

peri t o n eu m n e ar the b as e of t h e mese n tery ( M arsh all )
As th e ge n i t al ridges i n cre as e in size t h ey become co n
s t ric t e d at their po i n ts of a t t ach me n t an d fi nally h an g ,

supported by a p eri to n e al m embr an e I n th e m ale th e .

tes t is becomes co n n ecte d wi t h t ubes w hich grow ou t of t h e

re nal tubul es an d form t h e v as a effere n t ia Th e ge n i tal .

ridges in t h e two sexes h ave a si m il ar a ppe ar an ce u n til n e ar

the close of l arv al li fe wh e n those o f th e fem al e u n dergo a
much more rapid growth .

Th e begi nn i ng o f th e vertebr al c olu m n i s represe n ted by th e

n otochord bu t this struc t ure forms but a rel at ively sm all por
,

tio n o f t h e b a ckbo n e of t h e adul t frog L oose mesodermic .

cells or m ese n chyme produced from t h e p eriphery of the

, ,

1 M ac B ride Q u ar t f ou r Al ia 5
, . . . 62 V o l 33 .
,
1 8 92 .

2 M ar a sh ll
“
,
b mb y l gy
V e rt e rat e E r o o .

3 Ge m mill A / f A t P /y m Phys Ab th
,
re L . un . u . z . .
,
1 8 97 .

4 H all B ll M C mp 2 1 H
,
u . d V l 45
u s. o . 00 . a r v ar ,
o .
,
1 9 04 .
1 16 TH E BIO L OG Y O F - TH E F RO G CH AP .
'

somite coll ect arou n d th e n otochord formi n g a tubul ar in

, ,

vestm e n t F ro m th e dors al sid e of this m ass ridges or folds

.

grow up an d surrou n d t he spi nal cord Th e mesod erm c o y .

eri ng the n o toch ord th e n beco mes divid ed by tr an svers e se pt a

which alter n ate w i t h those be twe e n th e so mites bu t t h ese do ,

n o t c u t across th e n o toc hord i tself Th e segm e n ts they cu t .

o ff represe n t t h e ver t ebr ae they so o n becom e c artil agi n ou s ,

a n d fi n ally ossi fy Th e c artil agi n ous sh e at h grows i nw ard

.

a t t h e e n ds of the vertebr a e co n stricti n g an d fi n ally cu t t i n g

,

through t h e n otochord so th at in t h e adul t all th a t rem ai n s

,

of t his structu re are sm all po r t i o n s i n closed withi n the ce n tr a

o f th e vertebrae .

M et amorph osis — At the ti m e of h atchi n g th e t adpole i s

.

a fi sh lik e cre a ture h avi n g a lo n g vertic ally fl a tte n ed t ail by

-

, , ,

m e an s of whic h i t s w i ms through th e w ater Th e side s .

of th e t ail sho w th e m arki ngs o f t h e m uscle segme n ts

t h rough th e ski n The fl a t t e n ed exp an sio n s o f the in t egu
.

m e n t o n th e u pper an d l o w er sid es of t he t ail are thi n an d

n e arly tr a n sp are n t so t h at o n e m ay e asily observe wi t h a
,

m icroscope the blood flo w i n g in th e c apill ari es .

Th e m outh bre aks through i n to th e arch e n tero n a few

d ays aft er h atchi n g th e l arv a previous to this tim e livi ng
, , ,

a t th e expe n se of the food yolk in th e ali m e n t a ry c an al .

The i n tes t i n e i n cre ases very r apidly in l e n gth an d beco m es ,

coil ed in th e form o f a spiral which m ay o fte n b e see n ,

t hrough t h e ve n tr al body w all Th e exter nal gills grow

.

r a pidly a ft er t h e t ad p ole is h a tc hed a n d soo n are co n ver t ed ,

i n to lo n g br an chi n g t uft s Three p ai rs of ex t er nal gills are

,
.

developed t h e po s t erior p air m aki n g its a ppe ar an ce after

,

t h e fi rs t t w o The gill slits gro w abou t th e t i m e th e mou t h

.

is fully form ed an d t h e w ater w h ich i s t ake n in a t th e m outh

,

is p assed th ro ugh t h e gill sli t s to th e e xterio r I n a dd itio n to .

the exter nal gills t here are d evelo pe d somewh at l ate r fou r
1 18 TH E B I OLO GY O F TH E F RO G CH AP
.

p airs of wh ich are produce d by foldi ngs of the

m e mbr an e li n i n g th e gill slits B oth exter nal an d i n ter nal
.

gills r eceive an abu n d an t blood supply fr o m the vessels th at

fo r m th e ao r ti c or b r an chi al ar ches Th e dis appe ar an ce of
.

th e exter nal gills is associ ate d with th e growth o f a fold ,

the op er cu lu m which arises o n eith er sid e of the he ad an d

,

gr adu ally exte n ds b ackw ard Th e fre e posterio r edge of th e

.

fold fuses with th e body behi n d and below the gill regio n ,

le avi ng o nly an ope n sp ace o n t he left side of th e body ,

which is k n ow n as th e sp ir a cle Th e w ate r which p asses

.

ou t o f the gill sli t s com es i n to a ch amber bou nd e d exter

n ally by t h e ope rcul a r w all an d th e nce p asses th r ough th e
,

spir acle to the ou t side S oo n after the co mpleti o n of t h is

.

ch amber th e ex t er nal gills dis app e ar an d th e i n te r n al gills

fu n ctio n in th eir ste ad .

Th e j aws of th e t adpol e are fur nish e d wi th hor n y co ati ngs

.

which fu n c t i o n as te eth bu t these are shed in l ater l arv al

,

life I n addi t io n both uppe r an d lower lips also co n t ai n

.

tr an sverse rows of fi n e tee th wh ich v ary in n umbe r an d

,

arr an gem e n t in the differe n t speci es A r ou n d th e ou tsid e

.

of th e lip th ere are n um erous sm all p apill ae which also v ary ,

co n side r ably in t ad poles o f d iffere n t sp ecies of frogs Th e .

n as al pi t s d o n o t bre a k through i n to th e mou t h u n til som e

t i me a ft e r h at chi ng Th e eyes are si tu ated o n th e do r s al

.

sid e o f the h e ad an d lo ok obliqu ely upw ard There are

,
.

sever al ro w s o f se n se organ s o n th e ski n o f th e t adpol e ,

b u t th es e dis a ppe ar wh e n the an i m al assum es a te rres t ri a l

mod e o f li fe The ve n tr al sucker in th e r ece n tly h at che d
.

l arv a is in the form o f a horsesho e Th e ectodermic c e lls

.

coveri n g it are p ar t ly gl an dul ar and t hey form a muco u s

,

secre t i o n by m e an s of which th e l arv ae adh ere to v ario u s

obj ec t s L ater in l arv al life t h e sucker becomes divided in
.

two in the middle Th e t w o p arts beco m e c arried far th e r

.
 TH E D E V E LO PM E N T OF TH E F RO G 1 19

b ac k o n th e ve n tr al sid e of the he ad an d gr adu ally d ecre ase ,

in size an d fi n ally dis appe ar

,
.

Th e hi n d li mbs which are th e first o n es to appe ar bud

, ,

o u t as sm all p apill ae o n e i t her side of t h e b ase of th e t ail .

They gr adu ally i n cre ase in si ze become j oi n t ed in struc t ure , ,

a n d l ater bu d ou t th e toes at the

-
dist al en d The fo r e .

limbs develop in much th e s a me m an n er ; th e left li mb

p asses through the spi racle th e r igh t o n e pushi ng through ,

t he w all of th e o p ercu lu my

Towar d th e end of the l ar val pe riod th e t ail begi n s to dis

a ppe ar its tissues bre ak dow n an d are r esorbed servi n g , ,

d oub t le ss as food m ateri al for b uil di ng up th e oth er org an s

,

of t h e body D uri ng th e tr an sform at io n of t h e t adpole i nt o

.

the you n g frog th e i n testi n e shorte ns ; th e m ou th beco mes

,

much wider and th e h or n y j aws ar e shed th e to n gue in

, ,

c r e ases gre atly in size t h e legs grow r apidly th e rou nded , ,

body ch an ges in form an d th e gills becom e reso rbed th e ,

lu ngs the n d evelop r apidly and the t adpole freque n tly comes ,

to the surface for air .

The food o f th e t adpol e is m ai n ly veget abl e m atter .

S pirogyra an d oth er alg ae are commo n a r t icles of die t ;

an im al foo d howev er is gre at ly relished
,
T ad poles w ill
,
.

feed e agerly o n dec ayi n g i n sec t s e ar t h w orms o r al most any

‘

, ,

ki nd of me at Th ey will also eat bre ad or frui ts there are

.

few th i n gs app are n tly in the w ay o f food wh ich they

, , ,

disd ai n .

RE F E RE NC E S

m s t c m pl t cc u t s f t h d l p m t
T he o o e e a o n o e eve o en of th e fro g ar e co n

t i
a ne di M g n
“
s b k Th D l pm t f t h
or an
’
oo , e ev e o en o e F gsro
’
E g g, ”
an d

M sh ll s V t b t E mb y l g y
ar a
’
er e Am c d ra e r o o .
”
o re on en e s d cc u t is t
a o n o

be f u d i th
o n sm l l k “T h F g by t h
n e a w or on e ro ,
”
e l tt uth
a er a A or .
,

g l l m
e n e ra t
e ey d sc i p t i f t h d l p m t
e n ar e r on o e ev e o en o f th f g b s d
e ro , a e

m i ly t h
a n on k f M sh ll is c t i d i R s
e w or o ar a , on a ne n ee e
’
s I t duc t i t
n ro on o
1 20 TH E BIOLOGY OF TH E FROG CH A P .

Ve b t E mb y l gy L is t s f t h m s t im p t t li t t u
rt e ra e th r o o . o e o o r an era re o n e

sub j c t i ll h f u d i th
e k s f M sh ll d M g jus t f
w e d o n n e w or o ar a an or an re e rre

to T h f ll
. i g p p s d l i t h t h ch c t is t ics d m t m
e o ow n a er ea w e ara er an e a or

ph f t dp l s
o ses o a o e

B arf rt h D V such ub di V
u d lu g d F hl
, . er e er e erw an n er ro sc a r ve n .

A n at A n z . .
, Ed . 1 .

B ou len ger, G A .
. A Sy nop sis of th e T dp l s
a o e of E u ro
p ean B at ra
ch i s P c an . ro . 20 6 1 S o c L o n . . d o n, 1 89 1 .

Ca m er ano, L . O b se rva t io n i sui gi i i d gli r n e A n fihi an ui r . Bo ll .

M us . T i or n o, 8, 1 893 .

Cop elan d, E . B . H et e r o g e n eo us In duc t i on in T dp l s

a o e . S ci c
en e,

11 . s .
, V ol . 1 3, 1 90 0 .

H in k ley , M . H . On So m e if
D f e ren c s e in the Mo ut h S tr uc t u f re o

T dp l s
a o e of t he A n u usro B at ra ch i s f u d
an o n at M il t o n, Ma ss P c . ro .

B os . S oc . Na t . H is t V ol . 21 .

Ry der , J . A . T he V e n t ra l Suc k er or uc k i g D isk s

S n
”
of th e

T dp l s
a o e of D iff e re n t G e n era of F gsro an d T ds A m N t
oa . . a .
, V ol . 22 .
1 22 TH E BIO LOGY OF TH E F RO G CH A P .

An excelle n t ex ample o f fla t fen ea or sq u a mous ’

m ay be obt ai n ed in th e ou te r most ski n which is c ast off

duri n g the molt Th e cells of this l aye r are bro ad and ex
.

c e edin gly thi n an d show a rou n ded ,

nu cleu s n e ar t he ce n ter Th e cel ls .

o f th e perito n eu m are m ostly o f the

s am e fl a tte n ed type I n the coin /n .

na r the c ells are elo n

g ated pe r p e n d icul arly to th e su r
fac e an d are u su ally p r ism atic in
ou t li n e owi n g to m u tu al pressure ; ,

such epith eliu m 1 8 co mmo n in t he:

F lc 3 , A p m
1 ,
_
f t h mucous l ayer of th e i ntes t i n e I n
o on o e .

e
p i d miser
m an y pl aces as in th e ou te r ski n
s, s t m c ll
o a e
, ,

there m ay b e all tr an s it io n al st ages

.

be t wee n colu m nar epi th eliu m an d squ amous epi th eliu m .

Layers su ch as thi s which are seve r al cells d eep are c alled

s t r a z zfi ea ep z f/
' '

’
z e/
‘
z n nz .

I n so m e p ar t s of th e body the r e occurs a peculi ar v ariety

c alled ez lz a tea ep z t/z elz n nz in which th e cells are fur n i sh ed
’ '
’
’ ’

wi th cili a at th eir outer e n ds U su ally suc h cells are colum .

n a r bu t th ey m ay be cuboid or eve n so m ewh a t fl atte n ed

, .

C ili ate d epith eliu m occurs in the mou th and thro at of th e

frog in cert ai n p ar ts of th e p eri t o n e al li ni ng o f th e body
,
.

c avity o n th e i nn er li ni n g of th e oviduc t s in t h e m ouths

, ,

o f th e c ili at ed fu n n els of th e kid n ey in th e ve n tricl es o f the ,

br ai n an d in e arly l arval li fe o n th e ou ter surface o f the

, , ,

body I f t h e ro of of th e m ou th o f a frog be scr aped wi t h

.

a k n ife an d t h e cells re moved an d ex a mi n ed u n d er a mic r o

scope a s him meri ng movem e n t m ay be see n o n o n e sid e of

,

e ach cell This i s d ue t o t h e rapi d move me nt of th e cili a or

.

fi n e h ai rlike processes o n th e surface Th e cili a of all t h e .

cells of a p articul ar are a be at most stro ngly in o n e di r ectio n ,

vr H I STOLO G Y OF TH E FROG 1 23

an d the effec t o f this commo n moveme n t is to c r e ate a

cur r e n t which c ar r ies sm all o bj e cts away from th at regio n .

Th e actio n o f cili a m ay e asily be de mo n s t r ated by spri nkli n g

so m e powdered c armi n e o n th e roof of a frog s m ou t h S oo n ’
.

o n e m ay observe th at the subst an c e is slowly c ar ried b ack

w ard dow n the esoph agus i n to the s t o m ach .

Th e eon n eetz ve tiss ues emb race a l arge n u mber o f tissu es

'

whose ge n er al fu nctio n i t i s to support an d hold togethe r

th e v arious o the r p arts o f th e body While in the oth er .

ki nds of tissue the i n te r cellul ar subst an ce is r el atively very

s m all in amou n t in th e co n n ective tissu es it i s usu ally very
,

abu n d an t . Ne arly all of th e co nn ectiv e tissu e is d e r ived

fro m the middle ger m l aye r o r m esode r m I t ar ises chi efly
, .

from sc attered cells or mese n chym e an d in th e e arly s t ages

, ,

o f its differe n ti a tio n th e amou n t of i nt ercellul ar subst ance is

ve r y sm all an d of a j elly like co n sis t e n cy The in tercellul ar
,
-

subst an c e beco m es modified in v arious w ays in th e differe n t

v arie ties of co n n ective tissue I n som e c ases i t re m ai n s
.

soft in others i t beco mes fibrous in bo n e i t become s h ard

, ,

e n ed through deposits of c arbo n ate and phosph ate of li me .

The p r i n ci p al ki n ds o f co nn ective tissu e fou n d in th e frog

ar e the foll o wi n g

W/u te fi é r ous co nn ectiv e tissu e i s th e v ariety which has

'

the widest distr ib u tio n A good ex ample of this m ay be

.

obt ai n ed fro m the m embran es whi ch co n n ec t the ski n with

th e body w all I f a portio n is spre a d out o n th e slide an d
.

ex a mi n ed wi t h the m icroscope it will be see n to be m ade up

,

of a cle ar homoge n eous por t io n or m a trix o f a gel ati nous

, ,

subst an ce in which are i mbedd ed n u m erous fibers th e

fibers are usu ally u nbran ched and h ave a ch ar acteristic w avy
a ppe aran ce Th ey are freque n tly u n ited in bu n dles which
.

ru n in all di rectio n s Wh e n tre ated with acetic acid they

.
,

swell up a nd dis appe ar an d whe n boile d becom e co n ve r t ed

, ,
1 24 TH E BIOLOGY OF TH E F RO G CH A P .

i n to gel ati n S c at t ered a mo ng the whi te fibe r s th ere are

.

ge n er ally a fe w y et/o w e/a s tz e fi oers ; these are str aight an d

'

n o t w avy ; th ey are n o t a ffec ted by acet ic acid an d d o n o t

yi eld gel at i n wh e n
boiled th ey fre
que n tly br an ch and ,

wh e n cut across t h e ,

e n ds do no t curl like
those o f th e white
fibers I mbedded .

in sp aces of the m a
trix here and the r e
are the co n n ect ive

t iss u e eo rp u se/es or
cells Th ese cells .

v ary co n sider ably in

their for m an d in
th e appe ar an c e of
th eir c y t o p l a s m ;
F I G 3 — F ib us c n c t i v t issu f m
. 2 . ro
u s u al l y t h e y ar e
o ne e e ro

f g
ro . c c t i t i ssu c p uscl s ;
6 , o n ne l s
ve e or e e, e a

ti
c fib s ; h t fib s
er w
( A ft Pw k1 ed br an ch ed
er an d th e
er ar er a n ,

P k “
.
,

ar )e
br an ches o f n eigh
bori ng c ells o ft e n u n ite o r an a stomose for m i n g an irregu ,

lar n e t work th e m esh es o f which are filled wi th the i nter

,

c ellul ar subst an ce These processes o f t h e cells ru n in

.

c anals wh ich allow a circul atio n of t h e fluid amo n g th e sp aces

or l acu nae in whi ch t h e cells li e Whi te fibrous t issu e v aries .

gre a tly in co n siste n cy and texture in differe n t p arts The .

lo o se t issu e bi n di n g t h e muscles togeth er is c alled areol ar

t issu e an d is composed of s heets an d str an ds i n tersecti n g
,

e ach other in all pl an e s I t form s a co a ti n g or f a sez a for

’

e ach m uscle and to w ard th e e n d s of t he mu scles i t is

,

frequ e n tly modifie d i n t o ten a on which is very de n se and ’

1 26 TH E BIOLO G Y OF TH E FRO G CH A P.

t ilages covered by a l ayer or pericho n driu m wh ich

are , ,

co n sists of an ou te r fib r ous m embran e below which are c o n ,

n ec t ive tissu e corpuscles which as th e c artil age grows si n k , , ,

i n to the m atrix an d beco me tran sform ed i n to o rdi nary car

t ilag e c ells H y ali n e c ar til age occurs at th e e n ds o f the
.

bo n es o f th e li mbs b e ,

twe e n th e vertebr ae an d
at th e e n ds o f their tr an s

verse processes at the ,

tip of the u rostyle in the ,

pubis o f the pelvic gir s

dle in th e hyoi d an d th e ,

s c artil ages of th e l ary n x a ,

an d at both e n ds o f the

ster nu m it forms th e
F lG 3 3 — C til g f m t h h d f t h
asis o f the c r an ium an d
a
e
ar ro e ea o e

f mu
e
. .

c lls ; c lls n p c ss f d i
r. 0, e
b e e
o

i ro e o

visi on
; m p ty c ll s p c ; m t ix the ce n tr al axis O f th e
6 . s, e e a e 772, a r .

( A ft P k
er d P k
ar ) er a n
lowe r j aw
ar er .

Calezfi ea ca r tilage which co n t ai n s a deposit o f li m e s alts

’
,

in the m atrix occu rs in th e supr asc apul a the pelvi s o f old

, ,

fr ogs an d at the e n ds o f so me o f th e l arge r bo n es o f th e

,

limbs viz th e he a ds o f the h umerus and th e fe mu r

. .

Th e structure of t o ne is si mil ar to th at o f c artil age in th at

it co nt ai n s cells i mbed ded in a solid m atrix I n bo n e t h e .

m atrix i s re n dered fi rm by th e d eposit of c arbo nate an d

.

phosph ate o f li m e B y i mmersio n in aci d th e lim e s alts m ay

.

b e removed an d a c artil agi n ous body h avi n g esse n ti ally th e

s ame hi stologic al struct ure as bo n e r em ai ns B o n e how .
,

ever is n o t m erely c alcified c artil age i t diff ers fro m i t both

,

hist o logic ally and ch emi c ally C a r til age is ofte n the p re cu r .

sor o f bo n e but in such c ases the fo r mer i s b r oke n dow n

,

an d bo n y t issue built u p in its pl ace

Two pri ncip al v arie ties of bo n e ar e u su ally disti nguished ,

v1 H I S T OLOGY OF TH E FROG 1 27

eo m bo e which i s v ery firm n d d e n se an d sp o ngy o r

p a etn ,
a ,

ca n cel/ ou s bo n e which is m a de u p of pl ates an d b ars formi ng

,

a structure which is comp ar atively loose an d l a cki n g in

stre ngth The l at ter is fou n d withi n th e ce n ter o f the verte

.

brae an d to a sm all exte n t w i thi n so m e of th e lo ng bo n es A .

goo d ex ample o f
c o m p ac t b o n e
m ay be obt ai n ed
by m aki n g a cross
sec t io n o f th e fe
mur The ce nt r al
.

p ar t of th e bo n e i s
hollo w and fille d
wi t h m a r r ow an d ,

t h e ou t er surface

is covered by a
l aye r of per ioste
u m w h ich is si mi
,

lar in structure to
th e perico n driu m F I G 34 A p t f c ss s c t i n f th f mu f ar o a ro e o o e e r o

th f g c liculi l cu 1m l m ll ;
. .

; l
su rro u n d in g t h e m m e ro . 6 n,;
c vi t y (A ft P k nd P k )
arro w a
ana o,

er
a

ar
ae ,
a e ae

, . er a ar e r.
c a r t il ag e T h e .

bo n y subst ance is arran ged in co n ce n t r ic l ayers o r la me/la , ,

which co nt ai n n u merous la eu n w in which lie the l one t ells , .

F r o m th e l acu nae fi n e br an chi n g tubes o r ea n a lieu li co n t ai n , ,

ing processes from t h e bo n e cells are give n off which exte n d ,

in all directio n s an d an astomose wi th the c an aliculi o f n eigh

bori ng sp aces .

B o n es i n cre ase in thi ck n ess by th e addi tio n o f succe ssive

l ayers to t h e outsi de Th e osteo bla s ts or cells formi ng the
.
,

i nn er l ayer of the p er io st eu m give rise co n ti n u ally to n ew

,

bo n e c ells w hich c ause the d epositio n of n ew l ayers of bo n y

subst ance s betwee n th e pe r iosteu m and the old bo n e New .
1 28 TH E BIOLOGY OF TH E FROG CH A P
.

l aye r s m ay also be add ed fro m wi t hi n by a l aye r of cells lin

ing th e i n n er surface of th e w alls of the m ar row c avity .

M u scle i s composed of elo n g at ed cells or muscl e fibe r s

u n ited by co nn ec tiv e tissu e Two v arieti es of m uscl e are .

com mo n ly disti n guish ed the s t ria tea or so c alled vo lu n ta ry

,
’
,
-

an d th e u n s tr ia ted or in volu n t a iy ,
.

I n th e l a t t er th e cell structure is
r el atively si mpl e ; the fibers are
co m mo nly spi n dl e sh aped with a -

si n gl e n ucleus n e ar th e ce n ter ,

whic h is u su ally elo ng ated in the

di rec t io n of th e fiber Th e e n ds o f .

th e fib ers are so me t i m es b ran ched ,

b u t they are more commo nly en

tire The le n gth o f th e u n stri ated
.

m uscle fibers v ari es gre atly ; they

m ay be very n arrow an d at t enu
ated as in th e w alls of t h e bl add er , ,

or short an d comp ar at ively thick ,

as in th e w alls of the s m aller blood

vessels Whil e t h e fibers usu ally .

sh ow n o cross stri atio n the cyto ,

pl asm shows delic ate lo n git udi nal

F ro 3 5 — U t i t d muscl
str an ds or fi b r il/
ns r a e e
m which are c o n , ,

fib s f m t h i t s ti
. .

er ro f sidered by most i n ves t ig a tors to

e n e ne o

th f g m d ( A ft
e ro '
be th e co n tractile elem e nts o f the
” u' l eu s ' er

Ho ) w es '

cell The c ell w all i s very thi n .

an d tr an sp are n t I n i ts actio n u n stri ated muscl e is slow ;

.

a co n sider able tim e el a pses before i t res o n d s to a stimulus

p ,

and i t is also slow to r el ax I t i s fou n d in t hose p arts of th e

.

body wh ere th ere is li ttle occ asio n for sudd e n movem e n t .

I t occu r s in th e muscul ar co at s of the ali me n t ary c an al in ,

the w alls o f t h e blood vessels and o f m an y du cts in th e ,

1 30 TH E B IOLO G Y OF TH E F RO G CH A P .

ing of alte r nate light an d d ark b an ds Th e lo ngi tudi nal .

stri atio n i s due to the exis t e n c e o f mi n u te s t r an ds t h e sa r ,

cos ty les o r fi br il/a which exte nd th e le ngth of th e c ell Th e

, .

fi b rillae w hich are supposed to represe n t th e co n tr actile ele

,

m e n ts o f th e fibe r are sep ar ated by a semi fluid subst an c e

,
-

the sa r copla s m There is an arr an ge m e n t o f th e fi b rillae

.

i n to bu n dles th e m u scle colu mn s which are s ep ar ated fro m

, ,

e ach oth e r by a thicke r l ayer o f s ar copl as m th an th at b e

t w ee n th e fi b rillae .

The appe ar an ce o f c r oss stri atio n i s b r ough t about by th e

divisio n o f the fi b rillae i n to seg men ts or sa r co mer es Th e
, .

s arco meres are sep ar ated from e a ch oth e r by a ve r y fin e

d ar k li n e k n o w n as K r a u se s mem or a n e which exte n d s n o t
’

o n ly across the i n dividu al fi b rillae bu t across th e s ar copl as m

,

b etwe e n th e fi b rillae of th e fibe r K r ause s me m b r an e li es

.
’

in the ce n te r o f a co m p ar atively cle ar an d lightly st ai ni n g

b an d fo r med by th e oppose d e n d s of th e t w o co n tiguous
segme n ts The middle portio n of e ach s arcom ere form s
.

th e so c alled d ark b an d Across th e ce n ter of this b an d

-
.

there exte n ds a seco n d ve ry delic ate me mbr an e k no w n as ,

the line of H ensen Wh e n the fiber i s rel axe d th is li n e m ay

.
,

be see n to li e in th e ce n ter o f a co mp ara t ively ligh t b and ,

which is usu ally n o t evide n t wh e n t h e muscle i s in a c o n

tr acted st ate .Th e d ark b an d s of th e muscle fiber are
composed of m ateri al which is an iso t r opic or doubly refr act ,

ing while the lighter are as o n eith er side o f K r au se s m e m

’
,

br an e are isotropic or si ngly refracti n g like the s ar copl asm

, ,
.

Whe n viewed with pol arized light th e differe n ces betwee n

thes e two subst an c es are cle arly b r ought ou t .

A t r an sverse sectio n o f a m uscle fiber prese n ts th e ap

p e a ra n c e of a n u mber o f polygo n a l a re a s c alled C o lz n /
z eim s
’

fields which represe n t the cu t e n ds o f th e muscle colum n s

, ,

t h e sp aces betwe e n t h e field s be i n g filled w i th s arcopl asm .

v1 H IST OLO G Y OF TH E F RO G 1 31

E ach of th e fi elds shows a do tted appe aran c e due to th e

.

cu t e nds o f t h e i n dividu al fi brillae .

The muscl e fibers of t he he art diffe r fro m both of t h e

above cl asses They are cross s tri ated bu t e ach fiber c o n
.
-

t ai n s bu t a si n gl e n ucleus E ach muscle cell i s fur n ish ed

.

wi t h br an ches which co nn ect with th e bran c hes of co n tigu

ous muscle cells so th at th e whole m ass forms a sort o f
,

n etwork .

Th e tissu e o f t he n e r vous syste m co n sists o f n er ve fibers ,

a n d n erve or g a ng lio n cells

,
E ach n erve is com posed of .

u su ally a l arge n u mber of n e r ve fibers h eld toge t he r by c o n ,

n ec t ive n erve tissu e a n d su r rou n ded by a co mmo n she a t h .

A t ypic al n erve fiber prese n ts t h e followi ng p arts a ce n tral

s t r an d or a xis cy lin der ; a sh e ath of fat t y subst an ce arou n d
,

this c alled t h e m edu lla /y slz ea tlz or zo /z ite su bs ta n ce of ,

S c/izva n n an d a d elic a t e exter n al m e mbr an e n eu r ilem m a , ,

or shea th of S chw a n n A t i nterv al s co n strictio n s occu r

.
,

c alled th e n odes of R a n vier w here the white subst an ce is ,

i nt erru pted al t hough the axi s cyli n der an d n eurilemm a are

,

co n t i nuous I mm edi ately be n e at h th e n eurile mm a occu r

.
,

the n ucl ei e ach surrou n ded by a sm all amou n t of proto

,

pl as m . E ach i n ter n od al segme n t or sp ace be twe e n t w o ,

n od es of Ra nvi er co n t ai n s sever al obliqu e m arki n gs a cross

,

th e medull ary she at h which are k n ow n as th e in cis u r es of

,

S elim idt .

The axis c vlin der o f a n erve is si mply th e elo ng ated pro

c ess o f a g an glio n cell an d u n der high m ag nific atio n is
,

fou n d to be m ad e up much like a m uscle c ell of very fi n e

, ,

fi b rillae with an i n terve n i n g subst an ce of more fluid co n

,

siste n cy The white o r m edull ary subst an ce co n t ai n s a

.

l arge amou n t of fat t y m ateri al c alled myeli n i f a fresh n e rve

i s pl aced in w ate r this s u bst an ce will s w ell u p an d collect
,

in drops givi ng th e n er ve a ve r y irre gul ar ou tli n e

, T he .
1 32 TH E B I OLO G Y OF TH E F RO G CH A P .

m edull ary she ath is supposed to ac t as a sort o f i n sul ato r ,

like th e co ati ngs th at are wou n d arou n d an electri c wire .

Th e n erv e fibe r u n like th at o f muscle is a com posite

, ,

structure bei n g formed of c ellul ar eleme n ts of diverse

,

origi n The sh e aths of th e n erve repres e n t a seri es of cells

.

which h ave beco m e applied t o bu t h av e an e n tirely differe n t

-

or igi n fro m the axis cyli nd er Th e l at t er is alw ays an o u t .

growth o f a n erve o r g an gl io n cell an d is always o f e c t o der

mic origi n I n the d evelop me n t o f a n erve the axis cyli n d e r
.

is alw ays th e first p art to m ake i ts app e aran ce as i t grows

out pu shi n g i ts w ay th r ough the othe r tissues i t beco mes
, ,

FI G .
37 c lls d fib s f
. Nerve f sh
e fib
an B er o re n e rve er . ,

fib
n er ve i h h my li s ll h ugh t h bs p i
er w t t e f e n w o en t ro e a or t on o w at e r .

C c ss s c i
,
ro f fib s D g gli c lls
e t on o xis cyli d
n e rve er an on e . ax , a n er

xis cyli d p c ss f g gli c ll ; i cisu f S h mid t

.
,

p
a sa , a n er ro e o an on e n re o c ,

m m dull y sh th ;
. s, e ucl us ; l u il mm ; N R d f R
ar ea n, n e n ne r e a . no e o an

i ; p p p t p l smic p c ss f g gli c ll
, ,

v er , ro o a ro e o an on e .

'

surrou nded with n ucle ated cells w hich fl at te n o ut an d form

th e n eurile m m a the white subst an ce ap p e ars at a co mp ar a
1 34 TH E B IO L O G Y OF T H E F RO G CH A P .

C H A PT E R V I I

TH E DI G E ST IV E SYS T E M A ND I T S FUN CT IONS

O NE of th e ch ar acteri stics o f all forms o f li fe is th e n e ed

of food The m atter which composes th e bo dies of livi ng
.

o rg anis ms i s bei ng co n ti n u ally broke n do w n an d eli mi nat ed

as w aste pro du cts Ne w m at ter is co n sequ e n tly required
.

to m ake good th e loss i f th e vi t al process be ke pt g o i n g .

I n th e frog a p art o f t h e m at eri a l i s t ake n fro m th e oxyge n

o f the air an d from th e w at er absorbed t hrough th e ski n ;
b u t n either of t h ese sources supplies th e c arbo n n i t roge n , ,

a n d other ele me n ts which form esse n t i al p arts o f all livi n g

subst an ce L i fe phe n ome na are a ssoci ated especi ally wi th

.

cert ai n compou n d s c alle d pro t eids Th ese are co mplex .

subst an ce s co n t ai n i n g c arbo n oxyge n hydroge n an d n i t ro

, , ,

e n a n d in m an y c ases lso sulphur ph osphorus c lciu m

g , ,
a ,
a , , ,

potassiu m sodiu m iro n an d occ asio n ally o t her ele me n t s

, , ,
.

L ivi ng subst an c e or pro t opl asm i s o f pro teid n ature but

, , ,

it is prob able th at it is a group o f co mpou n ds r at h er

th an a p ar t icul ar compou n d w hich we m igh t express by a
defi n i t e che mic al formul a This livi n g m atter i s th e sub
.

j t of ch emic al ch anges which are spoke n of u n der the

e c

ge n er al te r m m eta bolis m The sy n t hetic or bui ldi ng u p

.
-

processes by wh ich this subs t an ce is form ed from si mpler

compou n ds are c alled a n a bolis m ; th e opposite or t e ari ng ,

dow n processes by which it is resolved i n t o si mpler sub

st an ces are k n ow n as ka ta bolism I f an org an ism gro w s i t is
.
,

evide nt th at th e anabolic side of th e process must p redo m i

VII TH E D I G E ST IV E SYS T E M A ND I T S FUN CT IO NS 1 35

na te over th e k at aboli c I f kat abolis m predomi n ates or in

.
, ,

o ther w or ds i f w aste exceeds rep air th e org an ism m ust

, ,

dimi n ish in size .

No w th e fun ctio n of food i s n o t merely to com pe n s ate for

th e m ateri al which i s broke n d ow n an d elimi n at ed but to ,

afford th e e n ergy n ecess a ry to c arry o n th e v arious ac t ivi

ties o f t h e org an ism F oo d i s t o t h e body w h at fu el is to

.

a ste a m e ngi n e Th e body i s co nti nu ally expe n di n g e n ergy

.

in t h e form of h e at Th e a mou n t of e n ergy lost in this w ay

.

depe n ds upo n circu mst ances and it m ay b e co mp aratively

,

s m all whe n the te m per a ture of th e an i m al i s o nly slightly

a bove freezi n g B u t so lo ng as li fe l asts the re i s so m e he at
.

p roduc ed an d thi s h e at resul t s fro m t h e bre aki n g dow n of

,

som e o f th e co n sti t ue n t s o f the body E very move me n t .

which t h e frog performs i nvolves th e expe n di ture o f e n ergy ,

w hich mus t co me ul ti m ately fro m its food su pply An .

organ ism h as o ft e n bee n comp are d to a vor t ex w hich m ai n

t ai n s i ts form while th e m ateri al o f which i t i s composed is
,

subj ect to co n ti nu al ch an ge Th e m at ter composi n g th e

.

t issues of an a n i m al is n o t th e s a m e duri n g successive ye ars ,

n o r quite th e s am e duri n g succ essive d ays I t i s bei ng c o n

.

t in u ally dr aw n t h r ough th e v o rtex wh ere i t gives u p a p art

,

o f it s e n ergy for th e m ai n t e n an ce of the vi t al processes .

Th e subs t an ces elimi nated by an an i m al possess th erefore , ,

l ess e n ergy th an the food m ateri al t ake n in Th e a mou n t .

of e n ergy obt ai nable from a gra m o f an y p ar ticul ar co m

pou n d such as c an e sug ar w he n i t u n d ergoes d ecomposi
, ,

t io n m ay be me asured wi th co n sider a ble accur acy

,
If w e .

m e asure t h e e n ergy resulti n g from the splitti n g up of a c er

t ai n a mou n t of food subs t an ce an d comp are i t wi t h th e

e n ergy obt ai n able from an e q u al a mou n t o f th e s am e ki n d

o f food m at eri al a ft er it h as bee n elimi nated from the
org an ism w e should fi n d the l atter to be much less in
,
1 36 TH E BIOLOGY OF TH E FROG CH A P .

a mou n t I f n o w we could m e asure th e e n ergy expe n ded by

.

the org an ism by r adi at i n g he at a n d performi n g work duri ng

the time t his m ateri al i s co n su m e d we should prob ably fi n d,

it t o be equ al to t h e differe n ce betwee n the pote n t i al e n ergy

o f the food an d t h at of t h e eli mi n ated products A ll o f our .

experi e n ce goes to prove th a t the gre at law o f co ns erv atio n

of e n ergy a ppli es as strictly to org an ism s as to t he phe
n o m e n a of th e i n org an ic world L ivi ng bei n gs are n o t
.

sources o f e n ergy in t he mselves bu t are d epe n d e n t u po n

,

their e n viro n me n t for e n ergy as much as th ey are for th e

m ateri al co m p osi n g th eir bodies .

I n order t h at food m ateri al m ay be a ssi mil a ted or built

u p i n to th e tissu es o f t h e bodi es it m u s t be re n d ered solu
,

ble so t h at it c an p ass through the li n i ng o f the alim en

,

t ary c an al i n to th e blood a n d lymph an d fro m these fluids ,

through the w alls o f the cells in t h e differe n t p arts o f th e

body This process o f co n verti ng food i n t o a soluble st at e
.

re ady for absorptio n is c alle d d igestio n There are cert ai n .

m ec h an ic al proc esses i n volved in digestio n such as (in h igher ,

a n i m als ch ewi n g t h e food m ovi n g it abou t by t h e c o n t rac

) ,

t io n s of t he w alls o f th e sto m ach an d p assi n g it alo n g th e

,

i nt es t i n e by th e peri st altic co n t r a ctio n o f th e w alls Th e .

frog however like mos t lowe r ver t ebr ates does n o t chew t h e
, , ,

food t ake n i n to th e m outh but sw allo w s it whol e dow n t h e

,

very d iste n sible esoph agus i n to t he s t om ach wh ere i t i s ac ted ,

u p o n by th e g astric j u ice The pri n cip al p art of t h e process

.

of digestio n co n sis t s in the c hemic al ch an ges produced in food

by the ac t io n of th e v ariou s diges t ive fluids Th es e ch an ges .

are m ai n ly o f th e n ature of ferm e n t atio n s c aused by sub

st an ces c alled enzy mes or fer m en ts Wh at the ch emi c al

,
.

n ature o f e n zymes is still rem a i ns very much in th e d a rk ,

si n ce t h ey c ann o t be comple t ely freed fro m th ei r associ a

tio n wi t h other subst an ce s b u t it is prob able th at t hey are
,
1 38 TH E BIOLOGY OF TH E FRO G CH A P .

The E soph agu s — Th e esoph agus i s very

an d S t omach .

short an d rem ark ably dis te nsibl e as is prove n by th e rel a ,

t ively l arge an i m als t h e frog is c ap able o f sw allowi n g The .

i n n er surface i s throw n i n t o
lo ngitu di n al folds which ex
t e n d also i n to th e s t o m ach .

Th ere i s no sh arp li n e o f
d em arc atio n sep ar ati n g the
esoph agu s fro m the p/z a r
y u x o n th e o n e h an d and

fro m th e st om a clz o n th e
o t her Th e a n terior e nd .

o f th e sto m ach i s co n sider

ably w id er t h an the eso p h a

gus and th e o rg an t apers ,

gr adu ally to th e pos t erior

o r pyloric e n d wh ere it is ,

sep ar at ed by a c o n st ric
tio n t h e py lo r u s fro m t h e , ,

s m all i n tes t i n e Th e s to m .

a cli lies m ai n ly in the le ft

h alf of t h e body an d i s ,

curved so t h at th e co nvex
side is tow ard th e le ft I t .

i s suspe n d ed d o rs ally by a
fold o f peri t o n eum t h e m es
og as ter an d fro m th e ve n ,

F I G 3 8 A lim t y c l f R
. . e n ar
tr al side a r is es a
an a o
seco nd
a n a es

l t A
c u en a p i g f t h c t um i t sheet o f peri t o n eu m ( t h e
.
,
o en n o e re n o

the cl c Cl D du d um ; D as t o lz a to du oden al liga

o a a, ; u o en
p
, ,

i l um ; b u d ybt g r e
- -

e o h l t
n ar e w e en t e a

ter a n d h l g i t s t i R H 8 m en t ) which exte n ds to th e

t e ar e n e ne, ; , ,

u i y bl dd ; M s t m ch ; M duod e n u m and live r The

r n ar a er , o a z ,

s p l ; O s p h gus ; R p yl us
e en e, e o a y, or .
.

( A ft VKi d h m ) w all o f the stom ach is mu ch

'

er c c rs ei .
v1 1 TH E DIG E S T IV E SYS T E M A ND I T s FUN CT IONS 1 39

thicke r th an th at o f the esoph agus or th e i n testi n e The .

i nn er surfac e is t hrow n i n to sever al lo n gitudi nal folds which ,

beco me less pro mi n e n t posteriorly and n e ar t h e pylo r ic end

,

e n tirely dis appe ar .

I n a cross sectio n of th e stom ach o n e m ay observe a very

t hi n ou t er l ayer composed of much fl at t e n ed cells ; thi s i s

the ser o u s coa t or ser osa an d i t i s form ed by t h e peri to n eum

‘

, .

Withi n the seros a i s a thicker l ayer th e s u bser osa co n si s t i n g

, ,

m ai n ly o f co n n ective tissu e Thi s l ayer h as be e n frequ e n t ly

.

described as a l ayer of lo n gi tudi n al m uscles an d i t has t h e ,

appe ar an c e o f su ch ; bu t i f tre ated w i th the proper s t ai n s ,

it c an re adily be sh ow n to be m ai n ly co nn ective tissu e .

S om e writers (V alat ou r P S chu l t ze ) o n the o t her h and

,
.
, ,

h ave bee n disposed to de ny th e exi ste nc e of lo n gitudi n al

muscles in the frog s sto m ach I n sectio n s across t h e c ardi ac
’
.

e n d of th e stom ach ho w ever o n e m ay de tect a few mu scle

, ,

fibers a mo n g the co n n ective tissue an d in th e pylori c en d

, ,

a ccordi n g t o G au pp t here are a few lo ngi tudi nal fibe r s

,

which are co n t i n uou s wi th t hose o f the i n testi n e .

Withi n th e subseros a i s a thick l ayer o f cir cu la r m u scles

which beco mes thicker tow ard th e pylorus I n ter n al t o th e .

circul a r muscles i s a l ayer of co nn ec tive tissue t h e s u b ,

m u cos a in which there a re n u me rous bloo d vessels

,
The .

tissu e of t h e submucos a exte n d s i n to the folds o f the i n n er

co at B etwee n the mucos a and submucos a there is a thi n
.

muscul ar l ayer the m u scu la r is m u cosce com posed of an

, ,

i nn er l ayer of ci r cul ar fibers an d an oute r str atu m of lo n gi

t u dinal o n es .

The m u cosa o f the stom ach is a thick l ayer composed o f

gl an ds embedded in a suppor t i n g m atrix of co n n ective tissu e .

Th ese gl an ds represe n t i nv agi na tio n o f th e epi th elium li n i n g

t h e i n n er surface o f th e sto m ach They are elo n gat e d
.

t ubul ar st r ucture s set very closely to ether an d fre u e n t ly

g q ,
1 40 TH E BIOLO GY OF TH E FROG CH A P .

more or less br an ched Th e gl an ds differ in structu re at.

th e t W O en ds of t h e sto m ach I n th e c ardi a c regi o n the

’
.

gl a nds are very lo ng t h e mou t h o f th e gl an d i s qui t e d eep

, ,

a n d li n ed w i th elo ng ated cells w hose cle ar i n n er e nds

a re filled with a subst an ce which prob ably forms m ucus .

Ne ar the ou t er en d of the gl an d th e cells are m ore elo n g ated ,

li ke t ho se of t h e su rface epi th eliu m ; b ehi n d th e cle a r sub

st ance th e cy t o pl as m o f th e cells is gran ul ar th e n ucleus is ,

F IG .
39 . G l ds
an of th e s m ch
to a . A f m c di c d ;
ro ar a en B f m p yl ic
ro or

m uh; ck ; b b dy f gl d
, ,

end ; m ,
o t 72, n e , o o an .

elo n ga ted an d th e outer e n ds are dr aw n ou t i n to a lo ng

,

n arro w p rocess P assi n g do w n th e m outh o f th e gl an d t h e

.

cells bec o me shor t er th e n uclei m ore rou n ded a n d the t ail

, ,

like p rocesses fi n ally almost dis a pp e ar I n t h e n eck regio n s .

of the gl an d t here are usu ally a few r ath er l arge cells c o n

t ai n i n g a l arge cle ar v acu ole w hic h p u sh es t h e n ucleu s an d
'

mos t o f the cyt o pl asm to o n e side I t is u su ally in t h e .

regio n of th ese cl e ar cells th a t t h e gl an ds br an ch The cells .

co mposi ng the body of th e gl an d lie j u st below t h e cle ar

1 42 TH E BIOLOGY OF TH E FROG CH A P .

th e gl an ds o f the c ardi ac e n d o f the sto m ac h Th e m ou ths .

of th e gl an ds are li n ed w ith a short cyli n dric al epithel iu m

with occ asio n al cili a ted cells .

G ast ric D igest ion. I n the s tom ach th e food is subj ected
to th e actio n of th e gastric j uice which i s secre t ed by th e
,

gl an d s o f the muc o s a G astric j uice is acid in re actio n fro m

.

the prese n ce o f a sm all qu an tity o f fre e hydrochloric acid ,

a n d i t co n t ai n s also a ferme n t e s in wh ich a cts u po n the

pp , ,

proteids co n verti n g th e m i n to soluble p ep t o nes Nei t her th e

,
.

fat s n o r t h e c arbohydr ates u n d ergo di gestio n in th e s t o m ach .

B y diges t i n g out t he proteid porti o n o f foods in which fats

a n d c a rbo hydr ates a re co n t a i n ed th e g astric j u ice h elps to

re nde r th ese subst an ces more r e ad ily digestible by oth er

fluids .

The actio n o f th e g astric j uice of th e frog m ay be re adily

demo n s t r ated by sip ho n i n g o ff so m e of t his flui d fro m th e
s t o m ach by m e an s o f a be n t gl ass tube an d pl aci n g in i t a
s m all bit o f th e whi te of a h ard boiled egg The pi ece of
-
.

egg aft er a ti m e will be se e n to be corroded an d fi n ally i t ,

will become e n tirely dissolved .

Th e secretio n of the esop h agus h as a stro n g dige stive

powe r but its re a ctio n i s alk ali n e i n ste ad of aci d an d i t i s
, ,

c ap able of acti ng o n ly a fter i t h as bee n re n d ere d a cid through

mixture with the fluid o f the sto m ach ( Nu ssb au m ) .

Wh e n gastric d iges t io n i s completed th e fo od p asses ,

t hrough th e pyloru s i n to th e sm all i n testi n e .

Ch an ges in t h e G lands dur in g D igest ion — Th e ch an ges .

u n dergo n e by th e gl an ds of th e esop h agu s an d stom ach h ave

bee n stu died by P artsch S wi ecicki Nu ssb au m G riit z n er
, ,
'

, ,

L an gley an d S ew all I n frogs which h ave bee n kept f o r

, .

several d ays wi thou t food L an gley fou n d the cells o f th e

body of t h e gl an d to be e nl arged so as to pr ac t ic ally o blit er
a t e t he c e n t r al c an al
. The co n t ents of th e cell are u niformly
vu TH E DIG E ST IV E SYST E M A ND I T S FU NCT IO N S 1 43

gr an ul ar an d th e cell ou t li n es are very i ndis t i nct I n one .

t o t w o hours a ft er feedi n g th e lu mi n a begi n t o b e obvious ,

an d th e gr an ules to dis appe ar from the i nn er border of the

c ells. U p to t h e fi fth hour th ese ch an ges becom e

more an d more m arked an d at th e s ame ti me the cells an d
,

the re m ai ni ng gr an
ules t hey co n t ai n
becom e dis t i n c t ly
sm aller an d th e cel l
,

s u b s t an c e s t ai n s
m ore d eeply .

At t h e p e r i o d o f
m aximu m c h a n g e
t h e n ucleus is mu ch

l ar g e r c o m p ar e d
w i t h th e cell sub
st anc e th an it is dur
ing res t ; i t is s t ill
surrou n ded by fi ne A C

gr an ul ar PTO t O F I G 4 —
'

. Sh i g ch g s i th g s t ic gl nds
o . ow n an e n e a r a

p l a s m a n d it I S f th
o f g A gl
e d f
ro m . hu ng y f ,
g an ro a r ro

hich h d t b f d f fiv d ys T h
w a no een e or e a e
some t imes p l a C e d
.

c ll u t li s i dis ti c t d t h g ul s
e o ne are n n an e ra n e are

n e ar th e ou t er b o r sc tt d t h ugh u t t h c lls B gl d t h
a e re ro o e e an ree

h u s f m l ; th g ul s h
.
,

o t r a er a di p
ea e ra n e a ve sa
der O f the c ells Th e'

p d l g th i
e a re b d f t h c lls ;
a on e n n er or er o e e

retur n to th e n orm al lum f t h gl d isibl C gl d t n ty

en o e an v e .
, an w e

fi v h u s ft e h o y rm l ; th
a c lls
er a ea v ea e e ar e
appe ar an c e begi n s sh u k n d n t full f g n ul s ( Aft
r n en a o so o ra e er

gl y )
.

a bou t t h e fi fth hou r L , an e .

so t h at duri ng the
gre at er p ar t of the digestive pe ri o d the fo rm a tive processes
go o n whilst th e secre t ory are s t ill ac t ive I n twe n ty four .
-

hou rs the gl an ds h ave n e arly or al t oge t her re t ur n ed t o t he

hu n gry co nditio n .Th e ti m e an d ex t e n t of t h e ch an ges
p r oduced in th e gl an ds we r e fou nd by L an gl ey to v ary
1 44 TH E BIOLOGY OF TH E FRO G CH AP .

e n ormou sly wi th th e a mou n t of food give n an d the st ate

o f the frog “.I f a frog i s fed w i t h sever al worms so th a t
t he sto m ach is mu ch diste n d ed wi t h digestible fo od the ,

ch an ges are gre at er an d persist for a mu ch lo nger tim e .

I n twe n t y fou r hours th e gl an ds i n ste a d o f h avi n g retur n ed

-

t o th e hu n gry s t ate ar e s t ill sm all a n d co n sist o f some w h at

,

sm all c ells wi t h a more o r l ess dis t i n ct i nn er n o n gr an ul ar -

bord er ; th e lumi n a a re freq u e n t ly l arge Th e i n cre ase in

.

t h e siz e o f th e lu me n i s a ccom p an ied an d prob ably c aused

by th e decre ase in t h e size o f th e c ells “ .I n frogs to which

an excess o f food h as be e n give n th e n o n gr an ul ar i nn er
,
-

zo n e i s u su ally most obviou s abou t the eightee n th or

t w e n tieth hour a fter feedi n g The c ells th ere h ave i n cre ased
.

a n d a re still i n cre asi n g in size ; th e gre ater cl e ar n ess wi t h

which th e n o n gr an u l ar zo n e c an b e s ee n i s t he n p r ob ably
-

due to th e n et i n cre ase in th e cell gr an ules t aki n g pl ac e

m ore slo w ly th an th e i n cre ase in th e cell protopl as m .

L an gl ey fou n d t h at the effect of fas t i n g in wi n ter is n o t

very gre a t ; t h e cells of t h e gl an d be co m e so me w h at sm aller ,

b u t t h ey are fa irly well filled wi t h gr a n ules I f however the

.
, ,

wi n ter frogs are ke pt w arm o r i f frogs at o t h er t i mes of ye ar

,

ar e kep t fo r a lo n g t im e in a fas t i n g co n di t io n t h e cells

,

shri n k in size an d becom e cle ar alo ng th e i n n er bord er as

they do aft e r s ecretio n .

The mucige n co n te n t o f t he cells li ni n g th e m ou th o f th e

g astric gl an ds is l arge in a mou n t before an d for som e ti me
a fter a m e al but duri n g t h e heigh t o f t h e diges t ive process i t
,

beco mes m uch dimi n ish ed Th e ch an ges in t h e pyloric gl an d s

.

are much like thos e in t h e mou t h a n d n eck o f th e c ardi ac

gl an d s
.
“ The m axi mu m a mou n t o f mucige n i s co n t ai n ed
by t h e pyloric an d simil ar gl an d cells a ft e r a m oder at ely pro
l o nged fas t The m i n i mu m a mou n t o f mucige n i s co n t ai n ed
.

by these cells twelve to eightee n hours aft er a h e avy m e al ;

1 46 TH E BIOLO G Y OF TH E FRO G CH AP .

portio n of t h e cell goes o n so th at a cle a r z o n e i s formed .

The cle ar zo n e ste adily i n cre ases u n til t h e sixth to t w elfth

hour or eve n l ater th e ti m e v a ryi n g with th e st a te of th e
, ,

a ni m al an d th e a m ou n t o f food give n Th e gl an d s the n

.

begi n to beco me more gran ul ar th e ti m e o f co mple te ,

recovery v aries e n ormously : in so m e c ases the gl ands are

throughou t gr an ul ar in twe n ty four h ours fro m the tim e o f
-

fe edi ng the an i m al in o thers they do n o t becom e so for

,

several d ays .
”

I f t h e frog i s fed with pieces o f spo n ge i n ste ad of food a ,

secretio n is set u p both in the s t o m ach an d the esoph agus ,

th e ch an ge b ei n g as a rule t he gre a ter t h e l arger the spo n ge

, .

S i mil ar ch an ges t ak e pl ace in t h e cells to those produced by

digestible food bu t they occu r m uc h more slowly begi n n i n g
, ,

ge n er ally o n ly three or four hou rs a ft er th e spo n ge i s pl ac ed

in th e sto m ach t h e gr a n ul es begi n to i n cre as e a g ai n in th e

esoph agu s o n ly a fter so m e d ays .

Nu ssb au m h as fou n d th a t a dire ct s ti mul atio n o f a p art ic

ul ar p ar t o f t h e mu cous me mbran e o f the e soph agus c auses
a dis a ppe ar an ce o f th e gran ul es fro m th at regio n Th e c o n
.

c lusio n s of Nussb au m th a t in n o rm al hu n gry frogs th e c ells o f

t h e esoph age al gl an ds h ave an out er cle a r z o n e an d th a t ,

a ft er feedi n g th ere is a n i n cre ase i n ste ad of a d ecre a se o f

gr an ules were prob ably d r aw n fro m u nh e althy speci me n s

, .

S luggish an d u n he al t hy frogs o fte n S how gl an dul ar c ells wi th

a n ou t er cl e ar zo n e but lively an d vigorous speci m e n s h ave
,

t h e cells filled w ith gr an ules H u n gry frogs wi t h foreig n

.

bodies in t h e s t o m a ch su ch as bits of le a f or o t her obj ects

,

s w allowed w ith th e food oft e n show a decre ase in th e granu

,

lar co n te n t of th e g lan d cells owi n g to th e irri t at io n t hus set

,

up Accordi n g t o G rut z n er there is a prel imi n a ry i n cre ase in

.

th e gr an ules in t h e es o ph age al gl an d s for a short tim e a ft er

fee di n g an d the n a m arked decre ase but L angley w as able
, ,
VI I TH E D I G E S T IV E S Y S T E M A ND I TS F UN CTIO N S 1 47

to obt ai n no d ecisive evide n c e o f such p relim i n ar y i ncre ase

in gr an ul atio n al t hou gh h e w as n o t disposed to de n y th at it
migh t t ake pl ace at le ast to a slight exte n t .

O f wh at sig n ific an c e are th ese ch an ges in th e gr an ul ar

co n te n ts o f the gl an d cells ? I t i s evide n t th at they h av e
so m e thi n g to do with th e form atio n of digestive fluids
o f the esoph agus an d sto m ach an d i t is prob abl e th at th e
,

gr an ules are composed o f a subst an c e which i s tr an sform ed

i n to p epsi n Th a t they are n o t co mposed of p epsi n itself
.
,

bu t o f som e subs t an ce which has bee n c alled p ep sin ogen is ,

i n dic ated by the followi n g experi m e n t s I f the esop h agus .

or sto m ac h of a frog be pl aced in glyceri n as r ap idly as

possible after re mov al fro m t h e body t h e glyceri n extr act ,

has o n ly a we ak p ep t ic po w er I f t h e esoph agus o r stom ach

.

o f a frog be kep t m oist for twe n ty hours before i t is pl ace d

in glyceri n the glyceri n extr ac t h as a very muc h gre ater
,

pep t ic power I f t h e esoph agus an d sto m ach which h as

.

bee n ex t r acted wi th say 5 cu cm of glyceri n for a wee k

, ,
. .

be w ash ed fre e o f glyceri n an d tre a ted with 5 cu cm o f . .

dilute hydrochloric acid th e n an e n ormously gre ater amou n t

,

of p epsi n is fou n d in th e acid th an is fou n d in the glyceri n

extr ac t .
”

The a mou n t o f pepsi n co n te n t i s gre ates t in those gl an ds

in whi c h t h ere is t h e gre a tes t n u mber of gr an ul ar cells .

Th e pepsi n co n te n t o f the esoph agus w as fou n d by S wiecicki ,

L an gley an d S ewall t o be gre ater th an t h at o f an equ al are a

,

o f th e stom ach I n th e pyloric regio n wh ere t h e gran ul ar cells

.
,

are few in n u mb er th e pepsi n co n te n t o f th e gl an ds is much

,

less th an in the c ardi ac en d L an gl ey fou n d t h at if pi eces of

.

equ al size were c u t ou t o f th e esoph ag u s c ardi a c en d mi d , ,

dle an d pyl o ric en d of the s t om ach an d th e pepsi n co n te n t

, ,

of e ach estim a t ed t h e p o w er o f co n verti n g proteid w as m uch

,

the g r e atest from the piec e fro m th e esop h agus and be c am e ,

1 48 TH E BI O L OG Y OF TH E F RO G CH A P .

less resp ectively in th e piec es fro m th e o ther regio n s n a m ed .

P artsch Nussb au m S wiecicki L an g ley an d S ew all h ave all

, , , ,

i n v estig ated th e rel ative digestive power o f the gl an ds well

filled with gr an ules an d gl an ds fro m which th e gr an ules h ave
m ai n ly dis appe ared an d all agree th a t th e p epsi n co n te n t o f
,

the former is much th e gre ater .

Rapidit y of D igest ion — Th e digestive processes o f th e

frog comp are d wi t h t h os e of t h e higher vertebrates proce ed
slowly du e prob ably to th e fact th at the frog i s a cold blood ed
,
-

a n i m al . Th e le n g t h of tim e t ake n to digest a m e al v aries

wi t h th e am ou n t o f food L an gley fou n d th at a s m all e arth
.

wor m w as digeste d in so m e w h at less th an twe n t y four h ours -

bu t i f sever al worm s were give n t hey do n o t dis appe a r fro m

,

th e sto m ach u n til a lo n ger period I t i s very prob able th a t .

th e te mp er ature o f t h e body is an i m p or t a n t fa c t or in d eter

m i n i ng th e r a te o f diges t io n bu t I a m acqu ai n ted wi t h n o
,

observ atio n s t o th a t e ffect .

S t ru ct u r e of t h e I nt est ine The sm all i n tes t i n e begi n s

.

j ust b eh i n d t h e pyloric co n s t ric t io n an d ru n s fo r w ard as t h e

,

du o den u m for some dis t an ce w h e n i t tur n s abrup t ly b ackw ard

,

as the ileu m which a fter coili n g abou t in a n irregul ar m a n n er

, ,

wide n s o u t abruptly i n t o t h e l arge i n tes t i n e n e a r th e pos t erior

e n d o f t h e body Th e di ameter o f the sm all i n tes t i n e which
.
,

is n e arly u ni form t hroug h ou t it s course is much l ess t h an t h at ,

of t h e s t o m ach an d it s w alls are much thi n n er The i n testi n e

,
.

i s fas t e n e d by a mese n t ery t o th e mid dors al portio n of t h e -

b o dy c avi t y an d it s du od e n al portio n is co n n ec t ed t o t h e
,

liver an d s t o m ach by the previously m e n tio n ed re m ai n s o f a

ve n t r al mese n t ery t he g as t ro h ep at o duode n al liga me nt
,
- -
.

A cross sectio n of th e sm all i n t es t i n e sh o w s th e follo w i n g

l ayers At t h e ou tside is a very thi n co at of p er ito n eu m
si mil ar to t h at c o at i n g the s t o m a ch Wi t hi n t h is is a well.

m arked l ayer o f lo ng itu dina l mu scle fiber s ; the n co mes a

150 TH E B I O LO GY O F TH E F RO G CH A P .

L eucoc ytes o ft e n fou n d bet w ee n th e epi th eli al cells

are ,

a n d also w an deri n g cells of l arger size wi th bodi es of v ari

ous ki n ds in their pro topl as m ( H eide n h ai n D e B ruy n e ) ,

.

FI G .
41 .c ss s c ti f t h sm ll i t st i f th f g bl bl d
P a rt o fa ro e on o e a n e ne o e ro . oo

ss ls ; g g bl t c lls ; p di y p i h li l c ll ; submuc s ;
,

ve e c , o e e e or n ar e t e a e e s,. o a

ci cul muscl s ; m l l gi udi l muscl s ; p p i t um

,

m . c, r ar e .
, on t na e e, er one .

( A ft H s) er ow e .

W an deri n g cells co n t ai n i n g pigm e nt h ave bee n fou n d to

occu r in t h e lower e nd of th e sm all i n testi n e ( O ppel ) .

The m ucos a o f t h e sm all i n tes t i n e i s throw n i n to n umer

ou s folds b u t th ere ar e n o tru e villi n o r defi n ite gl an ds n o r
,

crypts such as occur in th e higher ver t ebrates Just behi n d .

the pylorus t h e fold s t ake th e form of an irregul ar n etwork ,

b u t a short d is t a n ce far t her b ack th ey become a rr an ged I n

two seri es of t r a n svers e se milu n ar pli c at io n s the free e dges
of which are produ ced b ackw ard formi ng a d ouble serie s ,

o f pockets whic h te n d to c heck t h e flow of food in t h e

direc t io n o f the stom a ch Th e pocke t s are co n n ec t ed by .

sm aller folds wh ic h r u n m ai n ly in a lo n gi t udi n al direc t io n .

F ar ther b ack a little beyo n d th e m iddle o f the i n test in e

, ,
VII TH E DIG E S T IV E SYS T E M A ND I T S FUN CT IONS 15 1

th e fol ds lose t heir regul ar arr an ge m e n t an d in th e posterio r ,

third they assum e a lo ngi t udi n al direc t i on .

The la rge in testine is composed o f the s am e l ayers as th e

sm all Th e i nn e r surfac e is thro w n i n to fol ds w hich at th e
.
,

proxim al en d form an irregul ar n etwork b u t in t h e rec t u m ,

t h ey b ecom e lo n gitudi n al Th e epi theliu m of t h e mucos a

.

co n sists of cyli n dric al cells am o n g w hich n u merous goblet

,

cells are to be fou n d .

Th e P an creas — The p an cre as i s an elo n g ated gl an d of

.

i rregul ar sh ap e situ ated betwee n th e stom ach an d t h e duo

denu m a n d exte ndi n g fro m th e liver to w i t hi n a short dis
,

t an c e o f t h e pylorus I t is tr ave rsed by th e com mo n bile

.

duct i n to whi ch it s ducts e n ter O f these t h ere is a pri n ci

.

p a l duct ,
a n d sever al sm aller ducts fro m th e p ortio n of th e
gl an d n e ar th e liver .

Th e p an cre a s is a muc h b r an ch ed tubul ar gl an d th e ter

-

mi n al br an ch es o f t h e gl an ds bei n g o fte n curved an d t w isted

in an irregul ar m an n er The t ubules are co ated exter nally
.

w i th a b a sem e n t m embr a n e an d held toge ther by a delic ate

,

co nn ective tissue in wh ich lie th e bl ood vessels an d n erves .

Th e secre tory cells of th e tubul es co n t ai n n u merou s

zymoge n gr an ules w hich wh e n the frog is in a hu n gry st ate
, , ,

are fou n d in gre at abu n d an ce especi ally at th e i nn er or free

,

en d of th e cell . These dis appe ar a ft er th e an i m al is fed ,

like t h e gr an ules in th e gl an ds of th e sto m ach A peculi ar .

d arkly s t ai n i ng body ( p ar an ucleus n ebe n ker n ) i s u su ally,

fou n d n e ar th e n ucleus tow ard the outer or b as al e n d of

the cell .

The fluid secrete d by the p an cre as is alk ali n e m ai n ly ,

fro m th e prese n ce of sodiu m c arbo n ate ( Na2CO 3 ) and i t ,

co n t ai n s t hree ferme n ts s teapsin whi ch c au ses a splitti n g o f

,

fat s in t o fat ty acid an d glyceri n a my lop sin w hich co n ver t s ,

s t arch i n to sug ar and t iyp sin w h ich co nverts proteids i n to

,
1 5 2 TH E BIOLOGY OF TH E F RO G CH A P .

pepto n es Th e l at t er differs fro m pepsi n in th a t i t acts in an

.

alk ali n e or n eu t r a l m ediu m ; in a stro n gly acid mediu m i ts

a c t io n is e n tirely stopped .

Th e L iver The live r i s a m assive gl an d whose secre

.

tio n th e bil e is co n veyed to th e i n te sti n e through th e bile

, ,

du ct alo n g w i t h th e
flui d secre ted by the
p an cre as Th e o r .

g a n is of a d ark red

dish color an d is ,

d ivid ed i n to a right ,

a le ft an d a m iddle ,

lobe The middle lobe .

is sm all an d c o n
c ea le d fro m vi ew by

th e he art The left .

lobe is divi ded by an

obliqu e l n c l S l o n I nt o
an an teri or a n d a pos

t e rio r portio n th e la t ,

te r o c c u p y i n g t h e
middle of th e poste
rior p ar t of t h e liv er .

FI G 4 L iv c s ff g D Th e rea t e l F
:
g o r “

. 2 . nd p-
er a an re a o ro . c,

c mm n bil duc t ; D y cys t ic duc ts ; D/ t l o n o f th e h ver 15

o o e c , z,

h p t ic duc t s hich i t h th cys t ic Covered by a Closely

e a ,
w w e

duc t s c mbi t f m th c mm bil

o ne o or e o on e

duc t ; G g l l bl dd ; L L I L L l b s ad here n t l ayer o f p er

,
a a er 2 3 o e

li v tu n d f ds ; L /p h p t it o n e u m W h i c h is
, , , ,

of th e er r e o rw a r i e a o

du d l lig m t ; M s t m ch ; P p
,

o e na a en o a an
co t u ed to
, ,

c s ; P p c t ic duc ts n t i g t h
rea I
,
n an
in rea
fo r m a t e er n e
“

c mm n bil duc t ; Py p yl us ( A ft t ac h m en t s with t h e

o o e , or . er

W i d h im )
e ers e
p er ic ard iu m ve n t r al
.

body w all dors al body w all an d th e stom ach an d i n t es t i n e

, , .

The bile duct is form ed by th e co n flue n ce of the lz ep a tic

1
54 TH E BIOLOGY OF TH E F RO G CH A P .

The liver o f the frog ge n e rally co n t ai n s a co n si de r abl e

a mou n t of pigme n t T w o forms o f pigm e n t occur accord
.
,

in g to L eo nard the bl a ck or d ark bro w n an d t h e golde n

, ,
.

A cert ai n amou n t of pi g m e n t gr an ules occurs in t h e ordi n ary

cells o f th e liver p are n chy m a bu t most of t his subst an c e i s
,

fou n d in pigm e n t cells which are sc a t t ered abou t through

the whol e org an .

E berth h eld th a t th e pigm e n t cells li e withi n th e blood

vessels an d th a t th ey resulted in l arge p art at le ast fro m
, , ,

the tr an sfor m a tio n of leu cocytes P o n fi c k an d L eo n ard .

rega rd them as lyi ng outsid e the blood vessels in t h e p eri

v ascul ar lymph si n uses B r a us ho w ev er fi n ds pigme n ted
.
, ,

cells both in the blood an d in the lym ph vessel s .

There is n o evid e n ce t h at th e pigme n t c ells are d erive d

fro m t h e ord i n ary secreti n g cells o f the liver ( O ppel ) .

C olorless am oeboid cells h ave be e n observed in t h e lymph

sp aces of the liver an d i t i s no t im prob abl e t h at a l arge p art
,

of the pigme nt cells m ay resul t fro m the accu mul atio n of

pig m e n t by such cells which h ave w and ered i n to the l iver
fro m other sources .

The secreti ng cells o f the liver prese n t differe n t appe ar

an c es in rel at io n t o ch an ges in t heir ac t ivi t Th e gr an ules
y .

of t h e cells w ere fou n d by L angley t o i n cre ase in n u mber

aft er a m e al The ch an ges are mu ch more m arke d whe n
.

the cells h ave to s t ar t w ith a s m all ou t er n o n gr an ul ar zo n e

, ,
-

in such c ases in th e ot h to 8 th hou r of diges t io n t h e ou t er ,

zo n e is l arge an d in the 24 t h to 3 o t h the cells become

, ,

gr an ul ar t hroughou t The d ecre ase o f gr an ules w as fou n d

.
”
,

as a rul e to be a ccomp an i ed by an i n cre ase in th e glycoge n

,

in th e cells an d vice ver s a F rom a n alogy w i th th e b eh avior

, .

of simil ar gr an ul es in o t h er gl a n d cells L angl ey co n siders ,

the gran ules in th e liver t o be co n cer n ed in the secretio n of

bile L aho u sse fi n ds t h at gr an ules dis appe ar fro m the cell
.
VII TH E D IG E S T IV E SYST E M A ND I T S FUN CT IONS 1 55

a lmost e n tirely eleve n or twelve hours aft er feedi ng F ive or .

six hours a ft er food is give n t h e liver cells are co n sid er ably

e n l arged and th e c apill ar ies co ngested
,
B y the eleve n th .

F IG Th p h s s f t h h p tic c lls f th f g A c lls ich i

ree a e o e e a e o e ro . e r n

glyc g k f m f g du i g i t Th um us p
.
,

o en ta en ro a ro r n w n er . e re a re n e ro ro

te id g ul s u d t h lum
ra n e d s
ar o l l g f t gl bul s
n e d en , a n e ve r a ar er a o e t o w ar
th ut
e o ds f th c lls B c lls p i glyc g t k f m i
er en o e e .
,
e oo r n o en a en ro a w n

ter f g t h t h d b k p t f t d ys t t m p t u f C T h
ro a a ee n e or en a a a
‘

e e ra re o 22
°
. e

p t id g
ro e ul s sc tt d u i f mly t h ugh u t th c ll M uch th
ra n e are a e re n or ro o e e e

s m pp c is p s t d by t h h p ic c lls f f g i summ
.

a e a e a ra n e re e n e e e at e o a ro n e r.

C c lls e k f m f gs
ta en d f l g t im i summ T h c lls
ro a ro t a rve or a on e n er . e e

sh u k d h glyc g lm s t dis p p d ( F m F s t s
,

h
'
are r n e n an t e o en as a o a ea r e . ro o er

Physi l gy f L gl y )
o o ,
a ter an e .

hou r after feedi n g th e co n gestio n h as dis appe ared and th e ,

cells di mi n ish somewh a t in size .

F u n ct ion s o t h e B e
f il — The bil e which i s secre t ed by .
,

th e cells of th e liver m akes i ts w ay by m e an s of the g all,

 1
5 6 TH E BIOLO G Y OF TH E F RO G CH A P .

c apill ari es to th e h ep atic ducts and th e n ce i n to th e g all

,

bl adder wh e r e i t is store d u n til food p asses ou t of the

,

stom ach whe n i t is disch arged through th e co mm o n bil e duct

,

i n to the i n testi n e B ile i s an alkali n e fluid o f co mplex co m

.

positio n S o me Of its co n sti t ue n ts such as th e fatty sub

.
,

st an ce chol esteri n an d th e bil e pigme n ts are simply w aste

, , ,

products bu t others pl ay a cert ai n p art in digestio n I n

,
.

high e r ve rtebr a tes i t h as bee n show n th at th e bile h elps to

e mulsify fats an d fac ilit ates th eir absorptio n fro m th e i n tes ~

ti n e it also has a sli ght powe r of co n verti n g st arch i n t o sug ar

nt est in al D igest ion an d Ab sorpt ion —Th e food whe n
.

/ I
it is p assed from t h e sto m ach i n to th e duode nu m possess es ,

an a cid re actio n du e to th e a cid it y o f th e g a stric j uice wi t h

which it is mixed I n th e du od e nu m it beco m es mixed with

.

the p an cre atic j uice an d bile both o f which are alk ali n e an d
, ,

i ts acidity i s n eutralized The proteids wh ich m ay h ave

.

esc aped th e ferm e n t ative actio n o f the pepsi n in the stom ach
are ac te d u po n by th e trypsi n o f th e p an cre a ti c j uice an d

co nverted i n to p epto n es Th e s tarchy co n s t itue n ts of th e

.

food are co nverted i n to sugar m ai n ly by the secretio n o f th e

p an cre as though p erh aps also to a slight exte n t by the bile
, ,

an d the fats are p artly split i n to fatty a cid an d glyceri n a n d ,

p artly e mulsified by th e ac t io n o f th e p an cre a tic j uice T h e .

rOle of the i n testi n al j uice in the frog is u n cert a i n ; bu t in

so me of th e high er ver t ebrates i t h as th e p r ope r ty of c o n

ver t i ng st arch i n to sug ar .

Wh e n th e v arious co n stitue n ts o f th e food are digested o r

re n dered soluble by t h e ac tio n of the digestive j uices they ,

are absorbed t hrough th e w alls o f th e i n testi n e i n to th e

blood an d lymph I n th e higher ver t ebr ates m ost o f t he
.

fat i s t ake n u p by t h e lymph vessels o f the i n testi n e an d i t ,

is ge n er ally hel d t h at a l arge p art of t h e sug ar an d p epto n es

is absorbed by th e c apill ari es of the blood vessels Almost .
1 58 TH E BIOLO G Y OF TH E FRO G CH A P .

If the live r of a frog be left for some hours before boili ng

a n d the n tested fo r glycoge n i t will be fou n d th at the
,

a mou n t o f this subst an ce obt ai n ed i s comp ar atively s m all ,

an d i f a ppropri a te t ests be a pplied it m ay b e sho w n th at a ,

cert ai n a mou n t o f d ex t rose h as appe ared in its ste ad Th e .

liver co n t ai n s a ferme n t which has t he power of co n ve r ti n g

glycoge n i n to d extrose ; as t h e ferm e n t is d estroyed by
‘

boili ng a gre ater a mou n t o f glycoge n c an be ob t ai n ed fro m

,

t h e liver if it is boile d soo n a fter i t is removed fro m th e

body .

Th e glycoge n co n te n t o f the liver n o t o n ly i n cr e ases in

the fall an d decre ases in the spri ng and su m mer bu t i t ,

u n dergo es ch an ges in rel at io n to v ari atio n in th e a mou n t o f

food an d to ch an ges of te mperat ure of shor t dur atio n
, .

Afte r fe edi n g th ere is a sligh t i n cre as e in th e a m ou n t of

glycoge n in th e live r ; this slowly dis appe ars i f th e frog is
kept s ever al d ays with ou t fo o d I n wi n ter if frogs in which
.
,

th e liver i s well filled with glycoge n b e kept for a few d ays

in a w ar m room the glycog en co n te n t o f th e liver r apidly
,

d ecre ases O n th e oth er h an d if su mm er frogs which ge n

.
, ,

e rally co n t ai n li t tle glycoge n be kept a t a lo w temper ature

,

for sever al d ays the amou n t of glycoge n in th e liver beco mes

,

m arkedly i n cre ased .

Th e glycoge n stored in the liver m ay be give n ou t slowly

i n to th e blood in t h e form o f d extrose i n t o which i t is ,

ch anged by an e n zyme in t he hep a tic c ells Th e liver ac ts .

a s a sor t o f reservoir o f food stori n g it up in a comp ar atively

,

i n sol uble form wh e n it i s in excess an d expe n di n g it gradu

,

ally to tide over p eriods of fasti n g Th e frog begi n s it s lo n g

.

perio d of hiber nat io n with a l arge rese rve su pply of this

m ateri al which is s lo w ly u sed u p duri n g th e wi n ter an d
,

more r apidly co n sumed in t h e e arly spri ng .

Whil e glycoge n occurs in gre at es t abu n d ance in th e liver ,

VII TH E DIG E ST IV E SYST E M A ND I T S FUN CT IONS 1 59

formi n g at ti m es over 8 per ce n t of th e weigh t o f th at org an ,

i t is fou n d also in m an y other o rg an s of t he body The .

muscles co n t ai n a co n sider ably l ess per ce n t o f glycoge n th an

th e liver ; but owi n g to their mu ch gre ater bulk their tot al
glycoge n co n te n t m ay exce ed th at of the liver al t hough i t is ,

u su ally less S m aller qu an titi es o f glycoge n are fou n d in

.

th e o v aries ce n t ral n ervous syste m an d skeleto n

P eriodic Ch an ges in t h e L iven —The liver of the frog
.
, ,

u n d ergoes i mport an t ch anges in rel atio n to food an d tem

p e ra t u r
. e Th ere i s a regul a r se a so n al ch a n ge which a ffects
n o t o n ly th e size an d ge n er al a ppe a r an ce of t h e org an bu t ,

also the a mou n t o f pigm e n t co n t a i n ed in i t an d th e co n te n ts ,

of th e secreti n g cells I n th e sum mer th e liver is usu ally

.

l arge co mp ar at ively ligh t in color an d fur n ish ed with little

, ,

pigme n t ( Weber E berth L eo nard ) I n th e wi n ter an d

, ,
.

e arly spri ng before t h e feed i n g p eriod th e liver becomes

, ,

rel at ively sm all in size an d dark in color the n u mber o f ,

pigme n t cells i n cre ases an d t h ere are more pigm e n t gr an ules

,

co n t ai n ed in th e se cre t i n g cells M iss L eo n ard who h as

.
,

m ade a study of the p erce n t age of pigme n t in rel atio n to th e

w hol e m ass o f the org an in diffe r e n t times of ye ar arrives at ,

the followi ng result

November 7 per ce n t
, . Ju n e ,
per c e n t
D ecember per ce n t
,
July ,
. 68 per ce n t
April per ce n t
,

I t m ay t hus be se e n th a t the rel a tive a mou n t o f pigm e n t

co n t ai n ed in t h e liver i n cre ases through th e wi n ter , t h e n
dimi n ish es in t h e spri ng after t h e perio d o f feedi n g .

Th e s a me observer fou n d t h at in wi n t er and e arly spri n g

the aver age size of t h e secre t i n g cells an d also their n ucle i
w as sm alles t in e arly spri n g an d i n cre ased duri n g th e
,

summe r as is show n in the follo w i ng t able

1 60 TH E BIO LOGY OF TH E F ROG CH A P .

NO V EM ER B D E EM ER
C B A P RI L

A ve ra g di m
e a et e r . 0 29 2 mm . . 01 62 mm . . 01 2 mm . . 01 72 mm . . 0 27 4 mm .

o f c lls
e

A ve ra g di m
e a et er

o f ucl i
n e . 006 mm . . 00 44 mm . . 007 6 mm . . oo 6 5 mm . . oo 65 mm .

S i m il ar m e asure me n ts by F u nk e gave results approxi m ately

t h e s a m e as those ob t ai n ed by M iss L eo n ard Th e mi n i mu m .

size o f the c ells in R temp or a r ia accordi n g to F u n ke occu rs

.

in M ay th e m axi mu m in July an d August

,
I n R escu len ta . .

t h e mi n i mu m falls in J u n e an d t h e n or m al size is re ach ed

t w o o r t hree m o n ths l at er bu t th ere i s n o well de fi n ed period

,
-

o f m axi mu m gro w th I n both speci es the m i n imu m size o f

.

the liv er c ells a s well as th e live r as a whol e occurs at the

tim e of breedi n g .

The fat co n te n t o f th e liver w as fou n d by F u nke to v ary

in an irregul ar m an n er both in R escu len ta an d R temp o r a r ia . . .

I n the first species th e fat co n te n t of th e liver in m an y

i n st an c es almost e n tirely d is appe ared in J u n e D uri ng the .

su mmer fat i s stored in th e l iver an d in th e wi n te r i t suffers ,

v ery little d im i n utio n i f it d oes n o t actu ally i n cre ase in

a mou n t I n R te mp o r a r ia th e amou n t of fa t in the live r is
. .

very sm all co mp ared with th a t in R escu len ta an d n o defi .

,

n i te co n clusio n coul d be d r aw n reg ardi n g th e ge n er al course

o f its se aso nal ch an ges Accordi n g to L an gley s observ a .

’

tio n s upo n frogs in E n gl an d “ th e fat in the liver cells ,

re a ches i ts m axi mu m amou n t in F ebru ary an d M arch I n .

J an u ary it is as a rule som e w h a t less I n April i t r apidly .

d ecre ases from M ay to D ecember i t is prese n t in comp ara

,

t ively sm all t hough v a ryi n g amou n t s I t is u su ally prese n t .

in mi n i mu m a mou n t in S ep tember an d O ctober

”
.

M iss L eo nard fou n d th at th e r el ative propo r tio n of blood

1 62 TH E B IOLOGY OF TH E F ROG CH A P .

by a slow di mi n utio n du ri ng the wi n ter an d th e n a r apid

falli ng off in the spri n g Th e amou n t of glycoge n in the
.

liver alo n e w as fou n d to i n cre ase an d d ecre ase alo n g w ith

th at of the body as a whole The a m ou n t o f glycoge n in the
.

li ver w as fou n d to be more v ari abl e t h an th e glycoge n c o n

t e n t o f th e rest o f th e body exce edi n g the l atter in th e fall
,

a n d e arly wi n t er while in th e fall th e reve r se rel atio n obt ai n s

,
.

The v ari a t io n s in th e weight o f the liver as a whole h av e

be e n studied in det ail by G aule in R a n a escu len t a The .

weight of t h e liver w as fou n d t o be rel atively gre ater in

m al es t h an in fem ales an d to possess a som ewh at gre at er
r ange of se aso n al v ari atio n s The followi n g t able t ake n from
.

G aule s e stim ates sho w s t h e weight of t h e liver per gr a m of

’

body w eight in th e t w o sexes duri ng the differe n t mo n ths of

t h e ye ar

MAL E F EM EAL

Th e n u mbers in t h e t abl e repres e n t th e aver age weigh ts

of t h e livers an d b o dies o f a n u mber of i n di vidu als ( usu ally
1
5 to 2
5) s a crificed fo r e a ch d etermi n a tio n The
. v a ri a tio n s
VII TH E DI G E ST IV E SYS T E M A ND I T S F U N CT I ONS 1 63

in the size o f th e livers are thus s how n to correspo n d in

ge n er al to th e v ari atio n s in th e glycoge n co n te n t In .

Nove mbe r th e liver m ay beco me betwee n two an d th ree

ti mes as l arge as it is in Ju n e .

F at e of t h e D iff erent Kinds of F ood The fu n ctio n s of .

food as we h ave see n are to bui ld u p tissu e an d to supply

, ,

the org an is m w i th the e n ergy for c arryi ng o n its vit al pro

c esses O nly the pro teids are c ap abl e by themselves o f form
.

in g livi n g ti ssue as t hey alo n e possess all th e nec ess ary

,

ele m e n ts Th e fats an d c arbohydrates howeve r are also t o

.
, ,

a cert ai n exte n t tissue builders but th ey c an supply o n ly ,

three o f t h e el em e n ts of livi n g m atte r ; n am ely c arbo n , ,

oxyge n an d hydroge n
,
.

Th e fat stored in t h e cells of adipos e tissu e m ay b e o b

t ain ed fro m fat co n t ai n ed in th e food bu t it m ay also b e ,

derived fro m c arbohydra tes an d eve n fro m protei ds .

The pri n cip al fu n ctio n s of b o th fats an d c arbohyd rates is

th e p roduc t io n of e n ergy These com pou nds are split u p .

an d oxidized t o c arbo n dioxide an d w a t er yieldi ng e n ergy ,

in this w a for th e perform an ce o f bodily move me n ts an d

y
th e m ai nt e n an ce of th e temperatu re of th e an im al E n ergy .

is also derived fro m th e bre aki n g dow n o f proteid s so t h at ,

i t m ay be s aid th at all o f t h e pri n cip al cl asses of foods are

tissu e build ers and also e n ergy producers After the fo o d .

stuffs h ave pl ayed th eir p art and becom e broke n dow n i n t o

sim pler compou n ds th ey are elimi nated from th e body
,

through th e org an s o f respira tio n and excretio n .

RE F E RE NC E S

Cont ej ea n , C . S u r la dig s t i e on s t o m ach al e de la g ren o uill e . C R

. .

Ac . S ci ,
. P is T
ar , . 1 1 2, 1 89 1 .

Dew evre . No t e sur la fo n t ci on glyco g en iq u e ch z

e la g ren o uill e

d hi
’
ver . C R h eb d S o c B
. . . . i o l. , P is
ar , 1 8 92 .
1 64 TH E BIOLOGY OF TH E F ROG CH A P .

E b ert h , C I . . Die P ig m e n tleb er der F ro sch e und die M e l mia e .

V i ch
r ow
’
s A ch i r v, B d 40 , . 1 8 67 .

G riit z n er, P . Ub e er B ildu g n u nd A ussch e idu n g vo n Fe rm e n t e n .

Ar ch .
g es . P hys .
, Bd . 8 79
20 , 1 .

G riit z n er und S w ie cick i B e . m ku g er n en ub er die P hysi l gi o o e de r

V e d uu n g
r a b ei de n B at rach iern . A ch g B d 49 , 1 8 9 1
r . es . P hys .
, . .

H eiden h ain , M . U b e er die S t ru c t u r dc r Darm ep it h e lz e lle n . A ch r .

mi k . A n at .
,
B d 5 4, . 1 8 99 .

H eiden h ain , R . U n t er such u g n en ub er de n B au der L ab dr u sen .

Ar ch m i k. . A n at .
, B d 6, . 1 8 70 .

J
L an gley , . N . O n the H is t l gyo o an d P hysi o l gy o of t he P p si ne

f mi g G l ds P h il T
or n an . . r an s . Ro y S o c . .
, V ol . 17 2, p art 3 , 1 8 8 1 .

L a n gley an d S ew all . O n th e C h gsan e in P e p n fo r n si -

m i g G l ds an

d i ur n g Se c ti re on . J u P hys
o r . .
, V ol . 2, 1 8 80 .

L eona d, A r . De r E in flu ss der J h sz i t a re b z ll e auf de r L e er e e n vo n

Ra n a t emp or a r ia . Ar ch . A n at . u P h ys p hys
. A b t h S u pp l B d
.
, 88 7 . . . .
, 1 .

M oracz ew sk i, W . V on . Die Z uz a mm t g d L ib s
ense z u n es e e vo n

hu g n er n d en un d blu t m ar en F osch r en . A ch A t u P hys S u p p l B d

r . na . . .
, . .

1 9 00 .

Nu ssb au m M , . U b e er de n B au u nd die T h at igk e it der D us r en .

Arch mi k A n at
. . .
, Bd . 1 3, 1 5 , 1 6, an d 2 1 .

Op p el , A . Le h buch r de r v e r gl ich de en en mi k s k p isch A t m i ro o en na o e .

Pa rt sch , C . B i t ag e r e z ur K e n t n iss de s V d d m s i ig A m p h i
or er ar e e n er

bi en und Rep t ili en . A ch mi k

r . . A n at .
,
Bd . 1 4, 1 8 77 .

S t olk in ow b z ll i sb s d b i d
. V o rg ang e in de n L e er e en, n e on e re e en

P h sp h
o gift u g A ch A t u P hys S u pp l B d 8 8 7
o rver n . r . na . . .
, . .
, 1 .

S w i cick i H U t such u g iib di B i ldu g d A ussch idu g

e , . n er n en er e n un e n

d P p si s b i d B t h i
es e n A ch g P hys B d 3 8 76
e en a ra c er n . r . es . .
, . 1 , 1 .

W eb er C H U b di p i disch F b
, . d g lch . e er e er o e ar e n ve ra n e ru n en w e er

di L b
e d H uh e dd
er F O h l id t B ich t V h KO ig l
er n er u n er r sc e er e e . er . er . n .

Sachs G W iss L i p zig M t h p hys Cl 8 5

. es . . e , a -
. .
, 1 0 .
1 66 TH E B IO L O G Y OF TH E F RO G CH A P .

loop the tr ach e al process w hic h i s b e n t b a ckw ard an d

, ,

se r ves as a me an s of att ach m e n t fo r th e n ecks or roots of

FI G 44 y g —R s p i
s f th f g A
e l sp c T h igh t
ra t o r or an o e ro . ve n t ra a e t . e r

lu g l g h b l i d O p t h h i su f c I B th
. . ,

n r . u ,
as e en a en o S ow t e nn e r r a e . n e

l y xh b t th ugh h middl d h igh t h lf f h l y x

,

ar n as e e n cu ro t e e , an t e r a o t e ar n

and igh t lu g s f m th sid

r y n id c til g ; b by
ar e ee n ro e e . ar ar ten o ar a e .
,

m i p t f th hy id ; g l g l ; 1 t l y g t ch l ch mb ;
,

a n ar o e o , o t t is . r . c, ar n o -
ra ea a er

by p s t i h f hy d ; c l c d (Af H s )
<
’
p . c .
,
o er o r o rn o 01 v . ca , vo a or . te r ow e .

the lu n gs The a ry ten o id ca r tilages are a p air o f semilu n ar

.

v alves which rest upo n the cricoid c artil age ; the ir u ppe r
,

FI G .
45 .
— Cart il ag e s
l y x f th f g A f m b ; B f m of the ar n o e ro . ro a o ve ro

sid ; C y id c il g ; C l c i c id c i l g ; P
, ,

th e e C Z a, ar t en o art a e .
l to .
4 r o a rt a e

x p si c i c id ; Sp s p i us p c ss f h c ic id ;
, ,

e f han on o t e r o no ro e o t e r o

mi c s f h y t ids ( A f W i d h im )
,

p ro n en e o t e ar en o . te r e e rs e .

edges form t he l ater al m argi ns o f the glottis ; th ey .

a fford
VII I TH E VO CA L A ND R E S P I R AT ORY ORG A NS 1 67

a tt ach me n t to muscles by which t h e glot tis m ay be ope n ed

or closed The true sou n d produ ci n g org an s co n sist of a
.
-

p air of el astic b an ds t h e voca l co r ds ex t e n di n g lo n git u di

, ,

n ally a cross the l ary n x Th ey m ay e asily be see n from above

.

by s pre adi n g ap art t h e t w o si des o f the glot t i s or fro m b elow

,

by removi ng t h e m embr an o us floor of th e l ary n ge al c avity .

Th eir med i an edges are t hicke n ed an d lie n e ar e ach o t her

in th e m iddl e li n e S ou n d i s pr o duced by th e exp u lsio n o f
.

air fro m the lu n gs which se t s t h e fre e edges o f th e voc al

cords in vibr at i o n V ari at io n s in t h e sou n d are c aused by

.

alteri n g t h e te n sio n o n th e cords through t h e ac tio n of t h e

l aryn ge al muscl es The voc al app ara t us of t h e m ale frog is

.

much l arge r t h an th a t of t h e fem ale .

Th e m ales of m an y sp eci es of Rana possess a p air of

vo ca l sa cs si t u a t ed at th e sides of th e ph ary n x These s acs
.

are o u t —o c k e t in s o f the ph ry n ge a l w a ll w hich exte nd

p g a

b ackw ard be t we e n the ski n an d t h e body Th ey co mmu n i .

c ate with t h e mou th by sm all ope n i ngs in th e floor a short

dist an ce in fro n t o f th e an gl e o f the lower j aw B esides a .

li n i n g o f mucous m embran e th ey possess a mu scul ar co a t

which co n sis t s of fibers dr aw n o u t fro m th e su b hyo ideu s
muscle The voc al s acs are dis t e n d ed duri n g th e cro aki ng
.

o f the frog through the pressure o f t h e air in th e bucc al

c avi t y
. They serve as reso n at ors t o ree n force the sou n d
produced by t h e voc al cords Th ey are abs e n t in t h e
.

fem ale Their size in th e m ales o f R a n a p ip iens is very

.

v ari able ; in so m e of the v arie t ies o f t his sp eci es th ey are

abs e n t e n t i rely .

The L u n gs — Th e lu n gs are o v o id thi n w alled s acs of

,
-

comp ar at ively simple s tructure Th ey ar e c ap able o f gre at

.

diste n sio n an d m ay be re adily i n fl at ed thr o ugh the glot t is

they do n o t coll ap se w h e n t h e b o dy is c u t o pe n owi ng to ,

t h e fact th at t h e glo tt is u n der ordi n ary circu mst an ces re

1 68 TH E BIOLO G Y OF TH E FROG CH A P .

m ai n s closed Wh e n air i s l e t o u t o f th e lu n gs th ey shrivel

.
,

to an i n co n spicuous size Th e i nn er surfa ce of t h e lu ngs is

.

divid ed by a n et w ork of sept a i n to a seri es of s m all ch a m

bers or a lveoli by m e an s o f wh ic h th e a mou n t o f su rfac e
,

expos ed to th e air i s very gre atly i n cre ased Th e w alls of .

t h e alveoli are richly su pplied with blood vessels which bre ak

up to form a fi ne c apill ary n etwork Th e i n n er surface of .

th e alveoli is co vere d with a si ngle l ayer o f e pi th eli al cells

which are very thi n an d fl atte n ed exc ep t o n t h e edges o f th e
sept a where t h ey becom e cyli n dric al and cili ated O u t sid e
,
.

th e epitheliu m i s a co nn ec ti ve tissu e l ayer whi ch co n t ai n s

the bl o od an d lymph vessels an d n u merous u n stri ated ,

muscle cells which give th e lu ngs t h eir gre a t po w er o f c o n

tractio n The ou ter surfa c e o f the lu n gs is co a ted with
.

p erito n eu m .

The are a o f the i nn er surface o f th e lu n gs o f R a n a escu

len ta h as be e n c arefully c alcul at ed by K rogh I n a speci .

m en weighi n g 4 0 g it w as fou n d to be 9 8 sq c m
. Th e . .

tot al surface o f th e ski n w as estim ated to be 1 5 4 sq c m in . .

th e s a m e sp eci m e n .

Th e Resp irat or y M ovement s — S i n ce th e frog h as n o

ribs i t is u nable t o dr aw in air by e n l argi n g t h e c avi ty c o n
,

t ai n i ng th e lu n gs as th e high er an i m al s d o an d i t h as ,

recou rse th ere fore to a mo re i n direct me t ho d of i n spiratio n

, ,
.

I f o n e w atch es th e respir at ory move m e n t s o f a frog it will ,

be see n th at t h e floor o f th e mou th rises an d falls at qui t e

regul ar i nt erv als U su ally a t so m e w h at gre at e r i n terv als
.

t here m ay b e see n a co n tr ac t io n followed by a sudd e n

exp an sio n of th e b o dy w all an d acco mp anyi n g th e l atter

‘

move me n t th ere is a bri ef closure o f the n ares The respi .

rat o ry movem e n ts o f the frog fall i n to two cl asses ( )

1 the
oscill at ory t hro at movem e n t s and ( 2) th e move m e n ts directly
,

co n cer n ed in filli ng a n d emp t yi ng the lu n gs The thro at .

1 70 TH E BIOLOGY OF TH E FROG CH A P .

c avity a ft er e ach expir atio n fro m th e lu n gs th ey e nable co m

p ar a t ive ly pure a ir to be force d i n to th e lu n gs a g a i n a t the
n ext i n spir a tio n Th e bre ath i n g in wh ich th e lu ngs are
.

i nvolved is i n dic ated by movem e n ts of t h e fl anks or regi o n s ,

abov e an d be hi n d th e fore l egs . C ert ai n sm all m ov em e n t s ,

however occur in t h ese regio n s which ap pe ar to be i n cide n

,

t ally associ ated with the o scill a tory move me n ts o f the floo r
of the mou th an d pl ay n o p art in lu n g respi r atio n the true
fl an k movem e n ts are qui te well m arked Aft er e ach drawi ng.

in o f the fl an k or expir a tio n there follows i mm ed i a tely a

swelli n g o f t h e fl an k du e to i nspirat io n bu t there m ay el apse
,

a co n sider a bl e i n terv al before th e n ext expir a tio n occurs so ,

th at th e lu n gs are alw ays filled with air duri n g th e p ause

bet w ee n succ essive respi r atory acts E xpi r at io n is e ffec t ed
.

by t h e co n tr ac t io n o f the muscles of th e body w all aid ed by

th e el astici ty o f th e w alls of the lu ngs D uri n g th e ac t o f
.

expir a tio n t h e glo t t is o pe n s an d almost i mm edi a tely after

w ards clo ses I f the sides o f t h e body are cut ope n so th a t
.

the lu n gs c ann ot be compre ssed by t h e muscles o f th e body

w all air will be exp elled t hough m ore sl owly every ti m e t h e
, , ,

glottis ope ns A frog thu s o per ated o n i s s till c ap able of

.

bo t h i n spiri ng an d expiri n g air th e m ere el astici ty of the

,

w alls o f th e lu n gs bei n g sufficie n t for th e l at ter fu nc tio n .

I n filli n g t h e lu n gs th e bu cc al c avity acts as a sort o f fo rce

pu mp As the fl o or o f t h e mou t h rises th e n ares are closed
.
, ,

the glo t t is ope ns an d t h e air in t h e bucc al c avity thus sub

,

j ec t e d to pressure a n d h a vi n g n o o th er a ve n u e of esc a pe i s
force d t hrough th e glo t t is in to t h e lu ngs T h e glottis the n
.

closes an d t h e m ovem e n ts o f th e floor of t h e mouth m ay

,

co n ti nue for som e ti m e before t h e n ext in s p ir atio n t akes

pl ace Th e risi n g o f t h e floor o f th e t hro at an d th e closu re
.

of t h e n ares t ake pl ace al mos t at th e s ame t im e th a t air i s

expelled from th e lu ngs ; an d th e exp an sio n o f th e lu ngs
VIII TH E VO CA L A ND R E S P I R AT O RY OR GANS 1 71

follows almost i mm edi ately afterw ard M uch o f th e air .

exp elled from the lu n gs in expiratio n does n o t esc ape fro m

th e bucc al c avity but is forced b ack i n to the lu ngs agai n at
,

th e n ext i n spi r at io n I t i s mixed h owever wi t h th e co m

.
, ,

p artively pure air previou sly in th e mou t h c avi ty Th e val .

vu lar a rr an geme n t for closi n g th e n ar es i s an esse n ti al p art o f

th e m ech an is m fo r filli n g the lu n gs I t w as formerly though t .

th at th e n ares were closed by spe ci al muscles at t a ched to

t h e v alves b u t it w as show n by G au p p t h at this fu n c t io n is
,

performe d th rough r ai si n g th e tip o f th e lo w er j aw t hus ,

el ev ati ng th e pre m axill aries an d th ereby closi n g th ese ope n

i ngs I t m ay be re adily show n th at th e closure of t h e n ares
.

c an b e brough t abou t in this w ay by pressi n g upw ard ag ai n st

the prem axill aries w i t h th e fi nger S o lo ng as it s mouth is .

kep t ope n t he frog is u n able t o close i ts n ares bei ng u n able

to forc e air i n to t h e lu ngs such a frog will soo n er or l at er
,

die of asphyxi atio n D uri n g all o f t h e respir atory m ove m e n ts

.

the m ou th of t h e frog is held tigh t ly clo sed through t he

to n ic co n tr actio n o f t h e muscles o f th e lower j aw As h as .

b ee n expl ai n ed in a previ ous ch apter th e ti p of th e lower ,

w is i n depe n d e n tly m ov able o w i n g to the existe n ce o f t h e

j a ,

sm all m e n t o m eckeli an bo n es w hich are opposed to th e

-

prem axill aries The co n t r ac t io n o f th e sm all subm e n t alis

.

m uscl e which ru n s tr an sversely across th e tip o f the j aw

, ,

c au ses this p art to be r aised above t h e ge n er al level an d

by pressi n g upw ar d ag ai n st the pr em axill ari es closes t he
n a res .

As air i s fo r ced i n to the lu n gs th e pressure in t h e bucc al

,

c avity i s i n dic ated by th e slight pro t rusi o n of th e eyes an d

tymp an ic m embr an es S o metim es h o w ever whe n th e frog
.
, ,

i s m aki ng stro ng i n spir a t o ry effo rts t h e eyes are draw n i n w ard

,

duri ng e ach gulp o f air t hu s aidi ng th e process by di mi n ish

,

ing th e size of t h e bu cc al c avi ty .

1 72 TH E B IOLOGY OF TH E FROG CH A P .

Accordi n g to B aglio n i the exter n al aperture of th e n ares

does n o t re m ai n close d du ri n g t h e l ast ph ases o f th e elev at io n
of t h e floor o f t h e m ou th ; n ever t hel ess air d oes n o t esc ape
,

from th e bucc al c avi ty as m ay be show n by pl aci n g th e n ose

,

o f the frog be n e at h w a ter w he n n o bubbles arise from t h e

,

n os t rils . The muscles which draw th e hyoid app aratus an d

t o ngue fo rw ard an d upw ard c au se t h e ti p of th e j aw to be

depressed w h e n a cert ai n p osi t io n o f t h ese org an s h as bee n

re ached an d th e nares ope n Why th e n does n o t air p ass
.
, ,

ou t o f the n ares as the flo o r of t h e mou t h c o n t i n u es to be

r a ise d ? As B aglio n i m a i n t ai ns this i s bec ause t h e n as al
,

p ass ages are closed fro m behi n d by m e an s of t h e an terior

processes o f the hyoid c artil age w hich are so formed and

,

situ ated t h at t hey fi t n e atly i n to th e posterior n ares as th e

hyoid app ar at u s is dr aw n u pw ard an d forw ard in the ac t
of i nspir atio n.

Ch an ges in t he B lood in Respir at ion — Th e respir atory

move me nt s th at h ave bee n d escri bed are su bsidi a ry to keep
in g fresh air in close rel at io n wi t h th e blood O n t h e o ne
.

h an d w e h av e th e org an s o f re spir atio n an d th e d istributio n

wi t hi n th e m o f th e blood vessels which are so arr an ged th a t
,

th e blo o d is bro u gh t very close to t h e sur fac e over a l arge

a re a . A n d o n t h e oth er h an d we h ave a c o m plic ated an d
be au t ifully ad ap t ive m ech an ism for ke epi ng the l arge portio n
of th e respi rat ory surface i n clud ed in th e lu ngs in co n t act
w i th pure air . Th es e d evic es facili t ate th e exch an ge o f
g ases wh ich t akes pl ace bet w ee n t h e air an d th e blood by
me an s of diffusio n across t he i n t erve n i n g m embr an es T h e .

bl ood receives oxyge n from t h e air an d g ives off c arbo n

dioxide so th at t h e air which has bee n expired fro m the
,

lu ngs o r bucc al c avi t y always co n t ai n s less of th e form er an d

m ore o f t h e l atter gas Th e process o f res pir at io n falls
.

i nto two ph ases : ( 1 ) exte rnal r esp i r a tio n o r t h e exch ange

,
1 74 TH E BIOLO G Y OF TH E F RO G CH A P .

practi c ally t h e o nly respi rato r y org an F rogs m ay be kept

.

alive whe n subm erged in w at er at o to 1 3 C for sever a l d ays

0 °
. .

At a high er t emper a ture frogs t e n d t o com e to th e su rface

o ft e n er fo r air an d i f preve n ted fro m doi ng so t h ey m ay
, ,

die of asp hyxi at io n Th e ski n fu n c t io n s as a respir atory

.

org an bo t h in w a ter and in air I f th e n o s t rils of a frog be

.

plugged wi th w ax the an i m al m ay be kep t alive in co ol air

,

for sever al d ays .

Th e experi m e n t s o f seve r al i n vestig ators h ave S how n th at

more c arbo n di oxi de i s give n off t hrough th e ski n th an t h r ough
the lu n gs K lug fou nd th a t t h e r at io o f CO Q give n off by the
.

lu ngs to th at give n off through t he ski n v aried in t h e differe n t

speci m e n s i nves t ig ated fro m 1 to Th e frogs
wh ich K lug experi m e n ted u po n we re pu t in a ch a mber di vided
by a p ar t itio n which co n t ai n ed a n aperture surrou n d ed by rub
ber Th e frog w as pl ac ed so t h at it s h e ad proj ec t ed t h rough
.

th e p artitio n and w as h eld tigh tly by the rubber so th at o n e

,

ch amber w as co mpl etely shu t off fro m th e o t h er Air w as .

p assed through bo t h ch a mbers an d t h e amou n t o f c arbo n

,

dioxid e give n off i n to e ach m e asured an d comp are d Th e .

o n e ch a mber rec eived th e outpu t fro m t h e ski n o n ly w hil e ,

the other rec eived th at o f th e lu ngs toge t her w i t h t h e sm all

a mou n t exh aled fro m t h e ski n o f the h e ad The m ethod of
.

K lug w as an improveme n t over those o f his pred ecessors ,

al t hough n o t e n tirely fre e fro m obj ec t io n s th e pri n cip al o n e

,

bei n g th at th e pressure o f th e rubber n ecess ary to produc e

an air tigh t fi t would i mp ed e the n orm al movem e n ts o f res
-

p ir at io n E.xperi me n ts of lig a ti n g or ex t irp ati n g th e lu n gs ,

removi n g th e ski n tyi n g t h e cut an eo us blood vessels plu n g

, ,

ing the frog in oil n e arly up to it s n ostrils etc in order to ,

.
,

elimi nate o n e or th e o ther m odes of respi r atio n are all op e n to

th e s am e criti cism th at they do no t tell u s an y t hi ng o f th e
rel at io n of ski n an d pul mo nary respi r at io n u nd e r ordi nary
V 111 TH E V O CA L A ND R E S P I R AT ORY ORG A NS 1 75

co n ditio n s I f c arbo n dioxide i s preve n ted fr o m esc api n g

.

through t h e ski n m ore o f i t will be exh aled through the lu n gs

, ,

or if th e lu ngs are tied m ore c arbo n dioxide will be elimi nated

,

through th e ski n .

The rel atio n betwee n th e cut an eou s and pulmo nary respi
ratio n o f th e frog has rece n tly bee n quite exh austively studied
by K rogh Th e lu n gs were supplied with air by m e an s o f
.

ar t ifici al respir at io n an d the i n co m e of oxyge n an d t h e out

,

pu t o f CO , fro m bo t h th e lu n gs an d ski n co m p ared u n der

v ariou s co nditio n s I n R a n a f usca at a temp erature of 20 C
.
°
.

th e aver age r atio of oxyge n i n co m e to CO , outpu t in several

experim e n t s o n frogs t ake n at differe n t ti mes o f ye ar w as ,

in pulmo n ary respi ra t io n O , 1 0 5 CO , 4 5 in cut aneou s

,

respir atio n O , 5 2 CO 1 2 9 I t is thus evide n t th at in th e

, .

lu n gs th e oxyge n t ake n in is gre atly in excess o f th e CO ,

give n out whil e in th e ski n th e reverse rel atio n obt ai n s I n .

Ra n a escu len ta rel at ively m ore oxyge n i s t ake n in through

th e ski n an d rel a t iv ely l ess CO , eli mi n ated through the lu n gs .

T he r espir atory quo t ie n ts ( i e r ati o of O , to CO , ) in th e two

.

speci es at 2 0 C are as follows

°
.

CU E
T A N O US P UL M O NA RY
RES PI RA T I O N R E S PI RA T I O N

R . escu len t a

I n fl u en ce of E xt ern al Con dit ions u p on Respirat ion .

Th e respir atory fu n c t io n s of both t h e lu n gs an d th e ski n v ary

in differe n t periods of t h e ye a r eve n w h e n th e a ni m als are
pl aced u nd er t h e s a m e d egre e of temper ature Th e a mou nt .

of oxyge n t ake n in by the lu ngs is gre at es t duri n g t he breed

in g se aso n ; the n i t r apidly d ecre as es duri ng t h e su mm er ,
1 76 TH E BIOLOGY OF TH E FROG CH A P .

and re aches i ts mi n i mu m in t he wi n ter t he r atios o f oxyge n ,

absorp tio n a t a temper a ture o f 2 0 C bei ng as follo w s

°
.

spri ng ,
su mm er 8 2 wi n ter 5 4 Th e outpu t o f C O ,
, ,
.

by the lu n gs v aries in a simil ar m ann er ( sp r i n g 6 2 su m mer , ,

4 ;2 wi n ter
,
Th e cut a n eou s r espir a t io n i s subj ec t to
much less s e aso nal v ari atio n ; th e absorptio n o f oxyge n is
prac tic ally co n sta nt the eli mi natio n of CO , is co n si der ably
i ncre ased duri ng the bre ed i ng p erio d bu t for the res t o f th e
,

ye ar it v aries but litt le While th e a mou n t of oxyge n t ake n

.

in by th e lu ngs du ri n g th e spri ng a nd su m me r co n sid er ably

exceeds th at absorbed by th e ski n the cut an e ous absorptio n
,

o f oxyge n beco mes much gre ater th an the pulmo n ary in the
wi n ter I n wi n ter th ere fore th e ski n beco mes r el atively
.
, ,

m ore i mpo r t an t in respir atio n th an duri n g th e r es t o f th e

ye an
Wheth e r the ski n fu n ctio n s mo re e fficie n tly as a r espir a
tory org an in air or in w ater th e few an d co n tr adictory
results o f B oh r an d K rogh do n o t e nable o n e to d etermi n e .

F e w experi m e n ts h ave bee n m ad e u po n t h e rel at io n betwee n

temperature an d t h e r apidity of respi ratio n al t hough i t is ,

k now n th at r espiratio n t akes pl ace m uch m ore r apidly w h e n

th e temper a ture i s i n cre ased At low temper at ures respi
.

ra t o r
y ch a n ges are sligh t .

M o le sc ho t t an d F u b i n i h av e show n th at ligh t h as a m arked

e ffec t upo n res p iratio n o f th e ski n the a mou n t of CO , pro
,

du c e d at a give n t emper at ure bei n g m uc h gre at er in th e

light th an in th e d ark This w as h eld to be du e in p art to
.

a direct actio n o f ligh t u po n the ski n bec a use th e i n cre ase ,

occurs in frogs wh ose eyes have bee n removed al t hough to ,

a less exte n t t h an in n orm al speci me n s Th e m ore re frangi

.

ble r ays h ave t h e gre a test effec t u p o n ski n respirat io n as ,

w as show n by me asuri n g the CO ou t pu t in frogs exposed t o

,

di ffere n tly colored lights The r ati os o f CO , produc t io n u n der

.
1 78 TH E BIOLO G Y OF TH E FRO G CH A P .

M il n e E dw -
a r ds , H . De la i flu c
n en e des a g s p h ysi q u s l
en e su r a v e, i
P isar , 1 8 24 . L e go n s su r la h
p ys io lo i
g c et l t mi c m p é
’
an a o e o ar e de
l h mm im ux 8 5 7—1 8 65
’
o e et des an a ,
1 .

M oles ch ot t and ull i flu z d ll luc

F ub in i . S
’
n en a e a e m is t ch m t ic
a e ro a a

nell s l z i
’
e a a o ne di cid c b ic p 1 g ism
a o ar on o er
’
or an o an im l At t i d ll
a e . e
’

Ac d T i
a . or no, 1
5 , 1 8 79 .

Regn au l t et Reis e t . Re ch ch s ch imi q u s

er e e su r la re s pi t i ra on .

A nn . ch m e . et p hys .
, S er 3 , 26 T . .

W eden sk i, N . U b e e r die At h m u n
g des F sch es
ro . Ar ch .
g es . P hys .
,

Bd . 2
5 , 1 88 1 .
1 11 TH E S KI N 1 79

CH A PT E R I X

TH E SKIN

E xt er n al Ch aract er s —
The
ski n i s an org an of u n u su al
.

i mp o r ta n ce in t h e life of th e frog bec ause in addi tio n to the

, ,

fu n c t io ns which i t com mo n ly p erforms amo n g o ther an i m als ,

it h as a n u mber of speci al fu n ctio n s w hich are p eculi ar t o

th e A mphibi a an d which in m ost c ases re ach their fu llest
, , ,

develop me n t amo n g t h e A n u ra As in m ost o f t he Amphibi a

.
,

th e ski n of t h e frog is smoot h an d moist ; it is very loosely

a tt a ched to t he u n d erlyi n g muscul a ture by thi n b an ds of c o n

n e c t ive tissue wh ich sep ar a te th e l arge subcut an eous lymph

,

s p aces I t is every w here very tough but i t i s co n sid er ably

.
,

t hicker o n th e dors al sid e of th e bo dy th an i t is below In .

c ert ai n regio n s it prese n ts speci al th icke n i ngs ; such as th e

derm al plic ae which exte n d b ackw ard from n e ar t h e posterio r
,
'

an gles o f t h e eyes t h e su b ar t icu lar p ad s be n e ath th e j oi n ts

,

of t h e digits of th e feet th e swelli ng at th e b ase of th e first

,

fi nger of the arm t h e pro t uber an ce over th e six t h toe or p r e

,

h allux an d t h e upper eyelids an d lips S m all p apill ae ofte n

,
.

occur especi ally o n the dors al sid e of th e body some of

, ,

which th e t ac tile p apill ae are perm an e n t o thers th e sexu al

, , ,

p apill ae o f th e fe m ale occur o nly duri n g th e breedi n g period

,
.

H ist ological S t ru ct ure — The S ki n is composed of two

.

pri n cip al l ayers t h e ep ider m is an d t he cor iu m or cu tis A

, , ,
.

third l aye r of subcut an eous co n n ec t ive t issue n o t belo n gi n g ,

to th e ski n proper li es u n der n e ath t h e coriu m and forms th e

,

sept a u ni ti n g the ski n to the body w all .

1 80 TH E BIOLOGY OF TH E F RO G CH A P .

The ep ider m is or outer portio n o f th e ski n i s co mposed

, ,

o f severa l l ayers o f cells Th e cells o f th e i n n ermost l ayer

.

are colu m n ar ; b u t in p a ssi n g tow ard th e oute r surface th e

cells beco m e m ore an d more fl atte n ed u n til those of th e ,

ou t ermost or hor n y l ayer ( st r a t u m cor n eu m) beco me very

bro ad and th i n I t is the str a tum cor neu m th at i s sh ed duri n g
.

the m o lti n g process The gr adu al ch ange in sh a pe be t w e e n

.

the c ells o f th e i nn er an d ou ter surfac es o f t h e epi dermis

i s due to th e fact th a t there i s a co n ti n u al productio n of n ew
c ells in th e i nn e r l ayer wh ich are gr adu ally push ed ou t w ard ,

becomi n g more an d more fl a t t e n ed th e farth er th ey are

pressed aw ay fro m th eir poi n t o f or ig in .

Th e epidermis esp eci ally o n th e dors al S id e o f the body

, ,

usu ally co n t ai n s m ore or less d ark brow n o r bl a ck pigme n t .

This pigme n t i s p artly w ithi n speci al cells th e ch r om a to ,

p h or es
,
a n d p a rtly in a n d be twee n th e t ypic al cells o f th e
epid ermis I n cert ai n regio n s all o f the cells o f the epi
.

dermis m ay co n t ai n sm all pigme n t gr an ules E r m a nn fou n d .

t h at in the s am e regio n o f epide r mis p igm e n t would appe ar

an d dis app e ar in th e course o f a fe w m o n th s The c h ro .

m at o ph o res of th e epidermi s res emble t h e d a rk pigme n t

cells of th e coriu m W he ther t hey are d eriv ed fro m cells
.

o f the co riu m which h ave w an dered i n t o t h e e pidermis o r ,

whe t her t hey aris e through th e tran sfo rm a tio n o f cells o f t he

epidermis i t self is a m at ter o f c o n t ro versy L oeb an d S t ro n g 1
'

.
,

h ave co me to the co n clusio n th at t h e chro m a t o phores t h a t

a ppe a r in t h e rege n er a t ed epitheliu m of th e fr o g ar e d erived

fro m epi t heli al cells an d no t fro m c ells t h a t h ave wan d ered

,

in fro m t h e cu t is C hro m atophores in t h e epidermis are

.

n o t u su ally a bu n d an t The m ai n source of t h e color o f t h e

.

ski n is in t h e pigme n t c ells o f t h e corium .

The i nn er l aye r o f t he epid ermis co n t ai n s several s t ell ate

1 L oe b an d S tro n g , Am f . ou r . A n at V ol 3 p
.
, .
182 TH E BIOLO G Y O F TH E F RO G CH A P .

The str atu m comp act u m is m ai n ly com posed of a de n se

l ayer of co n n ecti ve tissu e w hose fibers ru n in a w avy course
,

p ar allel to the surface of the ski n At i n terv als this l ayer

.

is crossed by vertic al str an d s which o ft e n exte n d throu gh

,

th e str atu m spo n giosum i n to th e epiderm is I n additio n to .

fibrou s co n n ective tissu e t h ese str an ds frequ e n tly co n t ai n

,

s mooth mu scle fibers el astic fibers n erves an d blood ves

, , ,

sels I t is prob ably du e t o th e co n t r actio n o f these muscl e

.

fibers th at th e p ap illat io n of t h e ski n i s prod uced a ft er ce r

t ai n co n di tio n s of sti mul atio n The fibers m ay also aid in
.

S qu eezi n g ou t th e secretio n of th e cu t an e o us gl an ds .

The subcu t an eous co nn ec t ive tissue forms a loose l aye r

be n e ath t he str atu m comp actu m and a seco n d very th i n
l ayer n ext to th e muscles th e two l aye rs bei ng sep arated by
,

l arge lymph sp ac es except in the sept a wh ere th ey becom e ,

co n ti nu ous The oute r o f t h e two l ayers i s very v ascul ar

.

an d co n t ai n s n u merous stell ate c e lls wi thi n which are n u

,

m ero us gr ayish wh ite pigme n t gr an u les Thes e cells are .

esp eci ally abu n d an t o n th e ve n tr al sid e of th e body wh ere ,

they pr o duce the white coloratio n ch ar acteristic o f th a t

r eg io n .
_

G lands of t he S k in Th e ski n of th e frog like th a t

.
,

of m ost o f th e Amp h ibi a i s richly fur ni sh ed with gl an ds

,
.

These gl an ds are o f t h e si mpl e alveol ar type an d lie m ai n ly ,

in the str atum spo n giosu m of the coriu m O nly r arely as .

,

in the l arge gl an ds of the i nner fi nger do th ey exte n d i n to ,

th e d eeper portio n s o f t h e ski n Typic ally the gl an ds are

.

spheric al or ov al in form an d ope n to the surface through

,

a n arro w n eck whic h exte n ds through th e epidermis an d

termi nates in th e trir adi ate ope n i n g of a so c alled stom a -

cell at i ts ou ter en d .

.
‘

The ski n gl an ds of the frog h ave bee n studied by n u m er

ous i n v es t ig a tors bu t th ere rem ai n th e wid es t di ffere nces
,
ix TH E SKIN 1 83

of o pi n io n reg ardi n g m an y of the m ost i mportan t fe ature s

o f their s t ructure an d fu n c t i o n s Two v arie t ies of gl an d are
.

com mo n ly disti n guished wh i ch m ay be desig nated as th e

m u cu s g la n ds an d th e p oiso n g la n ds Wh ile H eide n h ai n .
,

Nico glu an d oth ers reg ard these t w o ty p es of gl an d as

,

specific ally disti n ct othe r i nvestig ators ( C almels L eydig

, , ,

S c z c esny Ju n ius ) co n sider the m as differe n t ph ases in th e

,

develop me n t of o n e an d th e s ame gl an d H owever this .

m ay be the gl an ds of the frog s ski n m ay be g ro u p e d in t o

'

’
,

t w o cl asses whic h are structur a lly an d fu n ctio n ally differe n t ,

and we sh all describe th em sep ar at ely withou t reg ar d to the

qu estio n as to wh ether they are ge n e t ic ally co n n ected .

Th e m ucus gl ands are sm alle r an d much more abu n d an t

th an th e poiso n gl an d s an d are fou n d over practi c ally th e
,

e n t ire surface of the body I n som e pl aces they are s o thick

.

th at th ey n e arly touch I n Ra n a fu sca accordi n g to E n gel

.
,

m an n they average abou t six t y to e ac h squ are m illi m eter o f

,

surface Thei r ducts are n arrow an d li n ed wi t h a l ayer o f

.
,

sm all fl atte n ed epi t heli al cells Th e body of the gl an d is

.

li n ed with epitheli al cells which form a si n gle l ayer except

n e ar th e op e n i n g of t h e n eck where there are two l ayers
, .

I t i s this e pithelium which forms the mucus which is dis

ch arged i n to th e lu me n o f t h e gl an d an d poured out through ,

the n eck ove r th e surface of th e ski n The appe ar an ce o f .

t h e secreti n g epitheliu m v aries gre a tly in differe n t gl an ds .

I n so m e c ases m o r e oft e n in the sm aller gl an ds the epi

, ,

t h elial cells are low cle arly m arked off fro m e ach other an d
, ,

from th e l arge lu m e n of t h e gl an d an d co n t ai n n uclei which

,

t a ke up a l arge p art of the cell I n other gl an ds th e cells

.

ar e elo n g ated so th at they fill a l arge p art o f th e lume n t h e ,

n ucleu s is rel atively sm a ll an d situ a ted n e ar the b ase of t h e

,

cell and n u m erous gr anules occur tow ard th e free e n ds

,

Duri ng secretio n th ese gr an ules swell up an d become c o n ,

1 84 TH E BIOLO G Y OF TH E FROG CH A P .

verted i n to a tran sp are n t subst an ce which is disch arged i n to

the ce n tr al c avity ( B iederm an n ) an d i t is prob able th at th ey ,

represe n t a st age in th e fo rm atio n o f m ucus Nu merous .

tr an siti o n al st ages be t w ee n t hese two v ari eti es of epi t heliu m

occu r an d i t is qui te cert ai n ,

th at the differe n ces are du e

to t h e age of the gl an ds and ,

th eir d iffere n t st ates o f secre

t io n C h an ges in th e form .

o f the c ells however are pro , ,

du c ed to a cert ai n exte n t by
t h e co n tr actio n of th e gl an d .

O utside o f the epi t heliu m

is a mu scul ar co at co mpos ed
o f smooth m uscl e cells whi ch

li e in a m eridio nal directio n .

Th e outermost co at o f th e
gl an d i s form ed by a l aye r
o f fibrous co n n ective tissu e .

Th e fu n ctio n o f th e muscle
cells i s th e expulsio n of th e
—
secretio n of the gl an d Th e
C ss s c t i n f t h ski n
.

FI G
47 ro e o o e
gl a n ds O f the S kl n are I n C0 11
. .

o f th f g D d mis
e ro .cu t i s ; E, er or ,

e p d m i ; b v bl
l er S d v ss l ; -
st an t mo tio n ( Asch erso n E m
oo e e ~

cu t us gl n d t t h ugh t h
.
,
an eo a cu ro e
a s m ay be see n by
Ge m“; C g l t h s m f m -
’
.
g e lm
e
ann
)a e ro o ne
,

sid ; a duc t f gl n d ; h f l f an ex ami n ati o n of the gl an ds

e , o a z .
'

h iz t l fib s f c n c in th e web o f th e foot Th ey
,
”
hf , or on a er o o ne

ti t i ssu ; h l ut h yl y .

ve e o er o r o rn a er

id mis l gh i
.
,

o f th p e e ; m l M er p i C h a n ge n o t o n ly in size bu t
a an ,

l y f th p id mis ; pg p i gm t also in form bei n g n o w

.
,

a er o e e er , en

c lls " ( A ft H ”
e er ow e
,

rou n ded an d n o w wri nkled

an d an gul ar C o n tr actio n m ay b e c au sed by stimul atio n of
.

t he ski n with irrit an t solu tio n s or by th e electric curre n t .

Th e poiso n gl an ds are l arger an d le ss abu n d an t th an th e

1 86 TH E B IOLOGY OF TH E FROG CH A P .

a bur n i n g t aste I t m ay b e c aused to exude fro m th e ski n

.
,

esp eci ally of the bull frog by pl aci n g th e an i m al u n der

,

chloro form O f its prop er t ies in th e frog comp ar a tively

.

little i s k n ow n P aul B er t fou n d th at a go ldfi n c h which w as

.

i n ocul at ed with the derm al secretio n o f Ra n a escu len ta died

withi n On e mi n ute an othe r frog o f th e s a m e species which
w as i n ocul a ted with th e poiso n d ied wi thi n an hour an d a

qu arter .

I n m an y othe r A mphi b i a especi ally t he to ads an d s al a

,
~

m an ders poiso n gl an ds are very ex te n sively developed an d

, ,

yield an abu n d an t secretio n

S ex D if f erences —Th e ski n o f th e frog p r ese n ts cert ai n
.

differe n ces ch ar a cteristic o f sex so m e of whi ch are perm a ,

n e n t while oth ers occur o n ly duri n g th e bre edi n g period

,
.

I n Ra n a f usca accordi n g to L eydig an d in R a r va lis

, , .
,

a ccordi ng to S t ee n st r u th e web o n the hi n d fee t o f t h e

p
'

m ale s is more fully d evelop ed th an in th e fem al es Th e .

swelli n g o n th e i n n er side of t h e firs t fi n ger o f t he m ale ,

which h as bee n m e n tio n ed in a previous ch apter is c aused ,

by modific atio ns both o f t h e coriu m and the epidermis .

This swelli ng is much l arge r in the breedi n g period th an at

oth e r tim es an d it doub t less subserves th e fu nc t io n of aid
,

ing the m ale to re t ain h o ld o f th e fem ale The cut an eous .

gl an ds in thi s regio n are much e n l arged an d beco me elo n ,

gated i n to a tubul a r form an d ex t e n d through the e n t ire

,

thick n ess Of th e Ski n Th e epidermi s in the bre edi n g period

.

is prolifer ated to form s m all p apill ae wi th a thick rough , ,

hor ny l ayer Aft er th e breedi n g p eriod th e epid ermis

.

beco mes smoo t h ag ai n and t h ere i s also a p arti al di s appe ar

,

an ce of th e pigm e n t o f the coriu m so th at t h e s w elli n g los es

,

its d ark color .

The o ccu rre n ce of d erm al p apill ae in t h e fem ale o f Ra n a

f usca duri ng th e bre edi ng perio d h as alre ady be e n su fficie n t ly
ix TH E SKIN 18
7

described ( see C h apter I I ) The m ales o f cert ai n species

‘

assu me at this time a blue color atio n w hich appe ars m a i n ly

o n th e ve n tr al side o f th e body I n Ra n a a rva lis ( R oxy

. .

r r h in u s ) i t h as bee n describe d by S t een st ru p and by S iebold .

I n Ra n a fu sca P alio described a blue colo r atio n a ppe ari n g

o n the thro at duri n g th e breedi n g p eriod L eydig fou n d
.

th at this color dis appe ared soo n aft er t h e anim al w as t ake n

fro m th e w at er B oth L eydig an d H aller who studied the
.
,

s a m e phe no me n o n in R a n a temp or a r ia regard th e blu e as

,

an i n terfere n c e color produced by mi n u te gr an ul es in the

ski n I t is prob abl e th at the appe ar an ce o f th e blue color

.

is associ ated w i th the absorptio n o f w ater F rogs which.

h ave lost th e blu e color whe n kept in t h e air soo n reg ai n it

whe n pl aced in the w at er ag ai n Aft er the bre edi n g period
. .

is over th e blue color quickly dis appe ars A reddish brow n

,
.

color duri n g t h e breedi n g se aso n h as bee n described by

L eydig in t h e fem ale o f R a n a fu sca an d S mi t h h as observed
,

a blue color a tio n of th e thro a t which h e reg ards as c h ar ac

t erist ic of the fem ale o f th at speci es at this ti m e .

The ski n o f th e m ale o f R a n a fu sca in the bre edi ng sea

so n beco mes swolle n an d m ay h an g do w n at the sides ,

assu mi n g wh a t L eydig desig n at es a q u a m m ig q u a i e

pp g s
-

A n sehe n . Th e str a tu m co mp ac t u m of the coriu m becomes

more or less gel ati nou s and th e subcu t an eous lymph sp aces
beco me filled wi th a m ateri al r ese mbli n g the V itreous hu mor
o f the eye .

Wi t h the e xceptio n of t h e swolle n first fi nger o f th e m ale .

an d t h e derm al p apill ae of t he fe m ale th ere i s n o evide n ce

as to wh a t fu n ctio n al sig nific an ce the above ch ar a cters pos

sess i f they possess any Th ey m ay be th e i n cide n t al prod

,
.

u c t s of th e i mport an t co n stitu tio n al c h anges whic h t ake

pl ace duri n g the breedi n g period wi t h o u t bei n g o f any
,
-

d i r ect v alu e to the o r gan ism .

1 88 TH E BIOLO G Y OF TH E F ROG CH A P .

S easonal Changes . M ost

o f th e se aso n al ch anges in th e
ski n are correl ated with th e sexu al differe n ces th at o ccu r
du r i n g th e breedi ng se aso n an d h ave bee n tre at ed u n der
,

th at he ad Th ere are som e othe r se aso nal ch an ges h ow ,

ever which occu r app are n tly wi thou t r eg ard to t h e develo p

,

m e n t o f the sexu al products I n the wi n ter an d e arly

.

spri ng frogs are d arke r in color th an in sum m er owi n g ,

prob ably in l arge p art to di ffere n ces o f temper at ure Ac .

cordi n g to D o n aldso n th e powe r o f the ski n to absorb w ate r

is g r e ater in su m mer th an in W in ter
Color Ch an ges —Th e power of th e ski n to ch ange i ts
.

colo r in r el atio n to su r rou n di n g co n diti o n s depe n ds upo n

ch an ges which o ccu r in th e pigm e n t cells o r chrom ato ,

p h o r es .O f th ese there m ay be disti n gui shed th e follo w i n g

v arieti es : bla c/z p igmen t cells ( melanoph or es) in terfer en ce ,

cells ( leucOp h or es) g

,
o lden
p igm en t cells
(x an t h Op h or es,

xant h oleu COph or es) , an d in som e speci es of frogs r ed p ig

men t cells .

Th e bl ack chrom atophores are stell ate c ells wi th irregu

larly br an chi n g p r ocesses Th ere is a si ngle n ucleus n e ar
.

th e ce n ter of th e cel l Th e d ark pigm e n t is in th e form

.

of n u m erous s m all brow n or bl ack gr an ules of a subst an ce

c alled mela n in which is a very resist an t compou n d rem ai n
,

ing u naffected by m ost re age n ts The bl ack chro m ato

.

p h o re s a r e m ost a bu n d an t o n the dors a l sid e of th e body ,

esp ec ially in th e bl a ck spots wh ere th ey ar e m assed togeth er

'

very thickly O n th e ve n tral sid e they are al most e n t irely

.

abse n t over a co n sider able are a They are fou n d mos t ly in

.

the superfici al l aye r o f the coriu m j us t below the epide r mis .

S c attere d chrom a tophores occur in t h e epi dermis an d the

de eper l ayers o f the coriu m Th ey t e n d to aggreg at e in
.

regio n s which ar e most abu n d an t ly su p p lied with blood

vessels Th e pigme n t of t h e chrom at ophores u n dergoes
.
1 90 TH E BIOLOGY OF TH E FROG CH A P .

i nvisible e xcept wh e n co n t ai n i ng pi gme n t V irchow V o n .

,

Wi ttich and B ied er m ann thi nk th a t th e ch an ges in th e

,

chrom atopho r es m ay i nvolve bot h o f t hes e facto r s Th e .

qu estio n is still n o t cert ai nly decided .

The cells which give th e ski n its golde n an d gree n colo r s

form a l ayer i m m edi ately be n e ath th e epidermi s U nlike the .

bl ack c hrom atophores th ey are u su ally rou nded or p o lygo

n al in fo r m a n d the li lit t le above the bl ack cells which
, y e a ,

co n stitu te a sort of d ark b ac k g r ou n d Th eir gold e n color is

.

due to a fatty pigm e n t or lipochro m e w hich is so m etimes ,

diffuse d throughout t he cell and at o ther ti mes aggreg ated

i n to l arge drops ( B i ed erm an n ) This pigm e n t is soluble in
.

alcohol chloroform an d eth er givi n g a golde n yellow solu

, , ,

tio n w hich tur n s to a yello w ish gree n w h e n v ery dilu te .

Th e s am e subst an c e accordi n g to K uh n e produces th e

, ,

yellow color o f th e fat body I n frogs wh ich h ave bee n p r e

.

s erved fo r so me tim e in alcoh ol t his pigm e n t dis appe ars ,

an d co n seque n tly the sp eci m e n s lose thei r golde n an d g r ee n

color atio n .

The golde n cells u su ally co n t ai n an addi tio nal source of

color in the form of the so c alled i n t erfere n ce gr an ules or
-

the iri desce n t pigm e n t of L eydig These gr an ules accordi ng

.
,

to E w ald and K ruke n berg are composed of gu an i n Th ey

,
.

are soluble in c au stic sod a or pot ash an d prese n t a n appe ar

an ce of cross st r i atio n B iede rm B y r an smitted ligh t

( an n
) t .

they are brow n o r g ray but in refle ct ed ligh t they are

,

usu ally blue .

The i n terfere n ce Cells are stell ate chro m atophores which

a re m ai nly co n fi n ed to the subcut an eous tissu e of the ve n tr al

side of the body w h ere t h ey produce th e ligh t color ch ar ac

,

t erist ic of th at regi o n They co n t ai n gu ani n g ran ules like

.

those in most of th e gold e n cells .

Red stell ate pigme n t cells h ave bee n described in R a n a

TH E SK IN 1 91
1 92 TH E BIOLOGY OF TH E FRO G CH A P .

f usca by V on Wittich Th ey occur in th e coriu m an d were

.
,

obser ved to u n dergo ch anges in th e distributio n of their

pigm e n t like t hose of the bl ack ch ro m atophores .

Ne arly all o f the color c h anges which th e ski n o f th e frog

u nd ergo es depe n d upo n t h e diffe re n ces in th e distributio n
o f t w o ele m e n ts th e bl ack an d t heyellow pigme n t Wh e n
,
.

t h e pigm e n t o f t h e bl ack ch ro m at o phores is exp an ded t h e ,

ski n beco m es d ark in color owi ng to t h e fact th at t h e bl ack

,

pig m e n t is spre ad over a gre ater a mou nt of su rfac e Whe n .

the ski n is ligh t in col or th e bl ack pigme n t beco mes c o n

,

tr acted i n to sm all m asses t hus allowi n g the ligh t to b e

,

reflecte d fro m t h e other p ig me n t c ells Th ese facts m ay

.

e asily be d emo n str ated by comp a ri n g th e ski n o f a d ark

frog with th a t o f a ligh t o ne whe n gre at differe n ces in the
,

chrom atophores will al m ost c ert ai nly be observed Al though .

the bl ack chro m atophores li e m ai n ly belo w th e gold e n c ells ,

their br an ch es cover th e l a tter to a gre a ter or less exte n t ,

a n d whe n th e bl ack pigm e n t is fully exp a n ded it cuts off ,

m uch o f t h e light which would oth erwise be r e fl ec ted fro m

them .

The golde n color th at app e ars in th e frog s ski n i s du e ’

directly t o t h e pigme n t in th e gold e n cells b u t th e gree n is

,

n o t produce d in so si m ple a m an n er Th ere i s n o gree n

.

pigm e n t in t h e frog s ski n a n d v arious expl an a tio n s h ave

’
,

bee n offered as to ho w this color com es to appe ar Th e .

subj ect h as bee n i n ves t ig ated by B riic k e H arless V o n Wit

, ,

t ich ,
E ber th B i ed erm ann an d E h rm ann e ach of whom
, , ,

dis agree s in cer t ai n p articul ars wi t h th e o t h ers B r ucke .

reg arded the gree n c o lor as a si mple i n terfere nc e p h en o m e

n o n c aused by t h e gr an ules of gu an i n bu t th at th e golde n
pigme n t is n ecess ary t o th e productio n of gre e n w as su b se
que nt ly sho w n by the fact t h at w he n t h e golde n pigm e n t is
dissolved ou t of th e cells the gree n color dis appe ars al t hough
‘
1 94 TH E B IOLO G Y O F TH E FRO G CH A P .

Th e g r ee n i s produ ced accordi n g to B ied erm an n w h e n

, ,

the bl ack chrom at ophores are exp an ded be n e ath the yellow .

The n m ost o f th e light is reflected from th e gr an ules Whe n .

t h e bl ack chro m atophores are co n tr a cted so th a t th e yellow

cells h ave a lighter b ackgrou nd ligh t m ay be refl ecte d from

,

o t h er ele m e n ts th an th e blu e gr an ules and a yello w or golde n

,

color m ay predo mi nate Th e r o le o f the co n tr actio n and

.

exp an sio n o f th e golde n pigm e n ts is n o t accu r ately deter

mi n ed . I t is prob able th a t th e gr ay o r gr ayi sh blu e color
which i s so m e tim es assumed m ay be produce d by t h e si mul
t an eo us co n tr actio n of bo t h t h e bl ack an d th e gol de n pig

me n ts si n ce frogs w ith th e bl ack p igme n t spo t s co n tr a cted

,

o ft e n exh ibit these hues wh e n the golde n pigme n t h as bee n

dissolved out in alcohol V o n Wi ttich fou n d in th e tree
.

frog th at a gr ay color w as associ a ted with t h e co n tr actio n

o f both ki n d s o f pi gm e n t I n th e ordi n ary color ch an ges
.

v ari atio n s in th e co n ce n tr a tio n o f the golde n pigm e n t ar e

m uch less i mport an t however th an th e ch an ges in th e bl ack
, ,

cells .

Th e color ch an ges in th e ski n are produced by n u merous

age n cies wh ic h ac t u po n th e pigm e n t cells eith er direc t ly o r

through the c e n tr al n e r vous system The ch r o m a tophores

.

of th e frog form a very delic ate and respo nsive system which
i s co n st an tly u n dergoi n g ch an ges in respo n se both to sti muli
from th e e nviro n m e n t an d th e v aryi n g i n tern al st ates of th e
an i m al .O n e o f th e most i mport an t of th e exter n al s t imuli
a ffec t i n g t h e ski n i s ligh t
. I t i s a well k n ow n fac t t h at frogs
-

ex p osed to a bri ght light becom e ligh t in color while i f ,

t h ey ar e kep t so me ti me in t h e d ark th e ski n tur n s much,

d arker Th ese ch an ges are much more pro nou n ced in tree
.

frogs ( H yl a ) th an in the speci es of Rana an d they bri n g ,

about an a d ap t at io n of t h e color of the an i m al t o th at of

it s e n viro n me n t w hich is oft e n very close Th e qu esti o n

.
ix TH E SKIN 19
5

w hether light affec t s th e chro m atophores d irectly or through

the c en tr al n ervous syste m h as received co n sider able atte n
tio n The l a tte r alter n ative w as espoused by L ister w h o
.
,

fou n d th at a bli n ded frog n o lo nger ch anges i ts colo r in

respo n se to ch an ges in t h e i n te n sity o f light L iste r s c o n
.
’

c lusio n h as bee n o n ly p arti ally co n fi r m ed by subsequ e n t

i nvestig at ors S t ei nac h fou n d th at i f both the n erves an d
.

bloo d vessels supplyi n g any portio n of t h e ski n were cut in

t w o there still re m ai n ed in th at p art a cert ai n c a p aci t y for
,

color ch ange in respo n se to light o f differe n t i nt e n si t ies .

Wh e n pi eces of d ark p aper were l aid over po r tio n s of the ski n

thus tre ated o r eve n upo n portio n s of ski n e n tirely removed
,

fro m th e body the are as covered were fou n d to be co n sid

,

e rab ly d arker th an those exposed to th e light S pecime n s

.

o f H yl a in whic h cert ai n p arts were sh aded whil e o t her p arts

were exposed to ligh t bec am e light colored in all except th e
sh aded are as This w as fou n d to occur both in n orm al frogs
.

a n d in fr ogs whose spi n al cord w as destroyed C olor ch an ges

.

were fou n d by Dut art re to t ak e pl ace more r apidly in n orm al

frogs th an in specim e n s which h ad bee n bli n ded bu t th e ,

s am e re actio ns occurred in both c ases There is n o doubt

.
,

therefore th at light bri n gs abou t color ch an ges both directly

,

an d through th e ce n tr al n ervous syste m .

The i n flu e n ce of th e n ervous system u po n the chrom ato

p h o r es is show n by the experim ents of sever al i n vestig a tors .

D estructio n o f the optic th al amus c au ses the ski n to become

mu ch d arke n ed ( S tei n er B i ed erm an n ) S timul atio n of the
, .

m edull a c auses the ski n to assu m e a lighter colo r The ski n .

of th e leg m ay be m ad e to tur n p ale through sti mul atio n o f

the sci atic n erve B i ederm ann h as show n th at color ch anges
.

m ay be brought abou t both through th e spi nal n erves an d

th e symp at hetic system I f the s p i n al n erves supp lyi n g the
.

leg be cut the ski n o f t h e leg m ay still respo n d to ch an ges in

,
1 96 TH E BIOLOGY OF TH E FRO G CH A P .

the ce n tral n ervous syste m provide d th e symp ath etic n erves

which accomp an y t h e blood vessels rem ai n u ni nj ured .

The co n di t io n of th e pigm e n t cells is profou n dly i n flu e n c ed

by ch an ges in th e circ ul atio n An arrest o f the blood flo w .

c auses a p ali ng o f t he ski n I f th e leg o f a d ark colored

.
-

frog be tightly lig atured arou n d th e k n ee th e p art below th e ,

li g at ure will soo n assu me a much ligh t er color Th e s a me .

resul t follo w s i f t h e blood v essels alo n e are ti ed an d i s ,

effected more quickly i f th e lig ature is m ade arou n d an

a rtery i n s t e ad o f a v ei n .

Raisi n g t h e te mper ature c auses t he pigm e n t of th e ski n

to co n tr act C old o n the oth er h an d c au ses the pigme n t
.
, ,

to exp an d and t he ski n to assu me a d ark colo r Th e d ark .

color o f wi nte r frogs is in p art at le as t th e effect o f cold ,

an d t h e ligh ter color o f sum mer frogs in p art th e r esul t o f a

higher tem p er a ture A d ark colored frog m ay re adily be

.
-

m ad e to tur n mu ch lighter i f it i s pl aced for sever al m in u te s

in w a t er of a te mper a t u re 2 7 C C h an ges o f te mp er ature
°
.

a ffec t t h e co n c e n t r a t io n of pigm e n t eve n in i sol a ted pieces

o f ski n .

V a rious ch emic al subst an ces affect th e chrom atophores ,

som e c ausi n g a c o n t r ac t io n oth ers an exp an sio n o f th e pig

,

m e n t C arbo n dioxide prod uces a d arke n i n g o f the ski n ;

.

c arbo n m o n oxid e o n t h e o ther h an d c au ses t h e ski n t o tur n

, ,

p ale C hlorofor m an d so m e o t her anaes th e t ics as well as

.

cert ai n irri t an t s su ch as cro t o n oil an d c an t h arides c au se

, ,

an exp an sio n o f t h e pigm e n t o n t h e p a r t s o f th e ski n to

which th ey are appli ed D ry n ess t e n ds to c aus e the ski n

.

to t ur n p al e while i mmersio n in w a ter p roduces the reverse

,

effect This h as be e n observed especi ally in R a n a fu sca

.

by B iederm an n an d in R ag ilis an d H yl a by W er n er
. .

B iederm an n h as sh o w n t h at color ch an ges ar e i n flue n ced

in a re m ark able w ay by co n t act s ti muli S peci me n s o f .
1 98 TH E B IOLOGY OF TH E FROG CH A P .

i n w ard C hloroform an d oth e r d epress an ts d ec r e ase the

.

r ate o f p ass age o f flui d fro m withou t a nd i n cre ase its r ate
of p ass age fro m withi n Th ese di ffere n ces in th e r ate of .

th e tr an sm issio n o f fluids in di ffere n t directio ns te n d to dis

appe ar aft er the ski n dies .

The a mou n t of flu id th at can b e forced th r ough the ski n

u n d e r pressure d epe nds also u po n the di rectio n of flo w .

C i m a fou n d th at as mu ch w ate r u n der a pressure o f 1 0 c m .

.

o f mercury would p ass th rough th e ski n o f t h e frog fro m

withi n outw ard in five m i n u tes as would p ass through in
the reverse directio n in thi r ty seve n m i n u tes -
.

O f the excretory fu n ctio n o f the ski n of th e frog p rac ti

c ally no thi n g is k n ow n .

RE F E RE NC E S

A sch er son . Ue b er die H au t dru se n der F ro sch e . A ch A n at u

r . .

P hys .
, 1 8 40 .

Be rt , P . V en i n cu t an é de la g re n o uil l e. C R S oc B
. . . i o l. T . 2,

1 885 .

B iede m ann , r is t W . Z ur H o lo gi e u nd P hysio lo g ic der S chle im s e

c tire on . S it z b d k A k W iss
. . . . . M at h .
-
n at . Cl .B d 9 4, A b t h 3 , 1 8 8 6 ,
, . .

Wi e n, 1 8 8 7; U b d F b e er en ar enw e chs el de r F r o sc h e A r g es .
, ch . . P hys .

B d 5 1 , 1 8 92
. .

B ou l en g er, G A . . The P o is ono us Se c ti

re on of B at ra ch i s an . Na t .

S ci , V o l
. . 1 , 1 892 .

Don aldson , H H . . O n t he A bs or
pt i on of W at e r by F ro g s . S ci c en e,

n s . .
, V ol
3 , 1 90 1 .
’

1 .

Dr asch , 0 B e o . b ch t u g
a n en an l b d
e en en D us en r , e t c. A ch An r . at .

u P hys
. .
, p hys . A b th .
, 1 889 .

Dut art re, A . S u r le s ch g m an e en t s de c u l u ch z l g ui ll

o e r e a re n o e

c m mu
o ne ( Ra n a escu len t a
) . C R H. . e bd m A S i T 3 8 9
o . c . c .
, .
,
1 0 .

E h m a nn , S r . Z ur Ph ysio lo g ic de r P igm t z llen C t f P hys e en . en . . .

,

B d 5, 1 89 1
. .

E n gel m ann , T . W . Die H au t drusen des F o sch s r e . A ch r .

g es .

P hys .
, B d 5, . 1 8 72 .
rx TH E SKIN 1 99

G adow , H . Co lo r in A m ph ibi a . P c ro . Roy I n t . s . G r eat B r t a i i n,

1 90 2 .

H a less, E r . Ub e er die Ch ro m a t op h o re n des F o sch es Z e i t

r . . w is
s .

Z ool B d 5 , 1 8 54
.
, . .

H eiden h ain , M . Die H au t dru sen der A m p h ibi en .

”
S it z b W u rz b . .

p hys m ed G es 1 8 93
-
. . .
, .

H ub e r , O . Ub e er B r un st w arz en b ei Ra n a t emp or a r ia . Z ei t . w iss .

Z l oo . B d 45 ,
, . 1 88 7 .

J u n iu s, P . Ub e er die H au t dru se n des F sch es Arch mi k An at

ro . . . .
,

B d 4 7, 1 8 9 6
. .

L ey dig , F . Ub e er die a l lg m ie e n en Be d ck u
e n gen hibi
der A mp en .

A ch r . m ik A n at . .
, Bd . 1 2, 18 76 ; Die an u re n B at rac hi d d u t sch
er er e en

F un a a, 1 8 77 ; I nt e gum ent b ru nst ig er F isch e un d A m p h ibi Bi en . ol .

Cen t .
, Bd . 1 2, 1 892 .

Ub e er das B l u i d F b d Th i Z l A
a n er ar e er er e , oo . nz .
, B d 8, . 1 885 ;

B lau farb ig e r W ss f sch Z l G t

a Bd 33 89
er ro . oo . ar en , .
, 1 2 .

O v ert on .
3 9 Th s iib di W dk mi d e en er e asse r ono e er A m p h ibi e n , et c .

V e rh p . hys
m ed G es W iirz b u rg , B d 3 6
.
-
. . . .

P f i z ne , W t
Die E p er r de r A p . id mis m h ibi en . M o rp h .
J ah rb .
, Bd .

Reid, W . O sm sis E x p im e n t s i t h L i vi n g
o er w an d D ea d Me mb ra n e s .

Reid an d H am b l y . T spi ti
O n th e f C b ran ra on o ar on D i xid e
o

t h ugh
ro th e i
S k n o f t he F g J u P hys V l 8 8 9 5
ro . o r . .
, o . 1 ,
1 .

S eeck , O . Ub e er die H td i ig A m p h ibi

au I
ru se n e n er en . n an g D iss
. .

I) o rp a n 1 89 1 .

S t einachUb F b chs l b i i d
, E W i b lt hi
. e er ar en w e e e n e ere n r e ere n

b di gt du ch di c t W i k u g d L ich t s f di P igm t z l l
e n r re e r n es e au e en e en .

C t f P hys B d 5
en . . 89 .
, .
, 1 1 .

S ti d L U b d B d H u t d F sch s A ch A t u
e a, . e er en au er a es ro e . r . na . .

P hys 8 6 5 .
, 1 .

S t ir l i g W O th
n E xt t t , h ich Abs pt i .t k n e en o w or on c an a e

P l c t h ugh t h S k i f t h F g J u A t d P hys V l
a e ro e n o e ro . o r . na . an .
, o . 1 1 ,

1 8 77 .

S t r ick er u nd S p ina U . n t er such u g ub di m ch isch

n en er e e an en Le is
t u g
n en de r a c in o se n D us r en . S it b A k W iss M t h
z . t Cl . . a .
-
na . .
, B d 80 . .

Ab th 3 , . 1 8 79 , W i en , 1 880 .

T ow n son , R . O b se rvat io n es p h ysio lo gicae de A m ph ib iis, G o t t in gae,

1 79 5
200 TH E BIOLOGY OF TH E FROG CH A P .

W er ner , F U b e er ut f b
die V e ra n deru n g de r H a ar e b ei e uro
p ais
V b d k k z l—
.

ch B t
en a rac h ie rn . er . b t G. Wi . . oo . o . es . en, B d 40 ,
. 1 8 90 ,
A lbi ismus
n un d Me l ismus b i R p t ili
an d A m p hibi
e e en u n en . I bid .
, Bd .

W itt ich W , g u F b d H ut u s F h i h
. v on . Die r ne ar e er a n er e r ro sc e, re

p hysi l gisch
o o d t
pa h l gisch
e un V a d g A ch A t u P hys
o o e er n er un en . r . na . . .
,

1 8 5 4 ; E tg g u g
n f H
e H nl ss
n
“U b di Ch m t ph
au e rr ar e
’
e er e ro a o o re n

l s F sch s I bid 8 5 4
ce ro e .
”
.
, 1 .
20 2 TH E BIOLOGY OF TH E FROG CH A P .

S t r uct ure an d — Th e kid neys

F u nct ion of t he Kidn eys
o f th e frog are ov al fl a tte n ed d ark red bodi es lyi n g dors al
, ,

t o t h e p e ri t o n e u m o f th e posterio r porti o n of th e body c avi t y .

The duct o f t h e kid n ey ,

o r u r eter i s j oi n ed at ,

abou t th e posterio r third

or fou rth of th e ou t er
m argi n ; i t th e n ru n s for
a shor t dist an ce alo ng
t h e do r s al surfac e an d
fi n ally becom es embed ded
in t he subst an c e of th e
kid n ey ru nni n g n e ar th e ,

m argi n to th e an terio r
e n d of th a t o rg an Th e .

ve n tr al surfac e of the kid

n ey is fl atte r th an th e
dors al an d i s tr averse d
lo ngi tudi nally by th e yel
lowish a dr en a l body Th e .

kid n eys are cov e red by

p erito n eu m o n ly o n th e
ve n tr al surface with t h e
F I G 5 —M l u i g i t l g s exceptio n of a very s hort
. 2 . a e r no en a or an .

A A t ; Cl cl
0, or a c ; C p stc l ,
oa a v, o ava

ve ni ; F K f t b di s ; H t s t s ; sp ace wh ere thi s me m

'

a o e o, e e

U u t s p i g i t t h cl c
,

r, re e r O t br an e is folded in over
en n n o e oa a a

S S ; V lv i s ( A ft W i d
’
r re n a e n er e er
th e edges
.
, ,

h im )
.

s e .

The kid n ey m ay be
reg ard ed as a co mpou nd tubul ar gl and m ad e up o f a , ,

l arge n umbe r o f coil ed u r inifer o u s t u bu les E ach u rin if .

ero u s tubule begi n s in a M a lpigh ia n b od y n e a r t h e ve n

t r al surfac e A M alpighi an body c o n sists o f t w o p ar t s

.
-

a k n o t o f blood vessels the g lo mer u lu s an d a sur , ,

 TH E E X CR E T ORY SYS T E M
r ou ndi n g m embr an e or , pB ow Th
ma ne ’
a rtery
s ca s u le .
,

vas afi er en s
’
e n teri n g t h e c apsule bre aks up i n to sever al
,

c apill aries whi ch a ft er fo r mi n g a few coils e m erge as th e

, ,

effere n t blood vessel fro m the s am e op e n i ng B owm an s .

’

c apsule is an exceedi n gly thi n me mbr an e the r e i s an i nn e r

fold closely applied to t he glom erulus which is co n ti nuous
with th e ou ter w all at th e poi n t wh ere th e bloo d vessels e n ter
th e c apsul e B owm an s c apsul e i s
.
’

si mply the thi nn ed out an d exp an ded

e n d o f a u ri n iferous tubul e which h as

beco m e push ed by th e glom e rulus as

o n e might push in t he e n d of a fi nger

of a glove . Th e c apsule howev er , ,

h as g r ow n arou n d th e glom erulus a n d

closely surrou n ds th e affere n t an d effe
re n t vessels At th e dors al sid e o f the
.

c apsul e and usu ally opposi te t h e poi n t

,

wh ere th e blood vessels e n te r t h e outer F I G ,

53 A n f . . u ri i er
w all p asses i n to the n eck of t he u rin if t b i a l ous u u e , co

f a iigg h n
FUb UI
e ro u s tubule Th e very th i n cells of d n C tfl

t; d’
.
.

E if
a ra 0
th is w all sh ad e off gradu ally in to cells 1 “ t b j ,
n e ro u s u u e

of colu m n ar epitheliu m which for a l di n g f m th l t ea ro e a

s h or t distan ce c arry very l arge c i1ia

t t th c ll .c ti g . er o e o e n

t ubul ( Aft N
.

e . er u ss

B eyo n d th e n eck wh ich is so m ewh at b um )

.

a .
,

n arrower th an t h e rest of th e t ubule th e c ells are li n ed wi t h ,

m uch shorter cili a E ach tubule i s li n ed with a si n gle l ayer

.

o f cells wh ich v arie s in ch ar acter in the di ffere n t p arts .

The course o f e ach tubul e is quite c omplic a ted At firs t it .

r u ns dors ally whe r e i t forms a more or l ess complic ated coil

, ,

the n i t proceeds to th e ve n tr al side o f the kid n ey forms a ,

seco n d coil an d fi nally ru n s dors ally ag ai n e mptyi n g in to

, ,

o n e of th e collecti n g c an als which exte n d tr an sversely a cross

the do r s al su r face of the kid n ey fro m the i nn e r m argi n to

204 TH E BIOLOG Y OF TH E FRO G CH AP .

th e u r eter Th e tubules are h eld toge t her by co nn ective

.

tissu e which fo rms a support also for the n u m erous blood

vessels with which the kid
n ey is suppli ed .

Th e ve n tr al surfac e of
the kid n ey is fur n ished
with n u m erous cili ated
fu nn els th e nep h r os to mes , ,

whose e xp an ded e n ds o pe n
i nto th e coelom A t t heir .

other e n d th e n ephrostom es
emp ty i n to br an ch es of th e
re n al vei n s an d th e cili a ,

with which they are li n ed

be at to w ard the u pper e n d
o f these o rg an s and thus
cre ate a curre n t o f lymph
fro m th e body c avity i n to
th e blood Thi s r el atio n .

o f th e n eph r osto m es is a
FIG 54. D i g m f k idn y sh
a ra o p eculi ar o n e and o ccurs
a e ow

i g th u t d c ll c t i g t ubul s
.

n e re e r a n o
o n ly in t h e A n ura Th e
e n e .

C c ll c t i g t ubul s ; L l
.

o e n g it di
e on u
lo er Amphibi preserve
, ,

na l c l f B idd ; S s mi l
an a o er
w
,
a e na

v sicl ;
e T t s t is ; U u t
e , e ; VE th e typic al arran gem e n t o f
, re e r ,

v s ff t i these o rg an s as the n eph r o

a a e er e n a .

st o m e s are co n n ected wi th the re n a l tubules This co n di .

tio n as M arsh all has fou n d occu rs also in th e e arly st age s

, ,

of t h e li fe of t h e frog bu t l ater t h e n ephrostomes los e th eir

,

origi n al co nn ec t io n wi th t h e t ubul es an d becom e u n ited

seco n d arily wi t h the re n al vei n s .

The kid n ey of t h e m ale frog st an ds in an i n t i m ate rel atio n

to the sexu al org an s The va sa efi er en tia or ducts which
.
,

co nvey the sp er m ato z o a from t h e tes tis p ass i nto the sub ,
 20 6 TH E BIOLO G Y OF TH E FROG CH A P .

n ec t ed wi th th e collecti n g tubul es wh ich exte n d across the

dors al sid e o f th e kid n ey to th e u reter Th e sho r t tubes .

wh ich co nn ect the lo n gi t udi nal c an al with th e collecti n g

tubules wide n o u t n e ar th e l a tter to form an am pull a Thi s .

e n l arge me n t is form ed by a M alpighi an body wh ich h as lost

its glom e r ulus an d co n seque n tly it s origi nal fu n ctio n I n .

Ra n a f usca there is a co mp ar a
t ive ly direct co nn ectio n es t ab

lish ed betwe e n th e v as a effe

r en t ia an d th e collecti n g tubules ,

an d the sperm a tozo a there fore , ,

are n o t fou n d in th e M alpighi an

u bodi es an d fu n ctio nal r e nal tu ‘

s e.

FI 5° D g m t ”I m t
th c u se f t h sp m t z
o r e o
la
bules B idd er s c a n al occu r s in
ra

e
o

er
n

a o
a e

oa
.
’

th ugh t h kid y f R
ro t h e kid n eys o f both sexes but
e ne o ana ,

f m mp fl i
.

in g
’

t ubul
‘t
l
l B i dd s l gi
e
i ts fu n ctio n in t h e fem ale is n o t
a ‘ a

er
C

on
; ,

t u din a l c an a l; t, u i if us
r n ero k no w n

t ubul e : 21 . u
m any of th e lower verte
re t e r : W »
va s
In
( M d fi d f om
$1212)
O l e r
br a t es ( E l a smobr a n ch s A m ,

p h ib ia ) t h e kid n ey is d ivided ‘

i n to an an terio r or sexu al portio n an d a posterior or ex , , ,

c re t o ry portio n Th e frog prese n ts o n ly the begi nn i ng of

.

such a di ffere n ti atio n Th e v as a effere n ti a are co n n ec ted .

with the an terior p art of the kid n ey but th e excre t ory fu n c ,

tio n of this regio n is still ret ai n ed The course o f th e sper .

m at o z o a t h r ough the kid n ey v aries c o n sid erably in di ffere n t

species of frogs as i s evi n ced by th e fact th at it is much ,

more direct in Ra n a f usca t h an in R escu len ta Th e l a tter . .

prese n ts doubtless th e more primi t ive co n di t io n

, ,
.

The kid ney is supplied wi th blood from two differe n t

sources ( I ) th e ren a l a r ter ies which rise fro m t h e uri n o ,

ge n i t al arteries o r direc t fro m th e aort a and ( 2) the r en a l

, ,

o r ta l veins whic h co n vey ve n ous blood from t he poste r io r

p ,
 TH E E XCRE TO RY S Y ST E M 20 7

portio n of t h e body Th e re n al arteri es of which there are

.
,

u su ally fro m four t o six e n ter the kid n ey at the medi an edge
,

or n e ar t h e l atter o n the ve n tr al surface Th e divisio n s .

of the r e nal arteri es are distribu ted to th e re n al tubule s

a rteri ae rec t ae ) a n d also to th e glom eruli v a s a affe r en t ia
( , ( ) .

The re n al p ort al vei n ru n s alo n g the dors al surfac e o f th e ‘

kid n ey very n e a r th e ou ter m argi n F rom the tr an sverse .

br an ches o f this vei n which exte n d a cross th e dors al surface

, ,

sm all vessels are give n off w h ich p e n etr at e the subst an ce of

the kid n ey an d form c apill ary n etworks arou n d th e r e n al
tubules Th e v as a effere n ti a which em erge fro m the glo
.
,

m eruli together wi th th e effere n t vei n s arisi n g fro m this c ap il

,

l ary n etwork go to form the begi nn i n gs o f the re n al vei ns

,

which co n vey the blood fro m t he kid n ey to th e posterior

ve na c ava Th e glo meruli are supplied o n ly with arteri al
.

blood while the re n al tubules receive blood fr o m the re n al

,

por t al vei n s an d also al t hough to a less exte n t fro m the

, , ,

re n al arteri es .

The fu nctio n of th e kid n ey is the eli mi natio n o f w aste

m a tters from th e blood Th e re n al excre t io n o r uri n e is a
.
, ,

fluid co nt ai n i n g a l arge n u mber o f compou n ds in solu tio n .

M o s t o f the n itroge n l e aves th e body in th e form of ure a ,

( N H 2) 2C O wh ich
,
i s a whi t e cryst alli n e compou n d very solu ,

ble in w ater U re a r eprese n ts the fi n al product o f the bre ak

'

ing dow n of th e n i troge n ous subst an ces of the body an d it ,

h as bee n sh ow n t h at t h e form at io n of this subst an ce t akes

pl ac e to a l arge exte n t in the liver fro m whi ch i t is give n to ,

the blood by a proc ess of in te rnal secretio n Th e kid n ey .

also excretes seve r al s alts such as th e chlorides sulph ates , ,

an d phosph a tes o f sodium pot assiu m c alcium an d m ag n e , , ,

sium and n u m erous o ther subst an c es in sm aller propor t io n s

, .

The specifi c r o les of th e glo m eruli an d tubules in re nal

excretio n h as lo n g bee n a m atter of dispute I t i s cer t ai n .
20 8 TH E BIOLO G Y OF TH E FROG CH A P .

th a t water and other subst an ces diffuse fr o m th e blood

t hrough th e w alls o f th e c apill aries o f the glo meruli i n to

t h e re nal tubules I t has bee n held especi ally by L udwig

.
,

an d his follo w ers th a t pr actic ally all of th e subst an ces excreted

, ,

by th e kid n ey p ass t hrough th e glomeruli an d t h at th e fu n c

,

t io n o f th e tubules is to absorb the excess o f w a ter an d

cert ai n other m ateri als which p ass dow n th e lum e n B y

-

o t her physiologis t s it h as be e n m ai n t ai n e d th at bo t h the

glomeruli and the re n al t ubul es are secre t ory but th at th ey,

el imi nate diffe r e n t produ cts Nussb au m s i nge n ious exp eri
.
’

m e n ts o n the fro g see med to offer a solu tio n of this problem .

As the glom eruli are suppli ed by bran che s o f th e re n al

a rteries Nussb au m co n clude d th at th e blood su pply o f t hese
,

organ s would b e c u t off if th e re nal arteries were t i ed Th e .

o ppor t u ni ty w as thu s prese n ted o f c omp ari ng th e excr etio n

of th e kid n ey in wh ich the glo meruli are r e n d ered fu n ctio n
less with th a t o f the n orm al org an I t w as fou n d th at in
.

frogs with th e re n al arterie s t ied th e s ecre t io n of u ri n e w as

much di mi n ish ed in a mou n t S olutio n s of sug ar pepto n es
.
, ,

an d egg albu me n which wh e n i nj ec ted i n to the blood o f

,
.

n orm al fro gs soo n m ake th eir a ppe a r an ce in th e uri n e could ,

n o t be detected a fter i nj ecti o n i n to th e blood in th e u ri n e

, ,

of frogs whose re n al arteri es were lig atured eve n after the

,

flow o f uri n e w as i n cre ased by the si mult an eou s i nj ectio n o f

ure a Nussb au m c am e to th e co n clu si o n th at album e n
.
,

sug ar and most s alts are excreted by th e glom eruli while

, ,

ure a is elimi n ated by the cells o f the uri ni ferous tubules .

Th ere i s a sourc e of erro r in s uch expe r i me n t s si nce lig at ,

ing t h e re n al arteries al o n e d oes n o t e n tirely cu t off th e

blood supply of t h e glomeruli ; t h ere are an astomoses wi th
t h e ge n it al arteries by m e an s o f which these org an s m a
y
receive blood in a so m e w h at rou nd abou t w ay A dami .

fou nd th a t some of th e glom eruli bec ame fill ed by i nj ecti n g

21 0 TH E BIOLO G Y OF TH E FROG CH A P .

r esti ng u po n a m e mbran e of co nn ective tissue Nu merou s .

goble t cells occur amo ng th e o ther epi t h eli al cells The .

middle l ayer of th e bl add er co nsists o f a n etwork of smoo th

muscle fibers Th e fibers are sometim es si ngle and so m e
.
,

t i mes u n i ted i n to bu n dles an d they exte n d in all di r ectio n s

, .

O utsid e o f the muscle l aye r is a thi n sh ee t o f co nn ec tive

t issue which is covered exter n ally by th e pe r i to n eu m .

Th e bl adder is very diste n sible as m ay re adily be show n ,

in a rece n tly killed frog by i n fla ti n g it by m e an s of a blow

,

pipe i nt r oduced i n to th e clo ac a Wh e n e n tire ly empty the .

,

bl adde r shri nks to a n i n co n spicuous size I t w as formerly .

doubted wh ethe r the bl adde r of the frog serves as a recep

t acl e fo r u ri n e as i t has no direct co nn ectio n wi th th e ducts
,

fro m t he kid n eys To w n so n whose co n clusi o n s were fol

.
,

lowed by Dum eril in hi s gre at work o n reptiles an d amph ibi a ,

reg arded the bl adder as a sort of reservoir for w ater absorbed

t hrough th e ski n Th e co n te n t s of the bl adder were st ated
.

t o be n e ar ly pure w ater an d the u r i n e proper w as supposed

,

to p ass ou t and through the clo ac a with out e n teri n g th e

bl adder at all Accordi n g t o Du m eril
.
l
la pr é te n due ,

vessi e uri n aire des G re n ouilles d es Rai n e ttes e t des Cra ,

p ands ai nsi que celle d es S al a m an dres est u n e sorte de

, ,

ci ter ne ou u n e hu meur a queus e presqu e pure des t i né e a

‘

, ,

l exh alat io n cut ané e se mbl e etre ap p o rt ie soit p ar les vei n s

’

, ,

s angui n es soi t p ar les lymp h atiqu es

, S ubsequ e n t i n vestig a .
”

tio n s by D avy Nussb au m and Ad ami h ave sho w n th at t here

, ,

is n o doub t th at th e fluid co n t ai n ed in the bl adde r i s derived

fro m the kid n eys si nce it co nt ai n s ure a an d other sub
,

s t an ces c h ar acteristic o f re n al excre tio n .

The e nd of the clo ac a i s co m mo n ly h eld closed by th e

co n t ractio n o f its ci rcul ar muscles an d t h e u ri n e which i s ,

thus preve n t ed fro m p assi n g o u t collects in t h e bl adder .

1 Du m e ril , E rp é t o l gi
o e g é n é i a le .
 TH E E XCRE T O RY S YS TE M 21 1

The co n te n ts of the bl adder are expelled sudde nly by the

co n tractio n of th e muscles of the body w all which n at ur ally ,

subj ects the bl adder to a co n siderabl e pressure The ex .

pulsio n of uri n e ofte n t ake s pl ac e whe n t he frog le aps and ,

it is very ap t to o ccur as a co n sequ e n ce o f the struggles o f

th e an im al if th e fro g is t ake n in the h an ds as every o n e who ,

h as h an dled frogs has doubtl ess discovered The belief th at .

the co n te n t o f th e bl adde r o f the to ad i s poiso n ou s is

e n tirely withou t fou n d atio n .

RE F ERE NC E S

J
Adam i, . G . O n th e Nat u re of th e G lo m u l us A c t i i t y
er v in t h e
K id y J u P hys
ne . o r . .
,
V ol 6, . 1 885 .

B eiss n er, H . De r B au der sa m e n ab le it e n de n W eg e b ei Ra n a f u sca

und Ra n a escu len t a . A ch mi k
r . . A n at .
,
B d 5 3 , 1 8 98
. .

Fa rr ing t on , 0 C . . T he Nep h ro s t o m es o f Ra n a . T ran s . Co n n . Ac .

S ci , V o l 8 ,
. . 1 8 92 .

F ra n k l , 0 . Die A u s fu h r w e g e der H arn sam e n n iere cle s F sch s ro e .

Z ei t . w iss Z . o ol .
, B d 6 3, . 1 897 . S ee s
al o Ar ch mi k . . A n at .
, Bd 5 . 1 , 1 8 98 .

Nu ssb au m U b di S c t i d ,
A ch g
M .P hys e er e e re on er Ni e re . r . es . .
,

Bd .6 8 78 F t g s t z t U t such u g
1 , 1 t . l Bd 7 or 8 78 e e e n er n en, e c . . e .
, . 1 , 1 .

U b di E t ic k lu g d
e er e m a b l it dn wW g b id A u e n er sa en e en en e e e en n re n .

Z l A
oo Bd 3
. 88 nz U b di E digu g d W im p t i h t i
.
, .
, 1 0 . e er e n n er er r c er n

d Ni
er d A u l
ere Bd 3 88 U b d B
er d di T h at ig
n re n . . e
.
, .
, 1 0 . e er en au u n e

k it d
e D ii A ch mi k A
er t
r Bd 7 886
se n . S ls A t A r . . na .
, . 2 , 1 . ee a o na . nz .
,

Bd . Z l A1 Bd ; d A ch mi k A
oo . t Bd 5 nz .
, . 20 an r . . na .
, . 1 .
21 2 TH E B IOLO G Y O F TH E F RO G CH A P .

C H A PT E R XI

TH E RE P ROD U CT I V E O RG ANS A ND T H E FAT B OD I E S

TH E reprod uctive syste m has th e fu n cti o n s of produci n g

t h e sex cells a nd tr an sporti ng the m outside of th e body .

Th e fi rst fu nctio n is disch arged by the go n ad s which are ,

k no wn in the fe m al e as ova r ies and in th e m al e as z es les

,
‘

or sp er ma r ies While th e ov aries an d testes are homologous

.

org an s th e sexu al prod ucts are c arried to th e outside in the

,

t w o sexes by very di ffere n t m ethods .

O r gan s of t h e F em ale E ach ov ary o f t h e frog is in the

.

fo r m of a sac whic h is more o r less lobul ated I t s i n t er nal .

c avity i s divide d by s everal p arti t io n s i n t o ch a mbers which

are filled by fluid E xter n ally the ov ary is covered by
.
,

peri to n eum which i s co n ti n u ed o n the dors al side to form

,

a doubl e m embr an e th e nzesova r ia in which suspe n ds t he

, ,

o v ary fro m th e do rs al body w all Th e blood v essels and

.

n erves which su pply the ov ar y r u n betwee n the t w o m em

bran e s of this su pporti ng str u cture Th e i nn er surface o f .

t h e ov ary is li n ed by a si ngle l ay e r of fl atte n ed epi t h eli al

cells th e origi n o f whi c h m ay be traced to o u tgrowth s fr o m

,

t h e kid n ey in e arly develop me n t Th e s lr a la nz m ediu m

.
,

or middle portio n of the w all of t h e ov ary v aries gre atly in ,

thick ness in di ffere n t p arts an d at differe n t ti m es I t i s .

composed m ai nly of ova an d follicle cells in v arious s t ages

o f d evelopm e nt Th e eggs lie withi n sm all ch ambe r s o r
.

follicles the se co n sist of a l ayer of cells ( me mbran a grau n

los a) lyi n g n ext to t he vi t elli ne m embr an e an d outsid e of ,

this a v ery v ascul ar n etwo r k th e f/z eea f ellien li Afte r th e

,
.
21 4 TH E B IOLO G Y OF TH E FRO G CH A P .

th e body c avi ty n e ar the b ase of th e lu n g At the poste r ior .

e n d it e n l a rges to form the thi n w alle d ve ry dist e n sible u ter us

-

, ,

the op e n i n gs of the t w o u teri lie close toge t h er o n the dors al

w all of th e clo ac a With t h e excep t io n o f the u terus an d
.
,

a short sp ace at th e an terio r e n d t h e oviduc ts possess a thick ,

gl an dul ar w all The i nn er surfac e o f th e oviduct i s throw n

.

i n t o lo ngi tudi n al ridges which are covered with cili ated

,

epi t h elium The grooves be t w e e n the ridges receive th e

.

op e n i ng s o f the n u merou s gl an d s wh ich secrete th e gel at i

n ou s co a ts o f th e eggs Th ese gl an ds are mostly of the
.

simple tubul ar type they are li n ed by a si ngle l ayer o f

cyli n dric al secreti n g cells which beco m e very much e n l arged
duri n g the b r e edi n g se aso n Whe n t h e secre t io n i s dis
.

ch arged th e outer m embran e o f t h e cells i s burst ( L ebru n)

, ,

a n d t h e co n te n ts w hich form ed a gre a ter p art o f the bulk

,

of the cell flow i n to th e lu me n o f th e gl an d

,
A fte r th e .

disch arge o f the secre t io n t h e gl an ds becom e very m uch

reduced in size an d th e whol e o viduct much t hi nn er an d of
, ,

a yellowish color from th e accu mul atio n o f fat As B o e tt .

cher h as sho w n th e oviducts in t h e breedi n g pe r iod possess

,

a rem ark a ble c ap a city for t h e absorptio n o f w a ter A p air .

o f ovidu cts whic h whe n j ust t ake n out of th e body weigh ed

,

g
. were
,
fou n d to weigh 1 0 8 4 g aft er they h ad l ai n so m e.

time in w ater i e th ey had i n cre ased in weigh t 1 1 3 tim es

. . .

Aft er the breedi n g s ea so n t his power o f absorbi n g w ater is

very much r educed The eggs as they are disch arged from
.
,

th e ov aries are t ak e n i n to th e mou t hs of t h e o viducts by m e an s

,

o f cili ary ac t io n Th ey are the n c arri ed dow n th e oviducts by

.

me an s of t h e cili a o n t h e ridges o f th e i n n er w alls D uri n g .

t h is p ass age t h ey receive t heir co ats o f j elly after which t h ey ,

collect in the u t eri whose w alls t hey gre a tly diste n d H ere
,
.

t h ey m ay rem ai n for sev er al d ays th e l e n g t h o f t im e depe n d

,

ing upo n th e prese n ce o r abs en ce of t h e m ale ( see p .

x1 RE P ROD UCT I V E O RG ANS A ND T H E F AT B OD I E S 21
5

Th e m ales of several species of Ran a possess a cu r ious

ho mologue o f the ovi duct o f th e fem ale I n R a n a p ip ien s
.

i t is very well developed and co n t ai ns an e n l argem e n t at its

,

po sterio r en d represe n ti ng a u terus I t lies j u st extern al to

.

the ureter and exte n ds as a fi n e tube som e dist an c e in fro n t

,

o f th e kid n ey I t s fun ctio n i f i t h as any is u n k n ow n C ases

.
, , .

ar e n o t u n com mo n in which org an s ch ar a cteristic of o n e sex

are fou n d in a r ud im e n t ary form in the oth er an d it is n o t ,

i mprob able th at the oviduct o f th e m ale frog is S im ply a use

less al t hough r ath er l arge rudim e n t o f this ki n d I n th e
, , .

bullfrog ( R a n a Ca les é ia n a ) thi s duc t is abse n t .

Or gan s of t he M ale —Th e tes tes are rou n ded or ovoid

.

org an s lyi ng ve n tr al to the kid n eys L ike th e ov aries they

.
,

are sur r ou n ded by perito n eu m which is exte n ded dors ally

,

a s a double me mbr an e th e mes or enin in to th e dors al sid e

, ,

o f t h e body c avi t y where it beco mes co n ti nuous with th e

,

ge n eral coelomi c li n i ng Th e ra w efi er en lia o r ducts o f

.
,

the testes co n sist of a v ari able n u mber of sl e n d er tubes

, ,

whic h ex t e n d wi t hi n th e mesorchiu m t o th e i nn er m argi n

of t h e kid n ey wh ere t hey co n n ec t wi t h B idd er s c anal Th e
,
’
.

v as a effere n ti a o fte n br an c h an d anastom ose m ore or less ,

in a w ay wh ich v a ri es gre atly in differe n t i n dividu als .

T he testis is m ade up esse n t i ally of a m ass of tubules ,

toge t h er wi t h blood vessels and n erves an d a sm all amou n t

,

of co nn ective tissu e bi n di n g th e tubules together The whole .

i s surrou n d ed by a c o n n ec t ive tissu e m embr an e th e t u n ica ,

a ll nzg in ea ou tside of which is t h e perito n eum

,
Tow ar d th e.

ou ter porti o n o f th e testis the tubules exte n d r adi ally an d ,

e n d bli n dly n ext to th e tu n i c a albugi n e a Ne ar th e poi n t

.

wh ere th e v as a effere n ti a e n ter they becom e co iled irreg u

larly . The v as a effere nt i a form a n etwork w i t hi n t he tes t is ,

i n to which t h e t ubules op e n at th eir i n n er e n ds E a ch .

tubule pos sesses a n o u te r nz ein é r a na p r op ria and an i nner

21 6 TH E BIOLO G Y OF TH E F RO G CH A P.

li ni n g o f cells som e of whic h (sp er nz a log on ia sp er m a zoey les

, ,
‘

a n d sp er nz a l irls repres e n t s t ages in t h e form a tio n o f sp erm a

)
t o z o a ; o t h ers form t h e so c all ed
“ follicle cell s an d t h e
- -
”
,

fl at t e n ed cells d escribe d by B ert ac c hini whi ch li e n ext to ,

FI G .
59 . A, c ss s c i
ro e ubul s f th t t is ; f bu dl s f
t o n o f o n e o f th e t e o e es s ) n e o

m t z p i h li l li i g f h t ubul B s g s i th d
,

sp e r a o oa ; L e, e t e a n n o t e e .
,
ta e n e e

v e lo p m en t o f sp er m t z
a o oa ( Af
. P k d P k
t er ar) er an ar er .

the ou ter m embr an e The fol licle c ells form a sor t o f w all
.

a rou n d groups of cells fro m wh ich th e sp erm at ozo a t ake

t hei r origi n .

Th e sperm atozo a o f th e frog p ass th rough the subst an ce

of t h e kid n ey i nt o th e u reter I n m an y speci es o f frogs the .

free p or t io n o f t h e u re ter i s dil ated to form a semi nal recep

t acle in whi ch t h e sp e r m at o zo a are st o red ag ai n st t h e ti m e
o f their d isch arge fro m t he body Th e semi n al recept acle .

is p oorly de v e lo p ed in R a n a p ip ien s an d R Ca ies bia na I n . .

the E urope an speci es R f usea it becomes very l arge an d .

divid ed i n t o a n u mber o f comp artme nt s .

C o rr espo n di ng to th e v arious st ages in t he d evelop m e n t

21 8 TH E BIOLOGY OF TH E FROG CH A P .

Aft er th e feedi n g p eriod begi n s the re i s a r apid stor age of a

yellowish fat in th e cells whi c h beco m e gre a tly i n cre ase d in ,

siz e .

Th e developme n t o f th e fat body is closely co nn ected

with th at of t h e go n ads B oth in fact arise fro m t he .
, ,

differe n ti atio n of th e ge n it al ri dge th e an terior portio n o f ,

which fo r ms the fat body the poste r ior po r tio n the ov ar y o r ,

testis .

RE F E RE NC E S

B oet U b d B
t ch er d di Q ll g f h ig k it d
, A . e er en au un e ue un s a e er

F ro sch il it V i ch s A ch i B d 3 7 8 6 6
e e er . r ow
’
r v, .
, 1 .

B o in M H is t g
u es d l G l, d G i t l F m l l ch z R
. o en e e a an e en a e e e e e an a

t mp
e i A ch d B i l T
or a r 7 9 a . r . e o .
, . 1 , 1 01 .

F nk e Ru U b di S ch k u g d F ttg h l t s d F t t f h
, . e er e w an n en es e e a e er e u

r end O g im K isl uf d J h s D k k A W iss M t h

en r an e re a es a re . en s r. c . . a .

na t Cl B d 6 8
.
9 Wi
.
,
.
, 1 00 , en .

G igl io T o E S l c p s g s d A m p h ibi s A ch I t l d
-
s, . ur es or ra es e . r . a . .

Bi l T
o .
5 8
, 9 6 . 2 ,
1 .

P l oe t A J Di V g g i d Fz ,
hh d u t d m Ei fl
. . e or an e n en ro sc o en n er e n u ss

d J h sz it
er A ch A
a t u P hys p hys A b t h S u p p l B d
re e .
89 r . na . . .
,
. . . .
,
1 0 .

T a ch anoff J Rr Z P hysi l gi d G h l ht pp t d , . . ur o o e es esc ec a a ra u s es

F sch s A ch g P hy B d 4 8 8 7
ro e . r . es . s .
, . 0, 1 .
xi1 T
I N E RN L A S E CRE TI O N A ND D U CT L E S S G L AND S 21 9

CH A PT E R XI I

T
I N E RN L A S E CRE T I O N A ND TH E D U CT L E S S G L AND S

TH E ide a of i n ter nal secre tio n w as first brought i n to

p r omi n e n ce by B row n S equ ard who fou nd th at extracts of
-

the testis o f m am m als wh e n i nj ected i n to th e blood prod u ce

a m arked sti mul a ti n g effect u po n th e org an ism Accordi ng .

to this i nves t ig ator t he t estis produc es som e su bst an ce which

,

p asses i n to the circul atio n S uch a process is term ed i n te r

.

n al secretio n in co n t r ast to the productio n of subst an ces

,

wh ich are co n veyed to th e outside o f a gl an d by a duct ,

as in th e secretio n o f s aliv a or bile I n rece n t ye ars the

.

subj ec t of i n ter n al secretio n has beco m e o n e o f th e mos t

i mport an t an d frui tful fi elds of physiologic al r ese arch .

All of th e cells of th e body give off subs t an ces i n to the

blood or lym ph but o n ly in a co mp ar atively few c ases has
, ,

an
y defi n ite p hysiologic al fu n ctio n of th ese products be e n

discovered Two i mport an t i n ter nal secretio n s sug ar and

.
,

ure a are formed by th e liver th e form e r arisi n g from t h e

, ,

glycoge n w hi ch i s stored in t h e h ep a tic cells Th e p ancre a s .

produces in addi tio n to the p an cre ati c j uice an i n ter nal

, ,

secre t io n which is of eve n gre at er i mport an ce to th e org an

ism Remov al o f the p an cre as from o n e of th e higher
.

an im als resul t s in th e produc t io n o f di abe tes which soo n ,

termi nates fat ally I n t his dise ase there i s an a b n orm al

.

amou n t of sug ar in th e blood u n der ord i n ary circu mst an ces

the u n du e produ ctio n of this subs t an ce is preve n t ed through
the age n cy of som e secretio n w hich is give n o ff fro m the
p an cre as i n to t he ge n e r al ci r cul atio n I f the duc t o f th e.
220 TH E B IOLOGY OF TH E F RO G CH A P
.

p an cre as be ti ed so as to destroy the ordi n ary fu n ctio n of

t hi s org an t here is n o ab n orm al produc t io n of sug ar an d the
, ,

an im al m ay live for a lo n g tim e A l arge p ar t o f th e p an .

cre as m ay be re moved o r th e whole o rg an re moved an d a

, ,

p art of i t gr afted in som e other p art o f th e body wi th ou t

produci n g fatal effec t s The an i m al m ay al so b e kept aliv e
.
,

eve n after co mple te extirp atio n of th e p an cre as i f extr acts ,

o f t his org an are i nj ected i n to t h e blood S o lo ng as the .

body receives subst an ces form ed by th e p an cre a s it m ay be

kep t alive bu t w he n these are com pletely wi thd raw n fat al
,

effects quickly follo w .

I n n e arly all vertebr ate an i m als th ere are sev er al org an s

the fu n ctio n o f w hi ch w as for a lo n g t i m e u nk n ow n M an y .

of t h e m were reg ard ed as rudime n t s o f org an s useful o n ce ,

b u t n o w fu n cti o n less Th is w as t h e c ase wi t h cert ai n sm all

.

s truc tures su ch as the thyroi d hypo p hysis an d adre n al , ,

bodi es I t is n o w k n o w n th at c er t ai n of these org an s far

.
,

fro m bei n g use less rudim e n ts are absolutely esse n ti al to th e

,

m ai n te n an ce of life M ost of th ese org an s belo n g in t h e

.

“
c at egory o f duc t less gl an ds so c alled bec ause t hey h ave ”
,

n o duc t or ex t er n al ou tlet The w ay in whi ch th ey fu n ctio n

.

h as bee n a m at ter of d ispute We k n o w th at t h ey ac t by

.

produci n g i n t er n al secre t io n s which a re give n off i n to the

blood an d it i s h eld by som e t h at t h es e subst an ces d estroy
,

poiso n s which are produ ced by t h e o th e r t issu es an d w h ic h

would c au se t h e d e ath o f t h e organ ism if allowed to ac cu m u
l at e. O t hers regard t h ese secre t io ns as affo r di n g the
sti muli n eed ful t o t h e disch arge o f t h e fu n ctio n s o f o t her
organ s I n cert ai n c ases the l at ter i n t erpre t at io n se ems
’

.
,

to be bor n e ou t ; but th is do es n o t prove t h at the i n t er nal

secre t io n s of o ther org an s d o n o t possess a n t itoxic proper
ties an d in fac t t here seems t o be good evide nce in som e
, ,

i n st an ces th at suc h i s th e c ase

, ,
.
222 TH E BIOLO G Y OF TH E F RO G CH AP .

co n vert th e p r o teid spli tti n g e nzym e of th at o r gan i nto an

-

active fo r m D oubt has however bee n rece n tly throw n u po n

.
, ,

th is co n clu sio n .

The Th yr oid G lan ds —Th e thy .

r oid gl and s o f th e frog are co m ‘

p le t ely s ep ar a ted fro m e a ch other ,

bei n g situ ate d o n either side of th e

A g hyoid app ar atus in a sm all sp ace
.

betwe e n its posterior l ater al an d

FIG 6 —D i g m sh
o a thyro hyoid processes G au pp h as
ra ow -

“ f a“ d escribed som e thyroid tissu e ac

. . .

gigi; 2 32 1312 i ;
l
(
1 t
a era l p c ss f hy id c esso ry t hyro rd) o n the ve n tr al side
ro e o o

hifiihgid f bf 12 3213 bee n able to co nfirm this observa

o f th e hyoglo ssus muscl e an d I h ave
ta “
,
ce s

tio n in Ra na p ip iens Th e t issu e o f th e thy r oid shows a

.

u n iqu e structure bei n g co mposed o f a m ass o f r ou n de d

,

follicl es u n ited by a
sm all amou n t o f c o n
n e c t iv e t i s s u e in

w h ich the r e is a ric h

supply of blood ves

sels . E ach folli cle
i s a perfectly closed
sac li n ed by a sin
gle l aye r of cubi c al
epitheli al cells In .

th e ce n t er of e ac h
fO HiCle i s a CO HO idal F I G 6 P t f c ss s c ti . 1 f th t h y
. ar o a ro e on o e

m ass O f tr ansp are n t d f R pp p i t h li l l y

ro r o f
an a i i en s . e, e e a a er o

v s i cl s ; m c ll id l subs t n c
e in e sicl o o a a e ve e
s u b s t a n c e W h iC h
.
,

prob ably represe n ts t h e secretio n of th e epith eli al li ni n g .

Th e t hyroid of th e frog like t h at o f th e high er vertebrates

, ,

has bee n fou n d to sec r ete a subst an ce r i ch in iodi n ( iodo

x1 1 T
I N E RNA L S E CRE TI ON A ND D U CT L E S S G L AND S 223

t hyrin, thyroiodi n) L i t tle is k n ow n o f i ts fu n ctio n in th e

.

frog I n th e high er vertebr ates its remov al is followed in

.

n e arly all c ases by fat al effects Re moval of o n ly a p art

.

o f th e gl an d as a rule c r e ates b u t li t tl e disturb an ce L ife

, ,
.

m ay be m ai n t ai n ed for a co n sider abl e period a fter complete

removal o f t he thyroid by givi n g i nj ectio n s of extracts of the
,

gl an d i n to th e blood I n m an .

t h e dise ase c alle d myxoed em a or

cre ti n ism c aused by th e atrophy

,

of t h e thyroid is ofte n muc h

,

h elp ed o r eve n cured by th e

ad mi n i s t r a tio n o f t hyroi d extr ac t .

The subst an ce t o which t h e thy

roi d owes it s i mportan t fu n ctio n i s F ro 6 Di g m sh i g t h 2 a ra ow n e

a p ro t el d W ith wh ich a co mp a r a
. .

p siti f th o t hymus M
on o e , ,

t ively l arge amou n t o f i odi n is in d p ss m dibul

"
e re or an ae

combi natio n T reu p el fou n d t h at

.
s l ; Ty t ym p
in um
o e ’
an '

frogs fro m which bo t h thyroid s were removed live d o nly two

o r three d ays bu t h e w as no t e n t irely cer t ai n th at th e r esul t
,

might no t be due to effects of th e oper atio n other th an th e

loss of t h e p arts in qu es tio n .

Th e Th ymu s The lliy in ns is a sm all ov al org an some

.
, ,

wh at reddish ih color situ ated behi n d the tymp anic m e m

,

br an e u n der the d epre ssor m andibul ae muscl e As in mos t .

higher forms th e thymus d imi n ish es in si ze with age M aurer

, .

fou n d th at in Ra n a es cu len ta th e thymus att ai n ed i ts m axi

mu m size in sp eci me n s o f two or three ce n timeters in l e n gth .

I n old frogs ( 7 to 8 c m ) th e org an is mu ch sm aller an d sho w s

.

m arks of d ege n er a tio n in st r ucture .

Th e thymus has esse n t i ally th e structure of a lymphoi d

gl an d I n it s fi n e n e t w o rk o f ade n oid tissue li e n u m erous
.

s m all rou n d ed cells ; The re are also several l arge cells of

,

co nce n tric s t ructure co n cern i ng whose origi n an d sig nific an ce

224 TH E BIOLO G Y OF TH E FRO G CH A P .

t here has be e n much discussio n but of whose fu n ctio n ,

n o t hi n g posi tive is k n o w n .

I t is prob able th a t blood co rpu scles are produced to a

cert ai n exte n t in the thymus ( M ayer) Accordi n g to Abe .

lous an d B ill ard if both th ym us gl and s of th e frog are

,

removed th e an i m al soo n dies after a period o f g r e at m us

, ,

c u lar we ak n ess ulcer a tio n of the ski n a n d a v a rie ty of ot h e

,
r ,

p a t h ologic al sy mptoms H am m ar ho w ev er failed to c o n

.
, ,

firm these results H e fou n d th a t bo th thymu s gl an ds m ay

.

be re moved from th e frog wi th ou t i nj ury an d co n cludes th at ,

t h e results ob t ai n e d by A b elo u s an d B ill ard were the effects

o f acci de n t al i n fec t io n .

The P seu dot h yr oid an d t he E p it h elial B odies — Th e

s eu do th r oid an d the ep itlielia l é e aies are org an s of si m il a r
’

p y
structure an d origi n They are derive d fro m t h e m o difi ca
.

tio n o f th e epi theliu m o f th e gill sli t s o f th e l arva and are

t herefore prod uc ts o f th e e n tod erm The two pseudo .

thyroids are the l argest o f these They are rou n ded .

r eddish bodi es lyi n g o n ei t her sid e o f the posterio r por t io n

,

of th e hyoid c ar t il age They were form erly mist ake n

.

for the thyroid s b u t th ey possess a very differe n t i n ter

,

n al structure ,
which i s esse n t i ally th a t o f a lympho id
gl an d .

The ep illielia l l adies are s m all rou n ded stru ct u res u su ally
,

m ore th an two in n umber o n e ach side a nd so me w h at v ari

able in positio n bu t ge n er ally si t u a ted n e a r th e p seu do t h y
,

roids As an org an prob ably belo n gi ng in th e s a m e c a te

.

gory as the prec edi n g m ay be m e n tio n ed th e p ro p eric ardial

body which i s a tran sverse ov al org an lyi n g ve n tral to the
,

hyogl o ssu s muscle be t w ee n t h e thyroids I t possesses a .

ly mphoid structure an d is l arge r in you ng th an in old frogs

ro m s mode o f d evelopme n t M au rer cl asses
( G au p p
). F i t

the c aroti d gl an d also amo n g t h e epi theli al bodies but its ,

‘
22 6 TH E BIOLOGY OF TH E F ROG CH AP .

from th e p e r i to n eu m whil e the ce n tral cells are ge n er ally

,

reg arded as m odific atio n s of the cells of th e symp athetic

g angli a I n th e high er vertebr ate s th e ce n tr al c ells form a
.

si n gle m ass whi ch is surrou nded by a defi n i t e c ortex but in ,

the frog they are sc atte r e d th r ough the co r tic al c ells in an

irregul ar m ann e r .

A b elo us an d L an glois fou n d th at i f both adre nals o f th e

frog were d estroyed the ope ratio n w as soo n follo wed by fat al
,

effects bu t i f o n ly o n e adre nal w as d estroyed th e an i m al ,

would co n ti nu e to live I f a fter th e destru ctio n o f both

.

adre n als portio n s o f o n e o f the bodi es were tr a n spl an ted in

th e dors al lymph sp ace life w as m ai n t ai n ed for a co n si de r

,

a bly lo n ger p eriod th an would oth erwise h ave bee n possible .

I t is well k n ow n th at th e adre n als produce an i n ter n al secre

tio n u po n which th e life o f th e organ ism is depe n de n t This .

m ate r i al (adre n ali n epi n ephri n ) m ay be extracted fro m th e

,

bod ies an d i ts physiologic al a c t i o n tested I t h as bee n

.

m uch experi me n ted wi t h a mo n g higher an i m als an d is n o w ,

u sed to a co n sid erable exte nt in m edici n e an d surgery I t .

h as the property o f gre a tly i n cre asi ng blood pressure by

c ausi n g a s tro n g co n tractio n of th e smooth muscle fibers o f
t he blood vess els .

E xp erime n ts o n the effects of th e extr act o f the adre nals

o f th e frog show th at this subst an ce h as mu ch the s am e
properti es as am o n g m am m als Whe n i nj ected i n to th e
.

blood of a m amm al it produces a m arked r ise in blood pres

,

sure an d o n the oth er h and i nj ec t io n of th e extr act fro m

, ,

th e m a m m ali an gl an d i n to th e frog produces v ery m arked

r esul t s which m ay be fat al if th e dose is l arge M oore an d
,
.

V i n ce n t fou n d th at “ a ft er i nj ectio n o f a glyc eri n extr act

equ ivale n t to about 5 g of th e fresh gl an d i n to th e

.

dors al lymph sac p ar alysis i mmedi ately com es o n

,
.

With l arger doses there are sp asms and fi b rillary twitchi ngs
x1 1 I N E RNA L T S E CRE T I O N A ND D U CTL E S S G L AND S 227

in v arious p arts With sm aller doses g of th e fresh . .

gl an d) O liver an d S ch afer fou n d a si mil ar p ar alysis bu t ,

i t c ame o n more slowly A fter h alf an hour the an im al .

appe ared to be n e a rly i f n o t quite in a n orm al co n ditio n

”
.
, ,

I nt ern al S ecr et ion s as a Mean s of F un ct ional Correla

t ion . F ro m
wh at h as bee n s aid it is evide n t th at i n ter n al
secretio n s pl ay an i mport an t rOle in securi ng th e c o Ordina
t io n of fu n c t io n s o f t h e v arious org an s of th e body They .

ac t as regul a t ive age n ts m aki n g possible the p arti al co n trol ,

o f o n e o r g an by anoth er i n dep en de n tly o f th e ce n tr al n ervou s

system O rg an s through their i n ter n al secretio n s m ay act
.

an d re act u po n e a ch o ther an d in this w ay bri ng about ,

the h armo nious fu n c t io n i n g of the differe n t p ar ts which is

esse n ti al to the life of the whol e .

RE F E RE NC E S
Ab elous , J . E . S ur l act io n ’
ant i t xi q u
o e des c p su l s
a e s urren ales .

C R S oc
. . . Bi l o .
, 1 895 .

Ab elou s et B illar d ch ch s l . Re er e sur es fo n t c i s d t hymus ch z

on a e

la gr en o uill e . Ar ch P hys N m t P t h
. . or . e a . Ann é 8 S
e,
5 T 8 2 , er . , .
,

1 8 96 .

Ab elous et L an g lois . No t e s ur le s fo n t ci s on de s c psul s su

a e rre

n ale s ch z l g uil l C R S B i l 8 9 L m t
e a re n o e. . . oc . o .
, 1 1 . a or des g uil l s re n o e

p es l d s t uc t i d c psul s l i 89 T i it d l
’
a r a e r on es a e s urre n a es, . e. , 1 1 . ox c e e ex

tra i t l h liq d muscl d g uil l s p i s d c p sul s

a co o ue l a e e re n o e r ve e a e su rr en a es,

l. e 89
.
, R ch ch s x p im t ll s
1 2 . l f c t i s d c p su l s
e er e e er en e e sur es on on e a e

l d l g
s u rre n a e s ui l l A ch P hys N m t P t h
e a T 4 89 re n o e . r . . or . e a . .
, 1 2 .

S url f c t i s d c p su l s
es on l l T 4 89
on es a e s u rre n a es, . e . .
, 1 2 .

B b r E C a R s ach s
e ,t h M i u t S t uc t u
. f t h Thy ids
. e e r e on e n e r re o e ro .

Ph il T s 8 8 p t 3
. ran .
, 1 1, ar .

B ol H Gl dul t hy id
au , dGl du l Thymus d A m p h ibi
. an a ro ea u n an a er en .

Z l J h b Ab t h f A t B d
oo . a r 8 99 . . . na .
, . 1 2, 1 .

G le A au B i l gic l Ch g s i th S pl
, . f th F g J u o o a an e n e e en o e ro . o r .

M ph V l 8 8 93
or .
, o .
, 1 .

G on f i R ch ch s p hysi lr n l f cti
. d gl d s e l er e o . su r e on on u an e su rr en a es .

R e v.m d S u i ss m d T 6 8 96
e . e ro an .
, . 1 , 1 .
228 TH E BIOLO G Y OF TH E F RO G CH AP .

H am m a r J ,
. A . I st die T h ym usdru se e ro b i m F sch e in leb e nsw ich
t ig s O g
e r an ? Ar ch .
g e s , B d 1 1 0,
. P hys
p 33 7 . . . .

rr ing P T
He ci P i ui y x c s o n t h e H t f
, . . T he A t on o f t t ar E t ra t ear o

the F g J u P hys V ol 3 1
ro . o r . .
, . .

M ay er S Z L h e v n d,
S hildd e n d Thymus b i d n
. ur e r o er c rus u e e

A m p h ibi n A n t A n B d 3 1 8 8 8
e . a . z .
, .
, .

M au rer F S ch ildd Thym us n d Ki m n e t d A m p h ibi n

,
. ru se , , u e e r s e er e .

M o rp h .
J ah rb .
, Bd . 1 3 , 1 888 . Die E p icle r m is und ih re Ak mm lin g e ,
1 895 .

M oor e m p t i Ch mis t y f t h S u p
an d V incen t . T h e Co a ra ve e r o e rare nal

C p sul s
a P c Ry Se . L d V l 6 897 ro . o . oc .
, on o n, o . 2, 1 .

Oliver an d S ch f r O t h P hysi l gic l A c t i f th E x t c t a e . n e o o a on o e ra of

th S up e l C p sul s J u P hys V l
rar e n a 6 8 94 a e . o r . .
, o . 1 , 1 .

Tr p l J euS t ff chs lu t such u g b i i m m it I d t hy i

e , . o w e e n er n e e ne o o r n

( Th i di ) b
y ro oh d lt F ll M
n yx d m d M it t h eil g i ig an e en a e vo n oe e un e un e n er

Thi h m it I d t hy i ( Thy i di ) M ii h
e rve rsu c e m d W ch o o r n ro o n . nc en er e . o en

s h 43 J h g 8 96
e r. a r .
, 1 .
 RA UL

F IG 63 A, S e —
k l t f R l mp e on Th ol ft limbs an a esh uld gi dl
o r a r ra e e o er r e,

m m b s f h l f sid f h skull m d C il gi us p t s d
. . .
,

e b ra n e o n e o t e e t e o t e a r e re o ve a rt a no ar ot
m s c il g b s i h i ck ; h s f m m b b s i i l i c c p i ls
.

na e of art a e one n t t o e o e ra n e o ne n ta a ta a

c u f hy id ; d o c bulum s t g lus ; o l y b s hy l
. .

i
a n ter o r o rn o o a a eta l AST a ra a i a r a ; C , ca
c lc um ; X x cci p i t l ; f mu ; f f f t ll s ; K f
-
.
, , ,
’
CA L a ane E O C, e o a FE e r on on o n an e e P PA , ro

p i ls ; U hum us ; il i um ; M X m x i ll ; lf p lf c t y c p sul ; t p
, . .
, , ,

ar e ta H er IL, a a o e o a or a e o r

c ss ; s i c u hy id m x ll ic
, .
, ,
,

p ro e
p f o ; R ik
ter o r/X p io rn ; Ro p o o ; R re a a P OT, ro t A
d i ul ; squ m s l su sc ul
.
,

ra o -
na
p h h m
S P ET H id S Q s ;en et p o ; p ; a o a S S CP , ra -
a a su s ,

s s c ss u s yl
. .
, ,

p i m
e nso r u ; fi b l ; i p p u a U n t r; V l
t ra n ve r e ro e ; ST , ro e c er t

t b V s c l b — digi s f u h b
- .
, ,

ve r e
; ra ; V 0a m
ra ; v e rt e B ra t h t t
vo er f I V t e o r ve r e ra s ee n
9
i f i zyg hysis c um l mi ; p u l sp i
.
, , , ,

n ro n t
yg a z p p a n t er o r
; ; l m a o en , e ntr a na n s ne ra ne ;

dicl s s c ss m k s ll s l gy sh
.
. .
, , .

p e ; e p ( F
t ra n v er e P d roH e Z o ro ar er an a w e
'
o o

l d f m H s)
. ,

a t e re ro ow e .
CH A P . x1 1 1 TH E S KE L E T ON 23 1

is plugged wi t h c artil age ag ai n s t which abuts th e i nn er en d

of th e colu mell a o f the ear .

I n the ve n tr al sid e of the skull is a l arge bo n e th e p a r a ,

oa sa l o r p a r a sp lien oizl which is in t h e sh ap e of a d agger with

, ,

E m 9

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out any h an dle the l ater al po r tio n s u n derli e th e two pro otics .

The f r on lo p a r iez a l bo n es form most o f the roo f of the

-
‘
23 2 TH E BIOLO G Y OF TH E F RO G CH AP.

skull Alo n g th e m iddl e li n e they are u ni ted by the s ag itta l

.

sutu re E a ch represe n ts t w o bo n es a fro n t al an d a p ariet al

.
, ,

an d in the e arly st ages o f th e d evelopm e n t of th e skull these

eleme n ts are sep arate bu t subseque n tly they fuse i n to a

,

si ngle bo n e .

The an terio r end of the c r an iu m i s sur r ou n d ed by a bo ny

r i ng th e elnmoia ( o r spnenellz moia ) bo n e This i s ove r
,
’ ’
.

l app ed by th e fr o n to p ar iet als above and th e p arab as al b e

-

low and i s sep ar ated fr o m the prootics beh i n d by quite a

,

lo ng i n terval o f u n ossified c ar til age Th e an terio r p ar t o f t he

.

eth moid is wide n ed out an d divid ed i n to t w o c h a mbers by a

m edi an vertic al p arti tio n Th e exp an ded po r tio n forms th e
.

posterior wall of th e nas al c avity ; th e l atter m ay be s ee n to

co m mu n ic ate with th e c ran i al c avity by a p air o f s m all ope n
i ngs through which th e olfactory n e r ves p ass The r est of .

the n as al c apsules are forme d m ai nly by c ar tilage .

Th e n asa ls are t w o n arrowly tri an gul ar bo nes lyi n g above ,

th e nas al c apsules thei r b ases which lie n e ar e ach other in

,

the middle li n e are sep ar ated fro m th e fro n to p ari et als by a

,
-

sm all p art of the r oof of th e e thmoid .

Th e vomers lie v e n tr al to the n as al c apsules ; e ach h as

thre e ou te r p r ocesses betwee n the two posteri or o f which
,

occu r the i nternal n ar es 3 the ve n t ral su r face be ars the

vomer ine leetn
—The
.

S u spen soriu m and J aw s j aws are att ach ed to th e

.

cr aniu m by me an s o f an i n termedi a te susp e nsory a pp ar at us

in which the followi ng sep ar ate bo n es are to be disti n
u ish e d
g
( )1Th e t m
y p a n ic
(q
s u a m os a l ) a T sh a ped bo n e th e,
m a i n -

li mb o f which exte n ds ou twar d an d b ackw ard t o the an gle

o f the j aws th e pos terior en d o f the cross piec e articul ates
w ith the p ro Ot ic while th e an terior e n d exte n ds obliquely
,

dow nw ard in fro nt B elo w t h e t ymp anic lies ( 2) th e p loz y

.
23 4 TH E BI O L O G Y OF TH E FRO G CH A P .

pro mi n e n ce the cor onoicl p r ocess wh ich gives att ach m e n t to

, ,

the muscl es for closi n g th e j aw .

Th e clen ta le li es o n th e outer side o f th e dist al en d of th e

an gul are overl appi n g M eckel s c artil age which there ru n s
,
’

bet w ee n the se t w o bo n es T he apic al portio n o f the m an .

dib u lar arch co n sists of two short mov able elem e n ts the ,

m en to mecleelia n bo n es which r esul t from the ossific atio n of

-

th e dist al portio n s o f M eckel s c artil age T hey u nderlie th e ’

.

pre m axill aries an d wh e n they are r a ised c au se a co r respo n d

, ,

in g elev atio n o f th e l atter bo n es .

The H y oid T he br an chi al skele to n of th e frog is co m

.

posed o f th e liy oicl ca r tilag e an d i ts processes The body .

of the hyoid is a l arge flat pl at e of h y ali n e c a rtil age more ,

or l ess qu adrate in ge n er al ou tli n e ly in g in t h e floor of t h e ,

bucc al c avity I t s an terior m argi n where t h e b ase o f th e

.
,

to ngu e is i n serted i s stro n gly c o n c ave At th e an terior en d

,
.
‘

o f the body a rise th e a n ter io r co r n u a whic h are lo n g slen , ,

d er r ods o f c artil age which p ass b ack w ard an d u pw ard o n

,

ei ther side o f th e thro at an d j oi n to the p ro Ot ic bo n es o f th e

skull .

O n th e i nn e r side o f th e b ase of e ach co r nu is a short

an t erior proc ess Th e a la ry p r ocesses fl atte n ed dis t ally
.
, ,

exp an ded pl ates of c ar t il age arise j ust behi n d th e an t erior ,

cor n u a The p oster o l at eral an gles of the body are produced

.
-

to form the poster o la ter a l p r oc sses The t/iy r o icl p r ocesses

- e
.

diverge from th e middle p art of th e posterior m argi n o f the

body an d lie o n ei ther side o f t h e l ary n x which they h elp to ,

suppo r t They are th e o nly p ar ts of the hyoi d app ar atus to

.

become o ssifi ed .

Th e Car t ilagin ous Cr anium — The skull of the frog is

compo sed of c ar t il age t o a much gre ater ex t e n t th an in th e
h igher ver t ebr at es O nly here and th ere does i ts origi
.

n al c a r t il agi n ous b asis becom e co n verted i n t o b o n e The .

XI I I TH E S KE L E T ON 23
5

v ar ious bo n es which co n sti tute th e skull m ay be divided as

r eg ar d s their origi n oo n es which r esult fro m
,

L
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m
23 6 TH E BIO LOGY OF TH E F RO G CH A P .

d eveloped from m e mb ran es The l atter are more superfici al

.

in positio n an d m ay be stripp ed off fro m the r es t o f th e skull ,

le avi ng a sort o f c artil agi n ous box with o ssifi c at io n s here and
there whi ch represe n t the so c alled c artil age bo n es I n th e
-

cr an iu m o n ly th e exo ccip it als th e pro o tics and th e eth moid

, ,

a re c ar t il age bo n es th e fro n to p ari et al p ar ab as al n as als

,
-

, , ,

a n d p al at i n es ar e d eveloped fro m m e mbr an es and m ay ,

re adily be sep ar ated from th e u n derlyi n g c artil age The .

c a rtil agi n ou s cr an iu m fo rms an al most compl ete c ase for the

br ai n ; th e r oo f however is i n co mplete there b ei n g a l arge
, , ,

oblo n g ope n i n g or f mz lcm e/Ze the f mesim

, ,
n e ar

th e middle an d a p ai r o f sm aller op e n i n gs f en esir x p a r ie

, ,

ta les farther b ack

,
.

Th e s uspen sor iu m an dj a w s also h ave a c ar til agi nous b asis

which is co n ti n uou s wi t h t h at o f th e cr an iu m proper The .

bo n es of th ese p arts w i t h the exceptio n of th e qu adr a to

,

j ug al an d the m e n to m eckeli an bo n es are d eveloped fro m

-

m e mbr an e A c ar t il agi nous b ar ru n s betwee n the pterygoid

.

an d tymp an ic t o the a n gl e of th e u pper

j aw wh e n ce it is ,

co n ti n ued forw ar d be n e at h th e m axill ary as far as the p al a

ti n e bo n e wh ere a tr an sve r se process joi n s th e posterior en d
,

o f th e n as al c ap sule The expo se d c artil age o n th e sid e of

.

th e cr an iu m i s perfor ated by a la rge for a me n for th e optic

n erve an d by sm aller o pe n i n gs for th e third and fourth
n erves . Th e an gul are an d de nt ary of th e lower j aw are
me mbr an e bo n es appli ed to th e c artil agi n ous core o r ,

M eckel s c artil age

’
The thy r oid p rocesses o f th e hyo id are
.

developed from c artil age .

The frog s skull represe n ts a typ e betwe en the skulls o f

’

th e lowe r fi sh es an d t hose of the higher v erteb rates In .

such forms as sh arks and sk ates the c r an iu m and i ts v arious

appe n d age s are e n tirely co mposed o f c a r til a ge H igher up .

in th e sc ale we m ee t with fish es su ch as som e of the g an oids , ,

23 8 TH E BIOLO G Y OF TH E FRO G C H AP .

The first vertebra or eri/a s differs fro m the oth e r s in hav

, ,

ing n o tr an sverse proc esses in th e abse n ce of the an terio r ,

z yg apop hyses an d in h avi ng in fro n t a p air of ov al co n c ave

, ,

facets for articul ati n g with th e occipit al co ndyles of t h e

skull Th e n i nt h vert ebr a h as the tran sverse processes very
.

stro n g and direc t ed obliqu ely b ack w ard i t h as n o posterior

zygapophyses an d th e post erior su r fac e o f the c e n trum be ars
,

a p air o f pro mi n e n ces

for articul atio n with th e

$ 3 90
urostyle .

Th e zz r esly /e is ele
vat ed o n th e dors al
side i n to a pro mi n e n t
keel which exte n ds
n e arly t o the posterior

en d The an terior sur .

— fac e p o ssesse s a IJ ai r
F IG . 66 T sv s s ct i ran f sh uld
er e e on o o er

gi dl c c id ; p
.

r e cor o ra p i ido; O f c
e a v it
.
ies
corfO ,
l ear t l l la
c o ra co
’

g ] gl i d c i ty ; b hum us ; p sc p tio n W ith th e n i n th

.
,

en o av u, er so . a

p su p sc p ul ; 3 t h i d v
,

u al ; s .s c ra -
a a r er
ver ebr Th e verte
,

t b f k k
,

P P t a"
e ra (.A t n d
er )
ar er a ar er .

b r al c anal is s m all an d
tri an gul ar in ou t l i n e There is a p ai r o f sm all ope n i ngs
.

through the sides of th e uros tyle n e ar the an terior en d for

the exit of the l ast p air of spi n al n erves .

Th e ce n tr a of th e vertebrae are j oi n ed togeth er by m e an s

of p ads o f hy ali n e c artil age ; co n n ecti ng lig am e n ts exte n d
alo n g both the ve n tr al an d th e do r s al su r faces o f th e ce n tr a

a n d th e arches an d n eur al spi n es are j oi n ed b l ig a me n ts

y .

Th e spi nal n erves m ake their exit through the z m er ver teé r a /
'
‘

f m be wee the sides of the eu r l rch es

'

o r a z rz a t ,
n n a a .

The P ect or al G ir dle an d S t er nu m The pector al girdle .

-

i s a bo n y arch which gives support to the fore li mbs Th e .

upper en d o f the girdle is fo r med by a flat dist ally exp an ded ,

XI I I TH E S KE L E TO N 23 9

p ortio n th e s u r a s cap u la
p ,
which is com posed of c artil age ,

which i s more or less c alcifie d at th e b ase The supr a .

FIG . 67 .
—
M iddlsh uld g i dl f th f g f m b l
e p art of C
th e o er r e o e ro ro e ow . 0,

c c id ; C p i c c id ; Cl cl icl ; E p p is um ; G gl id
o ra o o
'
e o ra o av e e t e rn ,
en o

c i y ; F f s t ; K C c il g b t sc p ul d cl icl ; 10
, , ,

av t e, ene ra a rt a e e w e en a a an av e 2.

xi p his um ; m j u ct i id ; S sc p ul ; S t s um
,

t ern f pi n on o e c o ra co s a a ,
t e rn .

f
, ,

( A W i d
ter h im ) e e rs e .

sc apul a articul at es belo w with the lo n g scap u la which is ,

oblo n g and co n stricted in th e m iddle t h e pos t erior side o f

t h e lower en d forms p a rt o f the g len o z a f ess a w hich receives
'
’
,

t h e h ead p f the fra m er zcs

. A n otc h occurs at t h e gle n oid .

foss a dividi n g the lower en d of t h e sc apul a i n to dors al or

,

glen oid p art from a ve n tr al acr o m z a ! por t io n

'

F ro m th e lo w er e n d o f th e sc apul a t w o b ars exte n d

tow ard the mid dle li n e The an terior of th ese co n sists of .

a b ar of hy al i n e c ar til age th e r oco r a coid in fr o n t o f an d

p , , ,
240 TH E BIO LOGY O F TH E FROG CH A P .

p artly i nclosi ng which i s a m embr an e bo n e the cla vicle

, ,
.

A t its outer e n d t h e cl avicl e i s be n t forw ard an d appl ied to ,

a forw ard proj ectio n of c artil age th e a cr o m z mz at th e lower

'

, ,

e n d of the sc apul a The poste r io r b ar th e cor a coz a is a

'
’
.
, ,

stout bo n e co n stri ct ed in th e middle and bro adly exp an ded

,

a t i ts i n n e r e nd a t i ts oute r extremi ty i t forms a portio n o f

th e gle n oid foss a B etwe e n th e two e nd s of the cor acoi d

.
,

a n d exte n di n g forw ard betwee n th e cl avicles are th e t w o ,

e z co r a cez a ca r tilag es which are u su ally more or less c alcified

' '
’

p ,

alo n g th e ve n tr al side o f t heir li n e o f u n c t io n

j .

Th e 8p l$ f€ 77 27l 772 lies in fro n t o f th e e p ic o raco ids an d co n

sists of a b as al p iece of b o n e an d a termi n al pi e ce o f c arti
l age w hich has an al most circul ar exp an sio n at th e an terior
,

en d .

The s ler mcm proper r esembles the epi ster nu m in ge n e ral

s h ape an d in bei n g co mposed of a proxi m al pi ece of bo n e
,

( m esoster n u m ) an d a dist al exp an d ed pi ece of c artil age .

Th e l atter h as a n otch in t h e middle of its posterior m argi n ,

which receives th e anterior abdomi nal vei n j ust before it

le aves th e body w all B oth the ster n u m an d the epi ster nu m
.

are c a p able of a li mited vertic al moveme n t esp eci ally a t th e

,

fl exible c artil agi n ous e n ds .

I n addi t io n to formi ng a pl ace o f att achme n t for th e

mu scles which move the fo r e li mbs th e pec t or al arch pro
t ec t s the v arious i n t er n al o rg an s su ch as t h e lu n gs he ar t
, , ,

etc an d serves to m ai n t ai n th e ge n er al form of t h e body

.
,
.

I t also gives att a ch me n t to the ve n tr al muscles which dr aw

b ack th e hyoid an d d epress the floor of the mou t h an d for
, ,

t h ese purposes th e di st al e n ds o f t h e ster n u m an d e p ist er

n u m ar e fl att e n ed an d exp an ded t heir c artil agi n ous cousis

,

t e n c y e n abli n g th e m to accom mo dat e th emselves to th e

ch an ges produ ced in th e b o dy w all .

The F ore L imb s — Th e u pp er bo n e of the fore limbs is

24 2 TH E BIO LOGY OF TH E FRO G CH A P .

dis t al end of th e r adio ul na is wide n ed an d e n ds in two -

epi physes o n e for e ach of the co mpo n e n t bo n es

,
.

Th e ca rp zcs or w rist o f th e frog co n t ai n s six bo n es arr ange d

, ,

in t w o ro w s I n th e proxi m al ro w t h e a lrz a r e an d r a clz a le

'

are si tu ated at th e e n ds o f th e ul n a an d r adius respec t ively , ,

and at th e i n n e r o r pre axi al sid e o f th e r adi ale is th e cen t r a le .

I n th e dis t al row th e first ca rp a l

occurs j ust behi n d the rudime n t ary
thu mb ; th e seco n d c arp al a very ,

sm all bo n e li es behi n d the seco n d ,

digit ; the outer c arp al i s of rel a

t ively l arge si ze an d i s formed by th e ,

fusio n of th ree or ig inally dis t i nct

bo n es o n e for e ach o f th e three .

outer digits of the h an d B eyo n d .

th e wrist are th e five metaca rp a ls ,

whic h form th e skeleto n o f t h e p ro xi

m al p art of the h an d The first m et a .

FIG . 6 9 — P l ic gi dl
.
c arp a l is rudi m e n t ary an d u su ally c a r
e v r e

o f th f g f om th t ilagino u s in th e fem ale but in th e

e ro r e ,

r igh t sid G t b m al e i t is l arger , and becom es c alci

e . ac e a

ulu m ; l l ilium ; 1
,

s,

ischium ; P p ubis fi ed or eve n o ssifi ed Th e other

,

.
.
,

( A ft er P k nd m et ac arp als are elo n g ated cyli n dric a l

ar er a

P k )
ar er .
bo n es so m ewh at exp and ed at the t w o ,

e n ds ; the i n n e r o n e o f th e fou r is stou ter in th e m ale th an

in the fem al e an d be ars a pr o mi n e n t crest o n th e i n ne r
,

m argi n The met ac arp als exce p t th e first are succeed ed

.
, ,

by th e p h a la nges o f which e ach o f the two outer digits h ave

'
,

three w hile t he two i n n er o n es h ave but two

,
.

Th e P elvic G ir dle —The pelz z c g z r a le w h ich supports

' '
'
.
r

the hi n d li mbs is a V sh aped m ass o f bo n e th e apex o f -

which lies at the pos t erio r e nd o f the skeleto n an d receive s ,

th e tip o f the urostyle t h e two an terio r e n ds are u n ited to

XI II TH E S KE L E TO N 243

th e l arge tr an sverse proc esses of th e n i n th or s acr al verte

,

b ra E ach h alf of th e pelvic girdle is composed of thre e

.

elem e n t s the z lz a m the z sclz z a m an d the p et als The first

' ' ' '

, , , .

n a med i s a lo n g bo n e a tt a ch ed to th e n i n th ve r tebr a in
,

fro nt an d m eeti ng its fello w behi nd o n th e dors al side is a

,

high thi n crest ; at the an terior en d of th e ve n tral side of

,

the ju n ctio n of the two ili a is a promi n e n c e c all ed the an te

rio r spi n e of th e p elvis The iliu m forms th e an terior p art .

o f the cup or acet abulum wh ich rec eives the he ad of th e

, ,

femur .

Th e pubis is a tri angul ar m ass of hy ali n e o r in older frogs ,

c alcifi ed c artil age o n th e v e n t r al side of the pelvic girdle ;

,

i t forms a p art of t h e lower sid e o f the acet abulu m .

Th e posterio r portio n o f th e p elvic girdle i s rep re

se n ted by t he ischiu m ; i t form s th e p o sterior p art o f
t h e ace t abulum an d exte n ds for w a rd dors ally as far
,

a s th e top o f a pro mi n e n ce th e posterior spi n e o f th e ,

p elvis which is situ ated above an d a little behi n d th e

, ,

a cet a bul u m here i t is fused wi t h the iliu m ; th e suture

be t wee n th ese bo n es can best be see n in you n g frogs .

The li n es o f u n io n of th e pubic an d i schi al bo n es of

op p osite sides are k n ow n r espe c t ively as the p u é zc and
'

'

isch ia l sy mp lzy ses .

Th e pelvic girdle of th e frog is re m ark abl e o n accou n t of

the elo ng ati o n of the ili a th e reduc t io n o f t he pubis an d ,

ischiu m an d t h e in tim ate fusio n of t he two l atter wi t h t h e

,

posterio r exp an ded p ar t o f th e iliu m to form an al mos t

circul ar m ass .

Th e H in d L imbs The skeleto n of th e hi nd li mb o f t h e

.

frog is co n structed o n esse n t i ally th e s ame pl an as th at o f

th e fo re limb I t co n sis t s of an u pper bo n e t h e fem u r cor
.
, ,

respo n di n g to the hu meru s o f th e fore li mb B elow this is .

th e z z é z o fi é a la correspo n di ng to th e r adio ul na ; an d fo ll o w
’ '
‘ - -

,
244 TH E B I OLO GY O F TH E FRO G CH AP .

in g th e tibi o fi b ula the z a r s zcs an d f eel correspo n d i n g to th e

-
‘
,

c arpus an d h an d The fe mur i s an elo ng ated cyli n dric al

,
.
, ,

very slightly sig moid bo n e ; i t articul ates by its exp an ded

an d r ou n d e d h e ad wi th th e ac et abulu m above fo r mi n g th e ,

hip j oi n t .

The tibio fi b ula is an elo n g a ted very slightly b e n t bo n e

-

som ewh at exp an de d an d fl at t e n ed at ei t h er e n d wh ere it i s ,

m ark ed o n both sid es by a groove which i n dic ates i ts form a

tio n fro m t w o bo n es the tibi a a nd the fibul a which were
, ,

origi n ally se p ar ate Th e t ibi a i s pre axi al in positio n an d

.

correspo n ds to t h e r a dius o f t he fore arm t h e fibul a is post

axi a l an d corres po n ds to the ul n a Ne ar th e middle t h e
tibio —
.

fib u la i s perfor ated by a for a me n for th e an terior t ibi al

a rtery.

Th e la rsa s of the frog i s p eculi arly m odifi ed in th at th e

proxi m al portio n is much elo ng ated an d co n sis t s o f but t w o
bo n es th e pre axi al bo n e n ex t to th e tibi a is c alled the al l
a le th e post axi al o n e n ex t t o the fibul a
,
th e fi aa la r e ; t h e ,

t w o bo n es are u n ited at their e n ds i n closi n g a n arrowly ,

ov al sp ace be t wee n them Th e ce n t r al e is represe n ted by

.

a sm all bo n e o n th e pre axi al side of the dist al e n d of t h e

t ibi ale ; a t i ts d ist al en d it supports th e preh allux The .

ta rs a lia ( which correspo n d to t h e c arp ali a of th e h an d ) are

much reduced bo th in size an d n u mber There is a sm all .

first t ars al behi n d th e b ase o f th e first m et at ars al bo n e .

B e hi n d th e seco n d an d third m e t at ars als i s a sm all b o n e

which rep r ese n ts the fused seco n d an d t hird t ars ali a Th e .

fourth an d fi ft h t ars ali a are abse n t Th ere are five m e t a .

t ars al bo n es all o f whic h are elo n g ated an d cyli nd ric al

,
.

O f th e ph al anges th e fi rst an d seco n d toes co n t ai n

t w o e ach th e t hird an d fi ft h toes three e a ch an d t h e
, ,

four t h toe fou r The sm all p r elz a llax is co m posed usu ally
.

o f two pi eces .
246 TH E B IOLO GY OF TH E F RO G CH A P
.

CH A PT E R XI V

TH E M U S CU L A R S YS TE M
TH E fu n ctio n of muscle i s th e productio n o f moveme n t
t hrough co n tr a ctio n Th e m uscles o f th e fr og re t ai n thei r
.

vit ality fo r a lo ng ti m e a fter they h ave b ee n r e m oved from

the body an d th ey are co n sequ e n tly well ad ap ted for
,

p hysiologic al experi m e n ts The l arge g a s t r ocn e/f z z a s or c alf

'

.
,

muscle o f the frog is so favor able a o n e fo r i n vestigatio n

,

th a t m uch of wh at i s k n ow n of th e ge n er al physiology o f
muscul ar activi ty i s de rived from a study of this obj ect .

Whe n a mu scle co n tracts it i n cre ases in t h ick n ess as i t de

,

cre ases in le n gth C o n tractio n m ay be brough t about as

.
,

i s re adily d emo nstr able with a fresh m uscle by a v ariety ,

o f c au ses si n ce it follows u po n the applic atio n o f n ervou s

, ,

therm al m ech an ic al ch emic al or el ectric al sti muli Th e

, , ,
.

respo n se to sti mul atio n in volu n t ary muscle t akes pl ac e very

quickly an d i n cre ases withi n c ert ain lim it s with th e i n cre ase ,

o f th e i n te n sity o f th e sti mulus I n i nvolu n t ary mu scl e the

.

respo n se i s much slower .

M os t of th e mu scles are att ached by o n e o r both e n ds to

bo n es Th e att ach me n t in som e c ases is direc t in o thers
.
,

i t is by m e an s o f a lef z aoa whic h is a b an d o f very tough

’
, ,

i n el astic co n n ective tissu e Th e ou t er surfac e of a mu scle

.

i s covered by a co nn ective tissu e m embr an e or fa scia , ,

whic h i s more or less el astic The te n do n s of m an y m uscles

.

are formed by a co n ti n u atio n of th e fa sci a which becomes ,

thicker tow ar d the end o f the muscle wh e r e it gradu at es i nt o

,
xrv TH E U U
M S C L AR SYST E M 24
7

a d e nse fibrous b an d Th e en d of a muscle which i s faste n ed

.

to a rel at ively i mmov able p ar t is c alled th e origi n th e more

movable end is th e i nsertio n Th e co n tr actio n of a muscle.

h as the effect of dr awi n g the origi n an d i n sertio n n e arer to

ge t her Th e g a s tr ocn em z a s muscle which m ay be t ake n as
’

.
,

an illustr atio n h as its origi n by t w o h e ads o n e fro m a t en

, ,

din o u s b an d exte n di n g fro m th e lo w er en d of the femur to

t h e ti b io fi b u la th e other by a n arro w te n do n which j oi n s
-

the t e ndo n of the triceps o n th e an t erior sid e o f the thigh .

At its lower extremi ty t h e muscle is i n serted by m e an s of a

stro n g te ndo n whic h p asses over th e an kle j oi n t an d spre a ds
out over the lower or pl an t ar surface o f t h e foo t Wh e n .

th e m uscle co n tracts it m ay pro duce two movem e nts I t

, .

m ay str aighte n ou t or exte n d the foot an d he n ce is spoke n

, ,

of as an ex tensor o f th at me mber I t m ay also be n d the .

leg u po n the thigh at th e k n e e a move me n t which is c alled ,

fl The g astro c n e mius i s desig nated t h ere fore

'

ex z oa
. as a n , ,

ex lea sa r of th e foot an d a f lex e r of the leg A muscl e .

which dr aws the limb posteriorly tow ard the lo n g axis of

th e body is c alled an aa cla clor o n e which pulls i t in th e
’
,

opposite directio n an a aa a ctor Th e n there are r e la lors

’
.
,

which c a use a limb to rot ate about i ts lo n g axis ; leva lo r s ,

which r aise a p art such as th e m uscles which r aise the lo w e r

,

j a w ; an d de r essors
p which
, p roduce th e opposite move me n t ,

such as th e dep r esso r m a n rlz b a la which lowers th e j aw Th e

'

.
,

ki nd of move me nt s a limb m ay m ak e is d epe n d e n t o n the

n ature of its j oi n ts and the n u mber an d att ach me n t s o f its

muscles ; an d the action s which the frog as a whole is cap

a ble o f p erfor m i ng are depe n de n t in a simil ar m an n er u po n

t h e org an iz a tio n of i ts skelet al an d muscul ar systems ,

although th e order an d combi n a tio n s o f the moveme n ts are

directed by i mpulses fro m th e c e n tr al n ervous syste m .

The actio n of muscles m ay b e w ell illustr ated by a study

248 TH E BIOLOGY OF TH E FROG CH A P . xrv

of so me of t h e m uscles of the h i n d leg I f th e ski n be .

stripp ed off fro m t h e leg an d t he frog l aid o n i ts b ack th e ,

followi n g mu scles m ay be see n o n th e ve n tr al side of the

t high

Th e sa r tor iu s a narrow m uscle h avi n g i ts origi n o n th e

,

ileum j us t in fro n t o f th e pubis I t crosses th e thigh o b

, .

liq u ely an d i s inserted by a te n do n a short dist an ce belo w

th e h e ad of t h e t ibi a Whe n it co n tr ac ts it flexes the leg o n
.
,

th e thigh an d pulls th e whole li mb forwa rd an d ve n tr ally .

Th e a a a a cler m ag n u s a l arge thick mu scle lyi n g behi n d

’ ’
,

th e s artorius which i t crosses at i ts lower e nd I t s o rigi n

, .

is fro m the pubis an d ischiu m an d it receives a sm all slip ,

w hich origi n a tes o n the te n do n of o n e o f th e h e ads of th e

sem ifen a in os a s I t is i n serted i n to th e dist al e nd o f the

’
.

fe mur I t s act io n i s to be n d t h e thigh ve n trally an d to pull

.

it an t eriorly or poste riorly a ccor di n g to th e positio n o f t h e

li mb Whe n th e th igh is pulled forward so th at a li n e c o n
.

n ec t ing the ce n ters of t h e origi n a n d i n sertio n of th e muscle

lies in fro n t of th e he a d of th e femu r the c o ntr actio n o f th e ,

a dducto r m ag n us h as th e effec t of m ovi ng th e li mb still

farther for w ard while pu lli ng i t ve n tr ally W h e n t he thigh .

is be n t b ack so th at the he ad of th e fe m ur l ies in fro n t o f a

li n e co n n ecti n g t h e ce nters of the origi n an d i n s e r tio n th e ,

co n tr actio n of th e muscl e h as the oppo sit e e ffec t Th e .

a cla a cf e r m ag n u s thus a cts as an adductor or an abductor

’

accordi n g t o circu mst an ces .

Th e a a a a cler lo ng a s a n arro w mu scle arisi n g fro m th e

’ ’

ve n tr al p art of th e ileum j ust dors al to the he ad o f th e

s artorius dist ally it j o i n s the a acla cz o r m ag n u s I t is p artly
’ ‘
.

covered by the s artorius but a sm all portio n o f it is exposed

,

alo n g th e pre axi al side of th at muscl e I t pulls the t high .

forw ard and ve n t r ally .

Th e t r iceps fem o r is a very l arge mu scle coveri n g the

,
 5
2 0 TH E BIOLOGY OF TH E F RO G CH A P .

whole fro n t of the thi gh I t arises by three h e ads Th e

. .

an terior h e ad ( cap a t a n lica m or cr u r a lis ) arises fro m

th e anteri or borde r of the ac et abulu m at the j u nctio n of the
ileu m an d pubis Th e middle he a d ( cap a l m ea ia m) is a
.
’

short flat muscle which ar ises fro m th e ve n tr al side o f th e

,

ileum n e ar th e middle an d j oi n s th e fasci a of th e r est o f th e

muscle at abou t th e proxi m al third o f the an t erior m argi n
o f the thigh The posterior he ad (cap a l p os lica m o a s z a s
.
,
‘

ex ler iz as g la fea s m a n us
, g ) a rises from the posterior e n d o f
th e c r est o f th e ileu m ; dist ally its fi bers j oi n those of th e
cr u r a lis or an terior h e a d . Th e whol e m uscle is i ns erted by
a very s tro n g te n do n whi ch p asses over the k n ee an d j oi n s

th e upp er en d o f t h e tibio fi b u la A te n di nous co nn ectio n

-

p asses also t o o n e h e ad o f th e g a s t r o cnem ius Th e actio n .

o f th e whol e muscle i s to exte n d the c r us an d d raw th e whole

l eg forward .

Th e g r a cilis m aj o r a l arge muscl e lyi n g alo n g th e poste

,

rior sid e o f th e ve n tr al su rface of the thigh I t arises by a .

sm all te n do n fr o m th e posterior m argi n of the ischiu m I t s .

d ist al te n do n d ivides i n to t w o p arts t h e o n e p assi n g be n e ath

,

th e te n do n o f the sa r lor ia s t o be i n serted i n to th e pre axi al

side o f t h e proxi m al en d of th e tibi a th e oth er p assi ng dors al
,

to the te n di n ous i n sertio n of th e semilen clirz osa s and j oi ni ng

th e posterior side of th e h e ad of the tibio fi b ula Ne ar t he
-
.

middle t he muscle is c r ossed by an obliqu e te n di n ous ih

scriptio n Th e g r a cilis m aj or pulls th e fe mur b ackw ar d and
.

ei t h er flexes or exte n ds th e crus acco r di n g to th e positio n

o f the l a tter in rel atio n to th e femur I f th e crus is flexed
.

so th at i t m akes an an gle o f less th an 9 0 with th e fe mu r °

,

t h e actio n of th e gr acilis is to flex i t s t ill mo r e I f how .

,

e ver t he crus i s p ar t ly exte n ded so t h a t i t m akes a c o n

,

side rab l gre ater a n gl e th an 0 wi t h the fe mur th e co n tr actio n

°
y 9 ,

of th e g r a cilis still fu r th e r exte nds it The diffe r e n ce o f the

.
xrv TH E M U S CU L A R S YS TE M 51
2

a ctio n of th e muscle u n d er th ese differe n t co n ditio n s dep e n d s

upo n th e ch ange in th e fulcr u m wh ich is b r ought abou t b y
the be n di n g of the leg .

The g r a cilis m ino r is a sle nd er mu scle which ar ises fro m

a t e n do n behi n d the i sch i a c symphysis D ist ally it j oi n s the .

te ndo n of i n sertio n of th e g r a cilis m aj or The muscle is .

also at t a ched to th e ski n of th e posterior side o f the thigh .

I t s actio n is si mil ar to th a t o f the g r acilis m aj or .

T h e two follo w i n g muscles appe ar o n th e dors al sid e of

the thigh : th e sem z m em or a n os a s a l arge mu scle lyi ng o n
’

th e posterio r side of the dors al surface o f th e thigh j ust

above th e gr acilis m aj or I t arises by a bro ad fleshy a t t ach
.

m e n t fro m the dors al h alf o f the pos terior m argi n o f the

ischium I t is i n serted by a short te n do n which p asses
.

be n e ath the te n do n of origi n o f th e ga s t r ocnem iu s i n to the

proxi m al end of the tibio fi b ula behi n d the k n ee There is
-
.

an oblique te n di n ous i n scriptio n r u n n i n g a cross this muscle

as in th e g r a cilis maj o r The semimem or a nos a s adducts th e

.

thigh o r pulls it b ackw ard and like the g r a cilis fl exes or

, , , ,

exte n d s th e l eg accordi n g to wh eth er i t is in a flexed o r an

exte n ded positio n .

The ileo fi oa la r is a sle n der m uscl e lyi ng be twee n the

-

s em imem or a nos a s an d th e posterior h e ad of th e t r icep s

fem or is I t arises fro m th e ileu m j ust behi n d th e posterior
.
,

e n d o f th e dors al crest an d it is i n serte d i n to the proxim al

,

en d of th e fibul a I t draws the thigh dors ally an d flexes th e

.

leg .

Th e semizemlirz osa s a sle n de r muscle which is cove r ed by

‘
,

th e g r acilis m aj or I t arises by two te n di n ous h e ads from

.

the ischium The ve n tr al he ad p asses be t wee n th e dors al

.

a n d ve n t r al h e ads of th e aa a a clo r m ag n u s an d a ffords

’ ’
a

poi n t o f a t t ach me n t for th e sm all third h e ad of th e l atter

muscle The two he ad s u n i te n e ar t h e mi ddle of the
.
25 2 TH E B IOLO G Y OF TH E FRO G CH AP .

thigh ; th e muscle is i n serte d by a n arrow te n do n i n to the

pre axi al sid e of the proxim al en d o f the tibi a I t adducts .

th e femur an d flexes th e leg .

The py r ifo r m is a sho r t sle n de r muscle exte n di n g from

, ,

th e ti p o f the urostyle to a sh ort dist an ce beyo n d the he ad

o f th e fe mu r I t lies bet w e e n t h e sem im em or a zzos a s an d
.

th e posterior h e ad o f th e t r icep s I t pulls the u r ostyle to .

o n e sid e an d dr aws t h e femur dors ally .

Th e iliacas ex ter m cs arises o n th e ou ter sid e o f th e do r

sal cres t of t h e ileu m fro m the a n terior third to wi t hi n a

short di st an c e fro m th e posterio r en d I t exte n d s b ack .

w ard p assi n g betwee n the middle an d posterio r h e ads o f

,

the tr iceps to b e i n serted o n th e pos t erior side o f th e h e ad

of t he fe mu r I t rot ates the femur forw ard
. .

The iliacu s in ter /t ics arises fro m th e ve n tr al bord er o f

th e ileu m a lit t le in fro n t o f th e an terior spi n e I n th e
, .

an terior positio n o f its bro a d o rigi n its fi be r s ru n belo w th e

lower m argi n o f the ileu m to be att ache d to th e m edi an sur

fac e I t s i n sertio n exte n ds fro m th e hi p j oi nt to about t h e
.

middle of t h e fe mu r This m u scle is bro ad a nd flat and

.
,

exte n ds bet w ee n th e middle an d an terior h e ads of the

t r icep s ve n tr al to the ilia ca s ex ter n a s I t d r aws th e thigh .

forw ard .

There are sever al sm alle r muscles arou n d th e h e ad of th e

fe m u r ; viz th e p ectineus o ota r a t or exter n a s o btu r a to r
.
, ,

in ter n a s ileo fem o r a lis ga a a r a t a s femo r is a n d ge mellzcs fo r

’
-
r
, , , < ,

a descri ptio n o f which th e stude n t is referred to E cker s ’

“A n ato my o f t h e F rog The muscles of th e l eg or crus

.
, ,

are as follows

Th e g a s t r ocnem iu s whi ch has bee n m e n tio n ed above i s

th e l arges t muscl e o f the leg I t arises by t w o h e ads the .
,

l arger o n e from a te n di n ous arch whi c h ex t e n ds from t h e

p osterio r sid e of t h e dist al e nd o f the fe mu r to t he h e ad
25 4 TH E BIOLO G Y OF TH E FRO G CH A P .

an terior side o f t h e l eg p artly covere d by t h e tioia lis a n ticu s

,

loizg a s I t arises by a sle n der t e n do n from th e dist al en d

.
,

o f th e femur I t i s i n serted d irectly u po n th e bo n e alo n g

.
,

t he an terior sid e of the t ibio fi b ula I t exte nd s th e leg -

. .

The t ioia lis a n tica s or eois is a short muscle lyi ng close

alo n gsid e o f the ex t e n sor cruris I t a rises fro m the dist al .

t h ird of t h e tibio fi b u la where it i s at t ach ed directly to th e

-

bo n e D ist ally i t e n ds in a te n do n which is i n serted in th e

.

proxim al e nd o f t h e tibi ale I t flexes the foot . .

There are n u merous sm all er muscl es for m ovi ng th e dif

feren t p arts o f t h e foot whose d escri ptio n w e sh all o mi t
, .

Whe n a muscle works in co mp an y with oth ers i ts actio n ,

is o ft e n qui t e differe n t fro m th at produc ed wh e n i t a c ts

alo n e This fact has bee n well illustr a ted by D r L o mb ard
.
l
.
,

w h o h as worked ou t th e double ac t io n o f th e muscles of th e

frog s leg in a very t h orough m ann er The tr iceps fem o r is

’
.
,

for i n st an ce wh e n worki ng alo n e acts as an ex t e n sor of the

,

leg bu t si n ce i t pulls th e thigh forw ard at th e s a m e ti m e ,

i t c auses the fle xo r muscles o n the b ack o f th e thigh t o

becom e te n se and thereby i n direc t ly bri ngs about a flexio n
,

of the leg The muscles o f th e frog s leg are so arranged

.
’

th a t an y force which pulls the leg forw ard c auses the crus to
flex ag ai n s t th e thigh an d th e foot to flex ag ai nst the crus ,

a n d a ny force which pulls the leg b ackw ard exte n ds the crus

an d foot The move me n ts o f the frog s leg in j u m p i n g an d ’

.

swi mmi n g co n sist m ai n ly in alter n ately bri n gi n g the leg up

ag ai n st th e sid e o f the body foldi ng the v arious p a rts ,

toge ther an d pulli n g it b ackward an d exte n di n g or u n fold

,

ing i ts differ e n t p arts A n y muscl e such as th e ilia ca s

.
,

in ter a ct s w hich pulls th e fe mu r b ackw ard bri ngs about a

, ,

flexio n o f th e crus through the te n sio n produced o n th e

1 Co ntr ibu t i s on to Me dic l R s ch d dic t d

a e ea r ,
e a e to V . C
. V a u g h an ,

p . 26 0 , 1 9 03 .
xrv TH E M U S CU L A R SYST E M 25 5

flexo rs o f t he b ack of t h e thigh A flexio n of th e cru s

.

stretches the tioia lis a n tica s lor g as o n th e an terior sid e of

th e crus and th ereby bri ngs about a flexio n o f t h e foot I f .

a mu scle pulls th e femur b ackw ard the t r icep s fem o r is is put

,

o n the stretch : this te n ds to str aigh t e n the leg th e t en ,

de n ey beco mi n g stro n ger the farther b ack t h e t high i s pulled .

E xte n di n g th e leg c auses th e g a s t r ociz em izcs to be p u t o n a

str ai n an d th e te n sio n is i n cre as ed t hrough th e pull o n the
,

te n do n t his mu scle rec eives fro m th e lower en d o f the

tr icep s . Through the pull o n th e g a s tr o cnem iu s t h e foot is
ex t e n ded O ther muscles are of course brough t i n t o pl ay
.

in this proce ss bu t wh at has be e n s aid w ill illustr ate t h e

,

pri n ciple o n which co ordi n at io n of m o vem e n ts em p loyed in

ju mpi n g or swi mmi n g is e ffec t ed Th e rel ative i n t e n si t y of
.

the n ervous i mpulses dis t ributed to t h e v arious muscles is

an i mport an t factor in de t ermi n i n g t h e ki n d o f movem e n t

th e limb will m ake bu t the b asi s for th e u n ity of ac t io n o f

,

th e p ar ts in th e ordi n ary movem e n ts of the limb lies in the

structur al arrangem e n t o f the bo n es and muscles .

As an exte n d ed tre atm e n t o f th e rem ai n i ng portio n s of

th e muscul ar syste m li es beyo n d th e scope of this w ork o n ly ,

a few of the more n otewor t hy muscles W ill be d escribed On .

th e ve n tr al sid e of t h e body t h e l arge r ect u s a oaomm is ex ’

te nds fro m th e pubis to which its fibers co n verge be hi n d to

, ,

th e ster n u m in fro n t I ts t w o h alves are sep ar ated in t he

.

m iddle li n e by the linea a loa an d there are five co nn ec t ive

,

t issu e sept a ( in scr i

p t iorzes t ea clia m whi ch cross it tr an sversely
)
an d divide it i n t o segme n t s .

The ooliga a s ex ter /ill s i s a l arge muscl e coveri n g m os t of

t h e sid es of the body I t origi na tes o n th e sides o f the
.

ileu m an d dors al fasci ae above an d is i n s er t ed i n t o th e sides

,

o f t h e rec t us abdomi n is I t s fibers exte n d obli q u ely upw ar d

.

an d fo rw ard .
2
5 6 TH E BIOLOGY OF TH E FROG CH A P .

Th e t r a ns oer s u s is a b r o ad muscle lyi n g be n e at h th e ex

ter nal obliqu e an d formi n g the i nn ermos t muscul ar l ayer o f
th e body w all I t s fibers ru n for the most p art tr an sversely
. .

A n terio r ly so m e o f its fibers are i nserted i n to the esoph agus

an d clos ely overlie th e peric ardiu m ; other fibers a tt ach t o

t h e cor acoi d an d xiphister n u m th e pos t erior portio n of the

m uscle is i n serted i n to a flat te nd o n which exte n ds dors ally
to th e r ect u s a oaom in is to th e m id ve ntral li n e All o f these
’
-
.

mu scl es h ave the ge n e ral effect o f co n tr acti ng the body

c avi ty .

The cu ta neous p ector is a p aired m uscle lyi ng o n th e ven

,

t r al side of th e an terior p art of th e body P osteri orly it is

.

a tt ached to th e body w all an d i t i s i n serte d a nte r iorly i n to

,

th e S ki n betwee n the fore legs .

Th e pector a lis maj or a l arge mu scle o n eith e r side o f th e

,

an terio r p art o f t h e bod y I t is composed o f three p arts an

.
,

abdomi nal por t io n a risi n g fro m the sides of th e a n teri or h alf

of the rec tus abdomi n is a m iddle po r tio n arisi n g from t he

,

ster n u m and xiphister n u m an d an an terior portio n ari si n g

,

fro m th e cor acoi d an d ep ic o rac o ids All three p arts are in

.

se rt ed n e a r togeth er o n th e ve n tr al crest of the h ume r us .

The s u bma x illa ry muscle ex te n ds tr ansvers ely ac r oss th e

floor of the bucc al c avi ty b et w ee n th e two r a mi o f the lower
,

j aw I.t r ais es the floor o f t he bu cc al c a v it y in respir atio n .

Th e s u bmen ta lis is a s m all muscl e lyi n g in th e an t erior

an gle o f th e lower j aw .I t s fibers ru n tran sversely an d b v,

th eir co n tractio n r aise the tip of the j aw an d thereby bri n g

abou t th e closure of th e n ares in respir a tio n .

The s u b/zy oia a sm all muscl e arisi n g fro m th e an terio r

’
,

cor n er of the hyoid n e ar the skull an d j oi n i n g ve n tr ally th e

posterior m argi n o f t h e s u bm ax illa ry I t r aises t h e hyoid
. .

The follo w i n g muscl es are at t ach ed to t h e hyoid and t ake

p art in th e move me nts o f respir at io n Th e g en iony oicl ,
25 8 TH E B IOLO G Y OF TH E FROG CH A P .

C H A PT E R X V
TH E CI RC L A U TO RY S YS TE M
TH E pri n cip al fu n c tio n s o f th e circul a tory syste m are to
c arry foo d m at eri al a n d oxyge n t o all p ar t s of th e body and ,

to re move t h e c arbo n dioxide an d o ther w aste produ c t s o f

tissu e m e t abolism t o t h e org an s w h ere they ar e elimi n at ed .

These fu n ctio n s are disch arge d by me an s o f two fluids the ,

blood an d t h e lymph We sh all d escribe t h e blood firs t

. .

The B lood — Th e blood of th e frog co nsis ts of a flui d th e

.
,

la s m a in which t h ere occur n u m erous fre e c ells or co r u s

p , p ,

cles . The corpus cle s are of three ki n ds : r ea co rp u scles ’

zo /
z ite co r
p u scles
(l )
eu cocy tes , an d s
p in dle cells .

The re d corpuscles are elliptic al in ou tli n e an d h ave an ov al

n ucleus in t h e ce n ter . Whe n see n o n edge t hey appe ar
fl at t e n ed an d sligh t ly bulgi n g in t h e ce n ter where t h e n ucleus
lies Th e n e arly tr an sp are n t cy t opl as m of th e cell co nt ai n s
.

a l a rge perce n t age o f hemoglobi n which give s it a yello w ish

,

color B lood appe ars red o n ly whe n a l ayer o f co n sider able

.

thick ness is see n .

The whi t e corpuscles u n like the r ed o n es v ary exceed

, ,

ingly both in size an d form The ou t li n e of th e w hi t e cor

.

p u sc l e u n dergoes ch a n ges much like those o f a n Amoeb a .

The ch an ges are sl o w b u t they m ay e asily be S h o w n by

,

m aki n g ou t li n e ske t ches o f th e corpuscle at i nt erv als o f o n e

or more mi nutes The cy t opl asm o f t h e w h i t e corp uscles is
.

n e a rly colorless but it oft e n co n t ai n s gr an ules o f v a riou s sizes

,

a n d s t ai n i n g re a c t io n s The form of t h e n ucleus v ari es

.
xv TH E CI RC L A U T O RY S YS TE M 25 9

gre atly in di ffere n t cells I n some c ases especi ally in th e

.
,

sm aller leucocytes i t is ne arly sph eric al I n other cells i t is

,

very ir r egul ar in ou tli n e an d m ay be deeply co n stricted in

seve ral pl aces an d n o t i n frequ e n t ly i t m ay b e divided i n to
,

two or mor e disti n ct n uclei I n the sm allest leucocytes the .

a mou n t o f cyto
pl asm is rel atively
sm all an d forms a
n arrow irregul ar
,

e n velope arou n d
th e spheric al nu
c leu s .The cyto
pl asm of th e l arger
c o r p u s c l e s is
r e l a t i v e ly m uch
g re ater in amou n t .

B y virtu e of
t h e i r a m oe b o i d
m ov e m en t s th e
whi t e corpuscles
h ave th e po w er
of i nd epe n de n t
F I G 7 —B l d c p uscl s f t h f g
. I . oo o or e o e ro . a, , 6 ,

locomotio n an d d bl d c p uscl s ;
re oo d a y u g st g s ;
or e a an o n a e

m tu c uscl — s i dl c lls dif

,

t h ey eve n p ass i f
’
c, p ; a / p re or e a z n e e n er

t st g s ; ly s t g ; s m htlt
,

’
en a e a , ea r a e e, a o ew a a er
through th e deli s t g ; fi g 1 ty p ic l p i dl c lls ; i—l f ms f
a e 2, a S n e e or o

l uc cy t s ; i y ly s t g ; j ld s t g ;
, ,

c ate w alls of th e e o e ve r ea r a e an o er a e

é c ll i th l b d ucl us ; 1 c ll i th f u ucl i
, ,

e w o e n e e w o r n e
c apill aries i n t o th e .
,
.

( A ft D kh y n )
,

er e u ze .

sp ac es betwee n
th e other cell s of the body They are n o t co n fi n ed there .
,

fore to th e blood v ess els like t h e red c ells b u t m ay b e

, ,

fou n d in almost every p art o f t h e organ ism An d they oft e n .

p ass o u t of wh at are s t rictly spe aki n g t he li mi ts o f the body

, ,

i nto the m outh and ali me n t ar y c anal .

260 TH E BI OL O GY OF TH E FRO G CH A P .

O ne i m p or t an t prope r ty o f th e whi te cells is th ei r powe r

of ingu lfi n g sm all bodies which are t ake n in m u ch as
,

an A m oeb a t a kes in its food B ac t eri a are devoured in

.

this w ay an d th e leucocytes thus afford th e body a m e as

,

ure o f protectio n ag ai n st th ese organis ms which are c o n ,

s t ant ly bei n g i n trodu ced i n to th e syst em in o n e w ay

or another and might i f u n check ed be produ ctiv e of
, , ,

serious i f n o t fat al effec t s A n irrit atio n set u p in an y regi o n

.

c au ses leucocytes to be attra ct ed to t h e spo t in l arge nu m

bers The i n trodu c t i o n of b acteri a i n to any p art of th e body
.

is followed by the i nv asio n of th a t p art by leu cocytes an d it ,

freque n tly h app e n s th at th e b acteri a are devou r ed by th ese

cells before th ey g ai n a stro n g foothold I t is very prob able
.

th a t o n e impo r t an t fa ctor whi ch c auses th e movem e n t s o f th e

leucocytes in such c as es is th e prese n ce o f subst an ces give n
off by the b acteri a which exercise a c hemo t a c t ic effe ct u po n
thes e w an deri n g cells c ausi n g the m to co ngreg ate about the
,

ce n ter o f di ffusio n Th e ch em ot acti c i n flue n ce of su ch sub

.

st an ces i s show n by the follo w i n g e xpe r i me n ts first performed

by M ass art I f a fi n e c apill ary tube s e aled at o n e e n d be
.

filled wi t h a culture flui d co n t ai ni n g th e b acteriu m S tap hy

lo coccu s py og en es a lou s an d i n tro duc ed i n to o n e of the l a rge
,

lymphs s paces u n d er th e ski n i t will b e fou n d in th e cou rse

,

o f t en or twelve hours t h at a swarm of leucocytes h ave m ad e

their e n t r an ce i n to the op e n e n d o f the tube I f a si mil ar .

tube filled o nly wi t h the culture m ediu m be i nt r oduced n o ,

leucocytes will be fou n d to e n te r i t Th e subst an ces pro

.

duc ed by the b a cteri a are app are n tly th e c ause of th e i n v a

sio n of th e tube by the w an deri n g cells .

B esides protecti n g the body agai ns t th e germs of disease ,

th e whi t e cells t ake p art in the re mov al o f tissu e wh ich h as

’

b ecome broke n dow n an d n o lo n ger o f service to the org an

i sm I n the dege n e ratio n o f th e t ail o f th e t adpole th e
.
262 TH E B I O L O GY O F TH E FROG CH AP
.

h and i f blood b e dr aw n i n to a vessel w h ose sid es are sm e ared

,

with oil it m ay be kept fr o m clotti n g for a comp ar atively

,

lo n g ti m e C old gre a tly ch ecks clotti n g th e process bei n g

.
,

del aye d alm ost i n defi n itely if blood is kept n e ar th e poi n t

of freezi n g I f blood be h e ated to n e ar 1 0 0 C its po w e r
.
°
.
,

o f clo t ti n g i s destroyed Th e form a tio n o f clot i s d e

.

pe n de n t in som e w ay u po n th e prese n ce o f c alciu m s al ts in

the pl asm a ; for i f t hese are preci pit ated out clo tti ng m ay be ,

preve n ted Th e proc ess of clotti n g is du e to t h e form atio n

.

o f fibri n fro m a subst an ce c alled fi b rino gen whi ch exists in a

st ate o f solutio n in th e pl asm a Th e ch an ge is du e like .
,

the form atio n of ch eese in m ilk to a fe r m e n t which tr an s ,

fo r ms the soluble co m pou n d i n to an i n soluble form The ,

form atio n of clot h as the fu n ctio n of preve n ti ng i n defi n ite

bleedi ng a fter an i nj ury h as b ee n received the co n t ac t w i t h ,

foreig n bodies c a usi n g th e clot to for m an d th ereby ch eck

ing fur t h er loss o f blood .

The L ymph — T h e lymph i s a colorless fluid devoid of ,

r ed corp uscles but fur nished wi th n u m erous leucocy t es I t s

,
.

pl asm a co agul ates but n o t qui te so re adily as th at o f blood

,
.

The P r oduct ion of New Corpu scles —The corpuscles of .

th e blood aft er fu n ctio n i n g for a cert ai n ti m e di e an d are

, ,

repl ac ed by n ew cells The process of r ege n er atio n of n ew

.

corpuscles does n o t t ake pl ac e u n i formly throughout the

ye ar as in higher an i m als bu t is m ai nly co n fi n ed to th e
,

spri n g an d e arly su m mer D uri n g the l at e fall wi n t er an d

.
, ,

e arly spri ng t here is a period of i n ac t ivi t y in th e produc tio n

of n ew cells O n ly aft er th e breedi n g p eri o d whe n the frog
.
,

begi n s to t ake food an d store u p n u t ri me n t in t h e body

,

1 8 th ere a r a pid rege n er at io n o f the blood The process .

re aches i ts m axi mu m in o n e or two weeks a ft er which the ,

produc t io n o f n e w corpuscles suffers a gr adu al dimi n uti o n

u ntil fall .
XV TH E U
CI RC L A T O RY S YS TE M 26 3

AS B izzozero Torre h ave show n t h e pri n cip al se at of

an d ,

th e form atio n of n ew corpuscles i s in the m arrow of t he

bo n es M arquis fou n d t h at the m arrow u n dergoes periodic
.

ch a nges correspo ndi n g to th e ch an ges in the blood In .

the l at e spri n g an d e arly sum me r duri n g th e period o f ,

most active re n ew al of blood cells the m arrow assumes a ,

lymphoid ch arac t er ; duri n g the su m mer i t gr adu ally ac

cu mul at es fat and ret ai n s a fatty ch ar ac te r duri ng th e fall
,

an d wi n t er . D uri ng th e spri n g t h e fat is more or less,

co mple t ely r esorbed an d th e lymph o i d cells wh ich are

, ,

co n cer n ed in the form atio n of ne w co r pu scles u n dergo ,

r api d mul t iplic atio n .

While m ost of the corpuscles both red an d white arise , ,

l
in th e m arrow of the bo n es it is c ert ai n th at both ki nds of
,

c ells are also produced by th e divisio n of pre exis t i ng cells in

the circul at i ng blood Thi s is n ot ably t h e c ase in th e white
.

corpuscl es ; an d divisio n s o f the you ng st ages o f t h e red

corpuscles h ave bee n wi t n essed by a n u mber o f observers .

T h e spi n dle cells fro m which red corpuscl es are d erived

u n d ergo i ndirect divisio n a t le ast aft er they h ave acquired
,

h emoglobi n . Their tr an sform atio n i n to r ed corpuscl es ,

accordi n g to Neu m an n occurs o n ly in the spri ng after th ey

, ,

acquire hemoglobi n bu t they are prese nt in th e blood

, ,

alth ough in less n u mbers du r i n g th e whole ye ar

,
Th e .

m ature red corpuscles d o n o t divide The v ery you n gest .

st age of the r ed cells is represe n ted accordi ng to Neum an n , ,

De khu yz e n an d oth ers by sm all leucocytes w i t h a rel atively

, ,

l arge n ucleus Tr ansitio nal st ages occur betwee n th ese an d

.

th e spi ndle cells an d betwee n t h e l at t er an d t h e you n g red

,

cells I n th eir you n g st ages the red cells h ave a more or

.
,

less circul ar bu t irregul ar form comp ar at ively li t t le h em o

,

globi n an d a rel at ively l arge irregul ar n ucleus H owever

,
.

differe n t the v arious fo r ms of corpuscles m ay be in thei r

2 64 TH E BIOLO G Y O F TH E FROG CH A P .

a dul t co nditio n th ey ari se from cells of a v ery si mil ar

, ,

al t hough p erh aps n o t id e n tic al s t ru cture

Th e S t r u ct u re of t h e H eart —Th e h e art of the frog is

.
,

situ ated in t h e an terio r p art o f th e body c avity ve n tr al to th e ,

live r . I t lies wi thi n a sac the p er ica r diu m whose c av

,
ity ,

is completely cut off fro m the c o lo m although origi nally e

co n ti n uous wi th i t in e arly develop me n t Th e p eric ardiu m .

is co mposed of two l ayers th e p a r ieta l formi n g th e ou t e r

, ,

w all of t h e sac a n d th e viscer a l w hich closely i n ves t s t h e

, ,

he ar t The t w o l ayer s are co n ti nu ou s p assi n g i n to e ach

.
,

other in t h e regio n o f th e tru n cus arteriosus below an d t h e

an terio r ve n ae c a vae abo ve Th e rel atio n is such as wou ld be
.

p roduced i f th e he art were pus hed i n to th e hollow p eric ar

di al sac fro m in fro n t al though as a m atter o f fact it is n o t
,

brought about in this w ay .

Whe n th e p eri c ardiu m is ope n ed o n the ve n tr al si de th e ,

follo w i n g p ar t s com e i n to vie w ; ( I ) The co n ic al ven tr icle ,

wi th its ap ex poi n t i n g b ackw ard this p art o f the h e ar t has

very t hick mu scul ar w alls a n d ap pe ars p aler th an the rest .

2 ) The a u r icles li e i mmedi a t ely in fro n t of th e ve n tricle

( .

Th e auricles are thi n w alled and are sep ara ted fro m e ach
-

othe r i n t er nally by a septu m but from th e outside th ey ,

prese n t Only a fai n t i n dic at io n o f t his divisio n ; t h ey are -

cle arly sep a rated fro m the ve n tricles by th e cor o n ary sulcu s .

(3) The o u lou s cor a is lyi

’
n
,
g in fro n t of t h e righ t side of t h e

ve n t ricle ; it is a th icke n ed muscul ar tube exte n di ng o b ,

liq u ely across th e righ t auricle an teriorly it i s co n ti n ued

i n to th e t hi n n er w alled t r u ncu s a r ter iosu s fro m which i t is

-

dem arc at ed by a sulcus ( 4 ) The tr u n cu s a r ter iosu s is

.

so mewh a t n arro w e r th a n th e bulbus ; i t soo n divides i n t o

t w o di vergi n g tru n ks w h ich give rise to three arteri es t h e
, ,

co m m o n ca r o tia th e a o r ta a n d t h e p u lm o cu ta n eo u s
’ -

.
, ,

O n th e dors al sid e of the h e a rt is the tri an gul ar thi n ,

26 6 TH E BIOLOGY OF TH E FRO G CH AP .

Th e i n ter nal s t ru cture of th e he art prese nts a complic at ed

an d be au tifully ad ap t ed m ech an ism for propelli n g t h e blood

al w ays in o n e directio n a n d keepi n g the pure an d i mpu re

blood s ep ara ted B y r em ovi n g th e ve n tral w all of the

.

au r icles ve n tricl es an d bulbus m ost of the fe a tures of th e

, , ,

i n ter n al st r ucture m ay be
expos e d to vi ew Th e .

in ter a u r icu la r sep t u m is

a il
so si tu ated th at the righ t
auricl e is mu ch l arger
th an th e left I n th e .

ri g ht auricl e close to th e ,

septu m i s the l a rge ,

s in u a u r icu la r
-

a p er t u r e,
through which blo o d e n
ters from the si nu s ve n o
sus I t is a tr an sverse .

ov al ope n i n g gu arded by
v alvul ar lips o n the an t e
FI G 7 3 — H t s n f m th d s l ear ee ro e or a
n ( I poster ior 5 1 des
. .

sid i t h th si us sus p n d p
.

e w e n
n o
ve n o
r a
o e e u ;
.

a o, ao r t ic t u k ; r igh t u icl ;
n au , r which preve n t th e blood
a r e au

l ft u icl ;
e a r e c t id t u k ; p
ca , aro r
t h a t has e n ter ed fro m th e
n . cu ,

p 1m
u 0 c_

mn a t u k , 15
eo u s p c v l
r n 7 . a, re a a

v in ;
e pt .
p st c
c, l v i ; p p l si n u s fro m bei n g forced
o ava e n . v, u

m o n ary i ve n si us
,
sus ;
s .v ,
b ack
n
ag a i n
ven o
whe n the v,

ve n r t icl ; e si u u icul v l v s
va n -
a r ar a e .

( A f ter H s )
ow e .
a u ricles co n t r a ct The .

left auricle receives bl o od

from the pul mo n ary vei n through a sm all ope n i n g n e a r t h e
s eptu m sligh t ly an t erior to t h e si n u au ricul ar aper t ure there -

is no v alve at t his poi n t but si n c e the vei n perfor a tes th e w all

,

obliqu ely th e pressu re c a used by the co n tr ac t io n o f the auricle

,

serve s to close the o pe n i n g an d thus pre ve n t s t h e b ack w ard

flo w of t h e blood B o t h auricles e mp t y i n t o t h e ve n t ricle
.

by a l arge ope n i n g t h e a u r icu lo ven tr icu la r ap er tu re which i s

,
-

,
xv TH E CI RC L A U TO RY S YS TE M 26 7

divided by the i n ter auric ul ar septum This ope n i n g is .

gu arded by four v alves two l arge o n es o n t he dors al an d

,

ve n tr al edges and a sm all v alve at ei t her en d ; sm all fibers

,

exte n d fro m the ve n tricul ar w all to be i nser t e d i n to the fre e

edges of th e v alves ; they preve n t the edges o f t he v alves
fro m bei ng turn ed b ack i n t o the auricles w h e n th e ve n tricle
co nt racts and t hus ke ep the blood fro m flowi n g fro m the
,

ve n tricle i n to the auricles The ve n t ricl e possesses in

.

addi t io n to th e ce n tr al c avity a t its b ase a n u mbe r of fi ssu re

like ch ambers in the thick muscul ar w all ; t h ese ch amber s

are sep ar a ted fro m e a ch o ther by muscul ar p a r t i t io n s a n d

e x t e n d outwardly n e arly to th e p eriph ery o f t h e ve n tricle ;

they receive a l arge p ar t of th e blood th at p asses t hrough
t h e he art and h ave t h e i mpor t an t fu n ctio n as will be see n
, ,

l at er o f preve n t i ng t h e mixi n g of th e blood th at comes in

,

fro m the two auricl es .

Th e ope n i n g fro m th e v e n tricl e i n to t h e bulbus cordis is

gu arded by three sem ilu na r va lves They are in the form .

of pockets ope n an teriorly whose w alls m ay be pressed

, ,

dow n wh e n blood is p assi n g out o f th e ve n tricle but wh e n ,

blood t e nds to p ass the oth er w ay they fill an d prev e n t its ,

re t ur n I n fro n t o f th ese v alves is a p eculi ar an d i mport an t

.

s t ructure k n ow n as the sp ir a l va lve I t co n sists of a lo ngi .

t u din al fold att ached alo n g th e dors al w all o f th e bulbus it s ,

ve n tr al edge lyi n g fre e At it s pos t erio r en d it is at t ach ed

.

t o the left side of the bulbus n e ar th e ope n i n g i n to the ven

,

t riele . I t p as se s obliquely across the bulbus an d wide n s ,

ou t at its an terio r en d i n to a cup like v alve Two sm aller -

.

v alves occur at the s am e level I n fro n t o f th e se v alves i s

.

the u n p aired portio n of the tru n cus which is p artly divid ed ,

by a ridge ex t e n di n g fro m th e poi n t o f u n io n of th e two

bran ch es to the middle of t h e an t erior e n l argeme n t o f the
spir al v alve E ach of the two br anches o f the tru n cus is
.
268 TH E BIOLO G Y OF TH E FRO G CH AP .

div id ed by two
s e pta i n t o thre e
co mp ar tm e n t s Th e .

an teri or co mp art
m e n ts whic h l e ad ,

to the com m on
c a r o t i d ar t e r i e s ,

bo t h e n ter th e u n
p aired divisio n of
t h e t r u n cus to the

righ t of th e septu m .

The middle co m
p ar t m en t s which ,

le ad to th e aor t a ,

ope n i n to the u n
p ai red portio n of
th e truncu s on
either sid e o f th e
s eptu m .

Th e posterior
co m partments ,

w h i c h are c o n
8 0 t in u e d i n t o t h e
F I G 7 4 — Di g m f t h t i l sys t m f th p u l m o c u t a n e o u s
a ra o e ar e r a e o e

f g s f m th l d ti c
. .
-

”
ro ee n t ro e ve n ra Sl e . ao ao r

ch ; igh t u icl ; l ft u icl ; o arches j oi n e ach

, ,

’ ”
ar au r a r e on , e a r e r , ,

b ch i l t y ; c t id ; g l c t id oth e r an d ope n by
,

ra a ar e r c c, a ro a aro

—
.

gl d ; il c mm ili c ;
,

an c ol m on a cc
e , c oe ra c o es

ic ; c li c ; cu t us ; l co m mo perture
.
,

' a n a
e nt er cce oe a cu , a n eo c ao,
.

d sl f m l ; g g s t ic ; l i n to the bulbu s
,

or a t ; f ao r a in , e o ra a r z,

h id l ; l p h p t ic ; l y p ig s t ic
,

h mm o rr o a i e a z e a r o

ve sic l ; k kid y ; l li gu l ; lg l ft lu g ;
a ne
cordis
,
b eh i nd
n
the a
,

”
e n

i m s t ic ; s i m s
, , ,

m t a n er o r p t e en e r v a lves a t i ts an t e o er o r e en

cci p i t l ; p p c t ic ; p
,
’
t i ;
er c

ulm c
o c,

us t ulm y
o

l rior
a
en d
c ,
an re a . cu ,
.

p o u ta n e o ; pu p o n ar ; r e , re n a ;
Th e Art er ies
-

sci t i c ; p s p l ic t u cus i
,

s c, a s en ; r a, tr n ar ter o .

sus ; t t s is t b l (A f s)
,

s, e t v , ve r e ra . H
ter ow e .

The a r ter ies, or

2 70 TH E BIOLO G Y OF TH E F ROG CH A P .

the middle li n e to th e b a s e o f th e S kull I n fro n t o f th e .

l ateral process of the p ar asph e noid bo n e i t gives off the

a la tine artery which cou rses forw ard alo n g th e roo f o f
p ,

the mouth ; a li ttle farther forw ard the cer ebr a l ca r otid
arises e n teri n g th e skull in th e regio n o f th e orbit an d sup
,

plyi n g the brai n Th e third br an ch th e op /z t/i a lm ic p asses

.
, ,

forward an d supplies th e eye an d so me of th e n eighbori n g

p ar t s .

Th e seco n d b r anch of th e tru n cus arte r iosus o r sy s temic ,

a r c/ z p ass es ou t w a rd an d th e n dors ally arou n d th e alim e n

,

t ary c an al m eeti n g i ts fellow of the o p posite side to form

,

th e do r sa l a or ta Ne a r i ts origi n e ach sys t e mic arch gives

.

o ff a sm all la ry ng ea l artery wh ich supplies the l ary n x and

,

cert ai n m uscles of th e hyoid The sm all esop h ag ea l arter .

i es are give n off at about t h e level o f th e seco n d ver t ebra ;

they are distributed to th e dors al side o f th e e so ph agus .

The occip ito ver tebr a l artery arises S lightly beyo n d o r so me

-

t i mes in com mo n with t h e l ast an d p asses forw ard a cross

,

t h e t r an svers e process o f t h e seco n d v ertebr a ; i t the n

divides i n to two br an ches wh ich ru n above the tran sverse

p r ocesses of the vertebrae n e ar th e c e n tra .

Th e posterior bran ch or ver teor a l artery exte n ds b ack

, ,

w ard alo n g t h e spi n al colu m n Th e an terior br an ch or .

,

occipit a l a rtery ru n s forw ard to the h e ad givi n g br an ch es

, ,

to t h e upper and lower j aw orbit an d n ose , ,

.

Th e l arge su ocla via n artery arises i m medi ately behi n d th e

occipi t o vertebr al and p asses l ater ally givi n g br an ches t o the
-

, ,

should er an d body w all an d the n as th e br achi al ar tery

, , ,

supplyi n g t h e arm .

The t w o syste mic arch es u n ite at abou t the level of th e

six t h ver t ebr a to form th e do r sa l a or ta which proceeds b a ck ,

w a rd be n e a t h the vertebr al colum n At t h e poi n t o f mee t .

ing of t h e t w o syste mic tru n ks bu t usu ally more o n t h e left ,

xv TH E U L AT O RY
CI RC S YS TE M 27 1

th an o n right the l arge ccelia co m esen ter ic ar t ery is give n off

,
-

which supplies the alim e n t ary c an al an d i ts appe ndages I t .

divides i n to an an terior br an ch or ccelia c artery an d a pos

, ,

t erio r b ran ch th e a n ter io r m esen t er ic artery

,
The former .
,

after givi n g off th e le ft g a s tric artery wh ich goes to th e left

,

sid e of the stom ach divides i n to the r zgnt g as tr ic artery sup

, ,

plyi n g th e r igh t sid e of th e stom ach an d p an cre as an d the ,

lz ep a tic whi ch a fter givi n g a br a n ch to the an terio r br an ch

, ,

of t h e p an cre as is distribute d to the l iver The a n ter io r

, .

mesen ter ic a rtery supplies th e sm all i n testi n e S plee n clo ac a , , ,

an d an t erior portio n of the rectu m .

Th e u r in og en ita l arteries are four to six sm all ar teri es whi ch

a rise from the ve n tr al sid e of the a or t a an d are distributed ,

to t h e reproductive o rg an s fat bodi es an d kid n eys They

, ,
.

v ary gre a tly in t h ei r m od e o f origi n bu t typic ally they arise

,

by very short m edi an t r u nks whic h divide i n to r igh t an d l eft

br an ches .

Th e lu moa r arteries are sm all vessels o ne to four in n u m ,

ber o n ei ther sid e which arise from the dors al sid e o f the
,

aort a an d ar e dis t ributed to th e body w all .

T he p oster io r m esen ter ic artery i s a sm all v essel give n off

fro m n e ar the posterior en d of th e aor t a I t su ppli es the .

p o sterior por t io n of the rectu m an d in t he fe m al e t h e , ,

m ed i an dors al w all of the u terus .

At its posterior en d th e dors al ao r t a divid es i n to two l arge ‘

iliac ar t eries wh ich are distributed m ai n ly t o th e hi n d limbs

,
.

A S hor t dis t an ce behi n d t h e bifurc atio n e ach ili ac artery gives

o ff a br an ch whic h divides i n to a n en a s tr ic a rtery supply ,

in g t h e v e n tr a l body w all an d a r ect o —

, ves ica l artery to the

rec t u m and bl adder A s m all ar t ery arisi ng from t h e ili ac

.

close to the above suppli es t h e se mi nal vesicles in th e m ale ,

an d t h e l at er al p art o f t h e u t erus in t h e fem ale .

A sho r t dist an ce beyon d the fo r egoi ng t h e femo r a l artery

2 72 TH E BIOLO G Y OF TH E FROG CH A P .

is give n off t o t h e ski n an d mu scles o f th e an teri or p art o f

t h e t high D is t al to t h e origi n of t h e fe mor al artery th e ili ac
.

artery i s co n ti n u ed as t h e scia tic whi ch p asses ou t of th e ,

body c avity alo n g w i t h the sci at i c n erve j ust behi n d the

, ,

p osterior e n d o f th e cre st of the ileu m an d supplies the hi n d ,

limbs .

The third br an ch of t h e tru ncus the p u lmo cu ta n eo u s ,

-

a rt ery p a sse s ou tw a rd a n d dors ally an d give s off p o steriorly

, ,

th e p u fin o n a ry artery to th e lu n gs ; it i s t he n co n ti nu ed as
t h e g r ea t cu ta n eo u s artery which p a sses u pw a rd an d fo rw ard
, ,

dividi n g behi n d t h e t ym p an ic membr an e in t o three br an ches

e a u r icu la r is w hic h supplies t h e t ymp an u m an d gives
( )
r t h ,

br an ch es to t h e t hymus lo w er j aw ph ary n x an d hyoid an d

, , , ,

a n as t om oses wi t h br an ch es of th e occi p i t al an d i n ter n al

c aro t id ; ( 2 ) the do r sa lis whi ch i s m ai n ly dis t ribu ted t o th e
,

S ki n of the b ack an d
(3) th e la ter a lis which p a sses outw ard ,

an d is exte n sively distribu ted t o t h e S ki n of the S id e o f

th e body Wi th t h e excep t io n of a few mi n or br an ch es th e
.

dis t ribu t io n o f the pul mo cut an eous artery i s such as t o bri ng

-

th e blood i n t o regi o n s where i t m ay u n d ergo oxyge n atio n .

The bl o od which do es n o t go to the lu n gs p asses i n to th e

gre at cu t an e o us artery the l argest br an ch es of which are dis
,

tributed t o t h e ski n w h ere m uch of the respir a tio n of the frog

,

t akes pl ace th e bucc al c avi t y an d ph ary n x which are also ,

r espi rat o ry org an s receive bl o od from t h e s am e source

,
.

Th e V ein s —W i t h t h e excep t io n of th e blood comi n g

.

fro m t h e lu n gs all o f th e blood i s re t ur n ed to th e he ar t

,

th rough t h e t h ree l arge ve n ou s t ru n ks w hich e n t er th e si n us

ve n osu s The t w o an terior ven m c av ae which e n ter t h e
.
,

a n t erior an g les of the si n us a re e ach fo rm ed through t h e

,

j u n c t io n o f t hree br an ches t h e ex te n a l j ug u la r t h e in n o m i
,
r
,

n a te an d t h e s u ocla via n
,
Th e mos t a n t eri o r o f t h ese t h e
.
,

ex t er n al j ugul ar p asses forw ard w h ere it receives bran ch es

, ,
2 74 TH E BIOLOGY OF TH E FROG CH A P .

from th e to ngue hyoid t hyroid an d pseudo t hyroid gl an ds

, , ,
-

a n d floo r o f th e mou t h .

The in n o m in a te vei n o r seco n d bran c h o f t h e an terior,

ve n a c av a p asses l ater ally an d divides i n t o th e in ter n a l

,

j g
u u la r which
,
rec eives blood fro m th e br ai n a n d v arious
p arts of t h e h e ad an d th e s u oscap u la r w hich ru n s alo n g th e
, ,

oute r side of the fore li mb an d rec eives br an ch es fro m th e

S houlder .

Th e s u bcla via n or pos t erior br an ch of th e an terior ve n a

,

c av a is formed by th e co n flue n ce of t h e or a c/z ia l w hich is

, ,
.

d is t ributed to the fore li mb and t h e l arge cu ta neo u s vei n , ,

which is ext e n sively dis t ribu ted over th e side o f the b o dy

a n d he a d The l at ter retur n s mos t of the blood c arried by
.

the cu t an eous artery .

The l arge p os ter io r ven a ca va ar ises betwee n th e kid n eys ,

an d ru n s fo rw ard ve n tr a l to th e dors al aor t a to t h e t a peri n g .

posterior e n d o f t h e si n us ve nosus Ne ar t h e l at ter i t .

receives the l arge but very sh ort lz ep a tic vei n s fro m t h e

liver .

Th e r en a l vei n s four to six in n u mber o n e ach S id e le ad

, ,

fro m t h e kid n eys i n to the pos t erior ve na c av a th e vei n s

fro m t h e reproduc t ive org an s (sp er m a tic or o va r ia n accord
"
,
!

in g to t h e sex of t h e a ni m al ) le ad ei t h er direc t ly i n to th e
ve na c av a or first i n to th e re n al vei ns Th e ve na c av a also .

receives o n e or two br an ches o n either sid e fro m the fat

bodies .

Th e blood from th e hi n d limbs do es n o t empty di r ectly

i n to th e p o sterior ve n a c av a as in the high er vertebr ates b u t ,

it is forced to p ass th rough a seco n d sys t e m o f c apill aries

be fore re achi n g th at vessel Two l arge vei n s co nvey t h e blood .

fro m t h e h i nd legs the scia tic which ru n s alo n g t he post

, ,

a xi al sid e o f th e thigh arid t h e fem o r a l which courses alo n g

, ,

the dors al an d an terio r S ide o f t h e thigh an d p asses u n d e r

XV TH E CI RC L AU T O RY SYS TE M 275

the il iac bo n e i nt o the body c avi ty These two v ei n s are

.

co n n ected close to the hi p j oi n t by th e tr a nsver se ilia c vei n ,

which p asses dors ally to the fe mur an d e n ters the body c avi ty
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276 TH E B IOLOGY OF TH E FRO G CH AP .

behi n d th e crest o f the il eu m Th e femor al vei n br an ches .

in fr o n t of the b ase of t h e thigh i n t o t w o p ar t s o n e o f w h ich ,

p asses ve n trally and j oi ns its fellow o f th e opposi te sid e t o

a n Za / é

l in l
‘

FIG .
77 .
—T h h
ic p t l sys t m sh i g it l t i s t th s t m ch
e ep at or a e ow n s re a on o e o a ,

i es i p c s d li b ch f m h i bd mi l
,

’
nt t ne, an rea an ver a ra n ro t e a n t er o r a o na

p t l i ; t é i bd mi l i ; d du d um ;
, .
,

t th
o e or a ve n a rz a a n te r o r a o na ve n a, o en

y t s m ; g g s ic i ; mb] g ll bl dd ; l Z igh t
.
,
'
d
’ ”
a ,
a rt e r o a e a tr ve n o a a er v v ,
r
b

d l f l b s f h li s p c i ly ; p t l i ; p p c s ; t
, , ,

an e t o e o t e ve r re e t ve or a ve n e, an rea s .

s m ch ( A f H s )
to a . t er ow e .

form t h e a n ter io r a é a e m z n a / vei n w hi ch ru n s for w ard in

’
’
,

t h e middl e o f t h e v e n t r a l body w all ; t h e other br an ch t h e ,

sses for rd dors lly j oi s th e sci

'

ex ter n a l z lz a e
'

p a w a a n d ,
a a n d n , ,

at ic vei n t o form th e ea m m o n M a c o r r en a l p o r ta l vei n ,

w hich ru n s for w ard alo n g t h e ou t er m a rgi n of th e kid n eys ,

i n t o t h e subs t an ce of w hich it se n ds it s br an ch es Th e re nal .

p o r t al recei v es t h e a orro Za m ba r vei n fro m the body w all and

’
-

, ,
2 78 TH E BIOLO GY OF TH E F RO G CH AP .

sep ar ate B ru cke s observ atio n s were exte n ded an d in most

.
’

po i n ts co n firmed by S ab a tier in 1 8 7 3 Th e i nterpre t atio n .

of the l at t e r author h as be e n fo llowed by G au p p in his

“
r ece n t r evisio n of E cker s A n ato mie d es F rosches
’
.
”

Wh e n the auricl es co n tract th e blood fro m th e l eft au ricle

, ,

which h as com e in from t h e pulmo n ary vei n an d is th erefore

oxyge nat ed is forced i n to the left sid e of the ve n tricle while
, ,

t he i mpure blood fro m the righ t auricle which co m es -

through the si n u s ve n osus pours i n to th e right side and

,

m iddle portio n o f th e ve n tricle The blood fro m th ese .

d iffere n t sources is preve n ted fro m becomi n g mixed by

bei n g received i n to t h e slit like ch ambers in the ve ntricul ar
-

w all D uri n g th e co n t r a c tio n o f the v e n tricle th e im pure

.

blood lyi n g n e ar the ope n i n g of th e bulbus naturally p asses

ou t firs t w h ile t he pure pulmo n ary blood fro m t he le ft side i s
,

forced out o n ly t ow ard th e close of t he ve n tricul ar c o nt rac

tio n Wh e n the ve n t ricle fi rst co n tracts th e w all of t h e bul
.
,

bus cordis i s rel axed an d th e impure bloo d flows freely ove r

,

the edge o f th e spi ral v alve i n to the le ft co mp artm e n t ,

whe n ce i t i s fre e t o issu e i n to th e pulmo cu t an e ous arch es -

t hro u gh t h eir co mm o n ope n i n g No w th e blood is u n d e r

.

less pressure in th e pul mo cu t an eous arches th an in th e -

others bec ause i ts rou t e i s shorter an d there are no i mped i

,

m e n ts to it s flo w I n th e c arotid arches th e blood m eets

.

wi t h a p arti al obstru ctio n in th e c aro ti d gl and and at t h e ,

outer e n ds o f the sys t emic arc hes th ere is a sm all v alve

( v a lv u l a p arad o xic a ) w hich a
,
lso te n ds t o ret a rd its flow .

The blood firs t issui n g fro m the h e ar t t akes th e li n e of le ast

resis t an ce n am ely t h e pul mo cu t an eous arches an d is
,
-

forc ed t hrough the firs t two p airs o f arch es o nly wh e n it has

n o e asier ave n u e of esc a pe To w ard the close of the c o n
.

tr a c t io n of t h e ve n t ricl e whe n t h e pure blood is p assi ng out

, ,

there is a co ntr actio n o f th e b ul b us c o rdtls Thi s bri ngs the .

XV TH E U
CI RC L ATO RY S YS TE M 2 79

w all o f th e bulbu s ag ai n st th e fre e edge o f th e spir al v alve

an d preve n ts t h e blood fro m fl o wi n g o v er i n t o t h e left or pul

m o nary sid e o f this divisi o n o f the he art Th e blood is pre

.

ve n ted fro m access to this sid e an teriorly by v alves so there ,

i s n ow n o course ope n to it bu t through th e c arotid an d

systemi c ar ch es S i n ce t h e co m mo n ope n i n g o f the pulmo
.

cut an eous arches lies behi n d th e v alves at the an terior e n d

o f th e bulb us it can receive n o blood wh e n th e c o m m u nic a
,

tio n betwe e n th e two sides o f th e bulbus i s cu t off I n this .

w a
y th e impu r e blood firs t se n t ou t o f the h e art goes m a i nly

to th e lu n gs an d ski n where i t i s purified whil e th e purer

, ,

blood p assi n g out tow ard th e clos e of th e co n tr actio n of th e

h e art is se n t to the v arious o t her p arts o f the body .

Th e h e ar t o f th e frog m ay be at for hours or u n der fav , ,

o rab le co n d i tio n s eve n for d ays a ft er i t h as bee n removed

, ,

from the body E ve n isol at ed p arts o f th e h e art such as

.
,

th e si n u s ve n osu s au ricl es or ve n tricl e m ay co n ti n ue be at

, , ,

ing although n o t wi th t h e s a m e rhythm

, I f the he art i s
.

r e mov ed so as to l e av e t h e si nus v e n osus withi n the body ,

t h e auricles an d ve n tricle be at wi t h a r ate l ess t h an th e n o r

m al but the si n us co n ti n ues to be a t with n e arly th e s am e

,

rhyth m as b efore I f the si n u s i s removed with th e r est of

.

the he art the be ati n g of the whole he art is more r api d th an

,

t h at of th e auricles an d ve n tricl e whe n re moved alo n e It .

i s app are n tly t h e si nu s ve n osu s which sets the rhyth m for th e

be ati n g of the oth er p arts o f th e h e art Aft er th e he art h as
.

c e ase d to be at spo n t an eously i t m ay be c aused to resum e

i ts activity by t h e applic a tio n of a sti mulu s .

Circulat ion in t h e W eb of t he F oot —The w eb of th e .

frog s foot affords a cl assic al obj ect for th e stu dy of th e

’

c apill ary circul a tio n I t m ay e asily be prep ared for observ a

.

tio n wi t h t he microscop e by tyi n g t h e frog dow n to a sm all

pi ece of bo ar d an d spre ad i ng its toes ap art so t h at the web
,
28 0 TH E BIOLOGY OF TH E FROG CH A P
.

is stre tched acr o ss a n otch o r hole th rough which ligh t m ay

be p assed fr o m below Th e to es m ay be h eld in positio n
.

by sm all piec es of thre a d tied t o th e tips an d faste n ed t o

their o t her e n ds to th e bo ard .

I n a web t hus prep ared th e blood m ay be see n flowi n g

r apidly in the s m all vei ns an d a r t eries an d m ore slo w ly in
,

the c apill ari es Th e red corpu scles w ill be fou n d t o beco m e

.

elo ngated an d n arrowed as th ey t h re ad their w ay slo w ly

through th e sm all c apill aries Th e leucocytes o fte n cree p
.

sl o wly alo n g t h e w alls o f th e vessels an d m ay be s ee n to

,

s t o p frequ e n t ly an d so m e t i mes to migra te through t he c ap il

,

l ary w alls I n the ar t eri es a puls a tio n due to th e be ati n g o f

.

t h e he art m ay be observed the c aliber o f the arteries oft e n

ch an ges owi n g to the co n tr actio n o f the mu scl e fibers o f
,

t heir w alls .

The c apill ary circul atio n m ay also be e asily studied in th e

t ail o f t h e t ad pol e .

Th e L y mph at ic S y st em — Th e lymph atic syste m o f the

frog i s rem ark able o n accou n t of th e ab un d an ce and l arge
size o f t h e lymph sp aces in v arious p ar t s of the body There .

are n o w ell de fi n e d lymph a t ic vessels such as occur in th e

-

m am m als ; th e lymph flo w s in irregul ar sp aces b e t w ee n an d

w i t hi n t h e differe n t o rg a n s ; t h e l a rger sp aces a r e li n ed by

fl a t t e n ed e n d o theli al cells but are e n tirely d evoid of a

,

muscul ar co at an d u su ally also o f a li n i n g of a co nn ective

, , ,

t issue.

Th e subcu t an eous lymph sp aces are esp eci ally well devel
oped t hey are sep a r ated fro m e ach o nly o ther by the nar
ro w sept a o f co nn ective tissu e by which the ski n is here
an d t here a t t ach ed to th e u n d erlyi n g muscles O n e of th e
.

l arges t of t h e lymph sp aces wi t hi n t h e body is the subver t e

br al l y m p h si n us or eis ter n a nz ag n a wh ich exte n ds above
, ,

m o s t o f t h e dors al sid e o f t h e b o dy c avi t y .

28 2 TH E BIOLOGY OF TH E FROG CH AP .

from t h e lymph s acs Th e lymph h e arts are furn ish ed wi th .

a muscul ar co a t composed o f a n etwork o f bu n dles of s t ri ated

mu scle fibers .

Th e be at i n g of the lymph h e ar ts m ay re adily be observed

in a rece n tly killed frog O ft e n th e puls atio n s of the pos .

t erio r lymp h h e arts m ay be see n be n e ath t h e ski n but th ey ,

are e a sily demo n str able in a very s atisfactory m ann e r by

re movi n g th e i n teg u m e n t o n e ach sid e o f the en d o f th e

u rostyle Th ei r puls a tio n s h ave n o rel atio n to those o f th e
.

h e art no r is th ere u niso n be t wee n th e beats o f the lymph

,

he arts o n the two si des of the body .

RE F E RE NC E S
B ru ck e , E . B e t ra i g e z ur ve r gl ich de en en A n at o mi e un d P hysi
o lo g ie de s G e fasssys t e m s, I . U b di
e er e Me ch i k d K isl u f s b i
an es re a e e

de n A m p hibi en . De n k skr . d k Ak d
. . a . W iss m t h iss Cl B d 3. a -
w . .
, .
,

W i en , 1 852 .

Dek h uy sen M C , . . Ub e er das B lu t de r A m p h ibi en. V e rb . An at .

,

G es .
, 6 V er s .
, 1 8 92 .

F u ch s , E i g K tm d F schbl u t s d d F sch
. B e t ra z ur en ss es ro e un er ro

lym p h V i ch s A chi B d 7 8 7 7
e . r ow
’
r v, . 1, 1 .

G l J B
a u e, b ch t u g tib di f bl s E l m t d F sch
. eo a n en er e ar o en e en e es ro

blu t s A ch A t u P hys p hys A b t h 8 8

e . r . na . . .
, . 1 0 .

M call m a S t udi s th B l d f A m p h ibi T s C di

u . e on e oo o a . ran . an a an

I st V l
n , 89 o . 2, 1 2.

M rq i C D a K ch m k d A m p hibi i d
u s, . hi as no en ar er en n en v e rs c e,

d e n enJ h sz i t I g D iss D p t
a re 89 e en . n au . or a , 1 2 .

N m n E H m t l g i h S t udi
eu a n, U b di B lu t bildu g
. a a o o sc e en, 1 . e er e n

vo n F osch V i ch s A chi B k 4 3 8 9 6
r
’

en . r o w
’
r v, . 1 , 1 .

S b ti r A
a Et u d s
a l c u
e t l ci cul t i
,
c t l d sl
. e su r e oe r e a r a on e n ra e an a

sé i d V t éb és A S i N t
r e es T 8 8 73
er r . nn . c . a . . 1 ,
1 .

T ii iik L Di T h il g d
r , th B lu t z l l
. b i A m p h ibi e e un er ro en e en e en .

A ch f mi k A t B d 3
r . . 888 . na .
, . 2, 1 .
xvr TH E NE RVO U S S YS TE M 28 3

CH A PT E R XV I

TH E NE RVO U S S YS TE M
TH E frog has th e powe r n o t o n ly of perfo r mi ng a l arge
n umber of complic ated m ove me n ts but of ad apti n g its
,

a c t io n s to th e v arious eleme n ts of its e n viro n me n t Th e

.

i n iti atio n an d co nt rol of th ese m oveme n ts are d ep e n de n t

upo n the r eceptio n of s t imuli eithe r from w ithi n or without
th e org an is m an d the t ran sfer of t h e i mpulses thus arisi ng
t o the muscl es w hich by t heir co n t rac t io n bri n g about th e

required ac t io n s Whe n the frog w ithdr aws i ts foo t whe n

.

it is irri t at ed or s naps at a movi ng i n sec t i t is p erformi n g an

, ,

ac t of an a d aptive n ature in r espo n se to an exter n al st im u

lus I t is evide n t th a t th e actio n s of the frog in rel atio n to

.

exter n al stimuli and the co o rdi n atio n of activities goi n g on

in differe n t p arts o f the org an is m n ecessit ate so m e hi ghly
speci alized m e an s for th e tr an sfer an d directio n of i mpulses ,

a n d it is with these fu nctio n s th a t the n ervous sys tem is

esp eci ally an d pri m arily co n cer n ed B ut th e n ervous sys

.

te m h as an other i mport an t fu n ctio n i n asmuch as i t a ffords

,

the m e an s for th e accu mul atio n of th e effects o f experi e n ces

w hereby th e an i m al is e n abled to profit by its form er beh av

ior an d modify its co n duct to sui t n ew situ atio n s This .

l at te r powe r forms the b asis of i n t ellige n c e a faculty r ath er

,

feebly d evelop ed in t h e frog it is tru e bu t as we sh all see

, , ,

l ater a n o t u n i mport an t eleme n t in th e life of t h e an im al

,
.

Th e n ervous system h as o ft e n bee n comp are d to a system

of tel egraph w ires by m e an s of which a n y o n e p art of a
c o u ntry m ay b e put i n to com mu n ic atio n wit h any o ther
'
28 4 TH E BIOLOGY OF TH E FRO G CH AP .

p art The n erves correspo n d to t h e wires an d th e g angli a

.
,

t o th e ce n t r al s t ati o n s wh ere m ess ages m ay be tr an sferred

fro m o ne li n e to an oth er All p ar t s of th e body are sup

.

plied with n erves wh ich are co nn ec ted with th e ce n tr al

n ervous syste m an d through th is ch an n el co nn ectio n s m a
, y
be est ablish ed betwe e n any t w o or more p arts o f th e organ
ism I n this w ay th ere i s re ndered possible th e c o Ordin a
.

tio n o f move me n ts in differe n t p a rts o f th e body an d t h e ,

abili t y of t h e org an is m to ac t as a whole in rel a tio n to

exter n al thi n gs.

The n ervous syste m is com posed o f t hree r at her closely

associ at ed divisio n s : th e t er ebr o sp in a l co n si sti n g o f th e
-

spi n al co r d an d brai n ; th e p er ipher a l co nsi sti ng of the

,

spi nal an d cr an i al n erves an d t he sy mp a t/ietie .

The S p in al Cor d —The spi n al c o rd of the frog is sh ort

.

an d so m ewh at fl at te n ed I t prese n ts two e n l arge me n ts o n e

.
,

in the b r a chi al regio n where t he l arge n erves to the fore

,

li mbs are give n off and o ne farther b ack where the l arge
, ,

n erves origi n a t e wh ich supply the hi n d legs B ehi nd the

.

pos terior e n l arge me n t the cord t apers to a n a rrow t hre ad ,

the fi ln m ter m in a le which exte n ds i nt o the urostyle At

,
.

it s an t erior e n d t h e cord wide n s gradu ally i n to th e m edu lla

oblo ng a ta ,
t h e posterior d ivisio n o f t h e br a i n B o t h the .

dors al an d th e ve n tr al sid es of the cord are d ivided by a

medi an fi ssure At the sides o f t h e cord th e r oots of t h e
.

sp in a l n er ves are give n off e ach n erve a rises fro m a d ors al

a n d a ve n tr al root whic h combi n e j ust a ft er they em erge

from t h e vertebr al c an al through the i nt ervert ebr al forami n a .

The roots o f t h e pos t eri or spi nal n erves are much elo n g ated ,

i n as much as th e shor t e n i n g o f t h e co rd bri ngs t heir origi n

far in fro n t of the vertebr ae t o wh ich t h ey co rrespo n d ; the
b u ndle o f ro ots thus form ed t oge t her wit h the fi ll/772 ter mi
,

n a le i s k n ow n a s t h e ea a a a e u in a
’
, q .
28 6 TH E BIOLO G Y OF TH E F RO G CH A P
.

i nto two l amell ae This l aye r i s very closely applied to th e .

ce n tr al n ervous system and is co nt i n u e d i nto v a rious fissu r es ,

of the brai n and the v e n tr al fi ssure o f th e spi n al cord

, .

A cross sectio n o f the c o rd sho w s i t t o be co mposed

m ai nly of g anglio n cells an d n erve fibers The ce n t r al p ar t .

o f th e cord i s form ed
of gray m at t er which
l C Z f .

co n sis t s c hie fly of gan

gli o n cells and n o n
m e d u l l at e d n e r v e s .

Ne ar th e ce n ter o f th e
gr ay m ass is a sm all
c anal the ca n a lis een ,

F I G 8 —C ss s c t i
. 0 . th ugh t h v t {r d /is li n ed by a si ngle
ro e on ro e er e

b l c lum
ra d sp i
o l c d sh i n g t h l ayer O f e l t h eh al cells
n , an na or ow e
,

or ig i f t h p i l
n o s c t l
e S na p n e rve . a c, e n ra
.

c l;
an a c t um ; d] d s l fissu ; This c anal is the re m
en , en r ,
or a re

a
’
m
. du m t ; ra d s l t f
n an t
a er
Of the lu
a r
'

me
.

n
or a ro o o

g y m tt ; g g ngli n f
, ,

n er v ; gm e ra a er n, a o o

d s l t; u l c h ; p n u l formed by t h e clo sl n g
,

or a ro o n a,. ne ra ar n s .
,
e ra

s p i ; p m p i m t ( th f c li o ver O f th e edges o f
ne .
,
a a er e re ere n e ne

sh uld s t p t t h m gi f t h c d ) ;
o o a e ar n o e or

t n v
. ert u k ; T p e t r s s p c ss ;
n
th e m r
edull ary
r g ro o v e
ra n v e r e ro e

l fissu hi m
,

v
f. v
,
n t e ; ra t t t duri n g developme n t ;
re w an , w e a e r.

( A ft H o ) at its an terio r en d i t
er w es '

w i d e n s ou t i n to th e ve n tricle s o f th e br ai n .

At th e sides th e gr ay m atter is produced both dors ally

a nd ve n tr ally i n to the dors al a n d ve n tr al eor n n a or lz o r n s .

The gr ay m atter o n the two sides o f t h e cord is co n n ected

bo t h above an d below t he ce n tr al c an al by m e an s o f t h e
do r sa l an d ven tr a l g r ay eo m m iss n r es w hi ch co n sist chie fly ,

of no n medull ated n erve fibers Just below th e ve n t r al gr ay

-

c ommissure is a co n spi cuous obliqu e crossi n g of medull a ted

fi b ers in th e whi te m at t er th e ven t r a l zo /z ite eo m m issn r e ,

.

B elo w th e w hi te commissure is th e ven tr a l fiss u r e which se p a ,

r at es t h e righ t a nd left c o lu m n s o f white m atter F ro m th e .

XV I TH E NE RVO U S S YS TE M 28 7

sh allow zlor sa l fiss u r e t h ere exte n ds a n arrow septu m as far

as th e dors al gr a y com missu re Th e n ervous elem e n t s of .

th e cord are bou n d togeth er by stell a te n eurogli a cells an d

by processes wh ich arise fro m th e t aperi n g ou te r e n ds of

FIG . 81 .
—Di ag ra m
s p i l c d sh i g th p ths t k by
of us
th e na or ow n e a a en n e rvo

im p uls s T h di c t i f t h im p uls s is i dic t d by

e . e re s on o e e n a e a rrow . c c,.

c l c l ; l c ll t l fib s ;
e n t ra an a l c ll i th
co c b l c t x;
o a e ra er c eor
. e n e ere ra or e

g sm ll c b l c ll ; c lls i d s l h f g y m tt
, ,

’
c a er e re ra i
e a c, . e n or a o rn o ra a er ; a r,

d s l t ; g g gli f d s l g gli c ll i d s l
,

or a ro o t ; g an on o or a ro o . c, an on e n or a

g gli ; g g y m tt ; M muscl ; m c ll i m dull h

,

an on an , ra a er e c, e n e a o

fib ; S ski ; f s s y fib ; p s p i l
.
,

l g t ; mf m
on a a . o to r er n s en or er s . e, na

c d; c lls i lh f g m l t f
, ,

or v c,
. e ; n t ve n t ra o rn o r ay a t t er v r.
,
v en ra ro o o

n e ; rv e m hi t m
w .
(Af ,
P k w d P k e ) att e r . ter ar e r an ar er .

th e cells li n i n g the c e n tr al c anal th ese processes b ran ch

repe atedly an d some of th em ex t e n d to the pe r iphe r y o f the
,

cord .

The whi te m at ter o f th e cord i s composed m ai nly of

m edull ated fibers M ost o f these ru n lo ngi t udi nally I so . .

l at ed ganglio n cells appe ar b u t t here seem s to be n o regu ,

larit y in their dis t ributio n S t r an ds of gr ay m at ter l argely .

,

epe n dym a fibers r adi ate fro m the ce n tral p art of t h e cord
,

to th e outer surface .

The cells of th e gr ay m atte r give off processes by m e an s

28 8 TH E BIOLOGY OF TH E FROG CH A P .

of w hich co n n ec t i o n s becom e est ablished betwee n di ffere n t

p arts of t h e cord I n t h e br o ad ve n tr al cor nu a th ere are
.

several g an glio n cells of u n usu al siz e from which processes

arise which form th e axis cyli n d ers o f the fibers o f th e ve n

tr al roots of th e spi n al n erves o t h er proc esses fro m these

cells cross to th e o p posi te sid e of t he cord in t h e ve n tra l
whi te co m missure and s till o t h er processes br an ch irregu
,

larly in bo t h th e gr ay an d whi t e m at ter o f t h e s a me sid e .

S c a tt ered abou t thr o ugh m ost o f th e gr ay subst an ce are the

co m missur al c ells wh ich give off axi s cyli nder processes
which cross to t h e o pposi t e si de o f t h e cord in t h e v e n t r al
gray comm issure an d th e n give o ff br an ch es which ru n in
t h e white m at t er bo th an teriorly an d pos t eriorly ; p ro t o plas

mic processe s are also give n o ff wh ich co nn ec t wi t h simil ar

processes from o t her cells in t h e gr ay m at ter in th e s am e
side O th er cells give off axis cyli n d er processes which ru n
.
,

in bo t h d irectio n s in the whi te m at t er o f the s a m e sid e of

the cord S till o t h er c ells o ccur wh o s e axis cyli n der pro
.

cesses divid e th e o n e bra n ch goi n g i n to t h e w hi t e m atter o f

,

th e s am e sid e o f t h e cord t h e o the r crossi n g through th e

,

ve n tral gr ay com missure t o th e whi t e m atte r of th e opposi te

side F i n ally th ere are n u m erous c ells wh ose proc esses do
.

n o t e n ter t h e whi t e m at ter ,

bu t br an ch an d co n n ec t wi t h
cells in t h e gr ay m a t t e r o f t h e s a me o r th e o p p o si t e side
.

A cross sec t io n t hrough a regi o n where t h e spi nal n erves

are give n off sho w s t h e fibers o f the dors al r o o t p assi ng
t hrough t h e dorso l a ter al portio n of t h e w hi t e m a tter to e n t er
-

th e gr ay subst an ce in a n a rro w bu n dl e M o st o f th e fibers

‘

o f th e dors al roots are processes o f c ells lyi n g in th e spi nal

gan glio n E ach fiber as it e n t ers th e cord gives off br an ches
.

w hich ru n in opposi t e direc t i o n s . C o n n ec t io n s are m ade

wi t h pr o cesses o f t h e l arge c ells which su p ply t h e ve n t ral or
mo t or roo t s of the n erves as well as with th e cells of the gr ay
290 TH E BIOLO G Y OF TH E FROG CH A P .

which are distribu ted to the

fore li mb and m uscles of the
should er .

Th e t/z ir a p air of n erves ’

,

a ft er givi n g a br an ch to th e

br achi al plexus suppli es th e ,

an t erior p art o f th e exter n al

obliqu e and t ran sversus mu s

cles an d gives som e twigs to
the ski n .

The f o u r tlz fif tlz and , ,

s ixt/ z n erves are sm all an d

ar e distribu ted m ai n ly to th e

ski n an d muscles of th e w all

of th e abdom e n .

The seven t/z ezgnt/z an d , ,

n in t/ z n erves p ass al most

directly b ackw ard an d an as
to mose with e ach other to
form the lu m oo sa cr a l or -

s cia t ic
p lex us Th e s eve n th .

n erve before i t e n t ers th e ,

plexus gives o ff t h e ileo ,

lzyp og a s tr ic n erve w hich is ,

FIG 8 —
. S pi2l v s d sym p
. na
distributed
n er e
to th e muscles
an a

t h t ic sys t m
e f th f g t h eigh t O f th e abdom e n The cml
o e ro e r .

si d s f om b l O ly t h r a lis n erve is give n O ff fro m

,

e ee n r e ow . n e

v t l b n ch s f t h s p i l
e n ra ra e o e
.

na

s sh
n e r ve S ym p t h ic sys m i bl ck
ow n I— X spi
a l et s; A te n a . ,
na n erve o,

sys mic ch f t ; o p l b chi l p l xus ; C c lc us b di s

.

te ar o ao r a r ra a e ,
a a reo o e ,

u d h s p i l g gli ; D A d s l t ; f m f m l
. ,

a ro n t e na an a ; o, or a ao r a e ,
e o ra n e r ve

y; sci i c l p l sci t ic p l xus ; S E skull ;

.

[ l A i1i
.
,
ac a r t er s e, at n e r ve ; so a e ,

s p l ch ic ms ic y ; Sy sym p h ic c d ;
.
,

lan n or c oe ia c o -
e enter a rt e r at et or

c mmissu s b t sym p h ic d s p i l s ; Sy g
, ,

tSj r. c
o re e w een at et an na n e r ve

sym p th t ic g gli ; U t u s yl ; V V c t f t b ; Vg
, ,

a e an a s ro t e 1 9 e n ra o ve r e rae
, , ,

va gus ( F m P rk
n e rve. d P k ) ro a er a n ar er .
xvr TH E NE RVO U S S YS TE M 29 1

the plexus ve n tr al to the posterior por t io n of the ileum ;

it is distributed t o th e muscles o f the abdome n an d ski n of
th e an terio r p art o f th e thigh The l argest n erve comi n g
.

fro m the plexus is the scia tic w hich emerges from th e ,

body c avi t y j ust behi n d the po sterior en d of t h e crest o f the

ileu m p assi n g bet w e e n the pyriformis and posteri o r he ad of
,

t he triceps exte n sor muscle an d ex t e n di ng do w n th e b ack

of th e thigh ; a short dist an ce proxi m al to th e k n ee i t

divi des i n t o th e tibialis n erve an d t h e p er o neu s Th e .

former exte n ds alo ng t h e pos terior or flexo r sid e of the

leg and e n erv at es th e g ast roc n em ius tibi alis pos t icu s an d
, , ,

n u merous m uscles of the pl an t a r s u rfa c e of the foo t The .

pero n eus ru n s u n der t h e te n do n o f the triceps an d exte nds

alo n g t h e exte n sor surface of the crus givi n g br an ches to ,
.

the p ero n eu s muscle th e t ibi alis an ticu s an d the muscles o n

, ,

the ex t e n sor surface o f th e foo t .

The ten t/z n erve wi t h a br an ch fro m th e nin tlz forms th e

is c/z io—coccyg ea l p lex u s fro m w hich br a n ches are give n off to
,

t h e bl add er clo ac a oviducts an d pos t erio r lym ph h e arts

, , , .

The te n t h n erves are of sm all size an d em erge from sm all

fo ra mi na in t h e sides o f t h e uros t yl e n e ar t he an terior en d .

A n eleve nt h spi n al n er ve some t im es occurs Wh e n prese n t .

it emerges fro m th e urostyle behi n d th e op e n i ng for th e

te n th and j oi n s the ileo coccyge al plexus
-
All o f t h e .

plexuses are subj ect to co nsid er abl e v ari atio n in differe nt

i n dividu als .

Th e B rain — The br ai n i s composed o f t h e followi n g

p arts t ake n in order from behi n d fo rw ard : th e medu lla
oolo ng a t a
,
or lz in l or a in ; th e cer eoellu m ; the m irt or a in ;
z - -

the tlza la men cep /z a lo n ; an d the fo r e or a in whic h co n sists o f

-

th e cer eor al nem isp lzer es an d olfa ctory looes .

Th e medull a oblo n ga t a is formed by a wide n i n g of th e an

t erio r end of th e spi nal co r d O n it s dors al side is situ ated
.
F IG 83 — l}ra in
f g A d s l sid ; B o f l s i d C l f sid D i
ro or a t ic l e ve n t ra e ; e t e ; n v er a

l g i ud i l s c i h ug h h middl co c b l um ; L H c b l h mis p h s ;
. .
, , , ,

on t na e t on t ro t e e e re e l er e re ra e e re

s i ch id l xus c m missu s c c ti g
.
.
, ,

/ pl
c z ix1 d / pl
ante r o r,
p t an
p ;
c z m x2 o e r o r, o ro e co o re o n ne n

igh t d l f h l s f th b i ; C C c u c b i ; D d i c p h l h l
. . ,

th e r an e t a ve o e ra n r r ra ere r z, en e a on, or t a a

M f m fM ; i i qu duc t f S yl ius ;
.
,

m ph l
e n ce ; f a on or o ra en i f
o o n ro te r, o r a e o v n un

ol m dull bl g lf c y l b Op t ] p l b ; p t
.
, ,

d b l m ; Al
i u u ea o
’
O lj Il e a o on ata ; o a to r o e ; O t o e o v,

p ic sicl ; p p i l b dy ; p t p i ui y b dy ; Sp s p i l c d ; hi d
.
, , ,

v3 ,
’
o t ve e in , n ea o i t tar o ca na or t r ven

f u th icl l—X c i l s ; 0p S p fi s t d s c d p i l
,
.
,

t i l ;
r c e v4, o r ve n t r e ; ra n a n e r ve 1 z r an e on S na

(F m P k dP k s Z Ol g f G p p d O sb )
, , ,

n rv
'
e es. ro ar er a n ar er O o v , a te r au an o rn .

29 2
294 TH E B IO L O GY O F TH E F RO G CH AP.

growths in th e mid dors al li n e th e p a r ap hysis a v ascul ar

-

, ,

outpocke t i n g o f the epithelium of the roof an d a short dis ,

t an ce behi n d the l atter th e ep ip hy sis a hollow thi n w alle d

, , ,
-

c anal which t ermi n ates bli ndly a t i ts an terior e n d A sm all .

a r iet a l n er ve ru n s alo n g th e dors al surfac e of t h e e piphysis

p
an d exte n ds forw ar d ove r th e p a r aphysis an d th e n p asses

through the s agittal suture o f the skull to end in the brow

spot Th e epiphysis whic h origi n ally w as co n ti n uou s wi t h
.
,

th e brow spot becomes co n stricted off fro m i t in early l arv a l

,

life O n th e ve n tral side o f the th al a me n ceph alo n is the

.

op tic clz ia s m a ,
o r c rossi ng of th e n erves which go to the
eyes . I n th e frog all of th e fibers cross to the opposite
side .

j ust behi n d th e o p t ic c h ias m a is

'

the inf un dib u la r lo be ,

a fl atte n ed bilob ed structure e m argi n ate p o steriorly and

,

divi ded by a medi an lo ngi tud i n al groove I t is formed of .

n ervous tissu e an d co n t ai n s a c avi ty which i s co n ti n uou s

w i t h th e t hi rd ve n t ricle .

The byp op /zy s is cer eb r i lies b ehi n d an d p artly covered by

the i n fu n d ibul ar lob e I t is com posed of an an terior and
.

posterior p art ; th e fo rm er is divided i n t o a medi an and t w o

l at er al por t io n s the posterior p art is fl at t e n ed an d more
, ,

or l ess qu adr a te in ou t li n e G e n etic ally the hypop hysis h as

.

n o co nn ec t io n w ith the br ai n but arises as an outgrowth from

,

the roo f o f t h e sto modeum .

Th e cer ebr a l lz em isp /z er es are elo ng at ed bod ies lyi ng in

fro n t of t he th al ame n ce ph alo n t hey t aper so mewh at a n t e
r io rly a n d a re sep a r a ted from e ach o t h er by t h e sag it ta l
fiss u r e Th eir c avi t ies th e la ter a l ven tr icles com mu n ic at e
.
, ,

wi t h t h e t hird ve n t ricle by the fo r a m en of [Mo n r o an t eriorly

th e l ateral ve n tricles exte n d i n t o the olfa ctory lobes .

The olf a cto ry lo bes lie j ust in fr o n t o f t h e c erebral h em i

spheres o f whic h they are bu t t h e co n t i n u at io n Th ey are
,
.
XV I TH E NE RVO U S S YS TE M 29
5

sep ar ated fro m th e l atter by a sh allow l ater al sulcus U nlike .

the cerebral hemispheres they are closely fused together in ,

th e middle li n e ; o n th e ve n t r al side however t hey show , ,

a well m arked medi an fissure

-
A n teriorly they give off th e .

olfactory n erves
Th e Cranial Nerves —There are t en p airs of n erves in
.

FIG . 85 i g m f h dis t ibu t i f h fif h s th i h d t t h

. D a ra o t e r on o t e t e ve n ,
n nt ,
an en

c i l s h fi st sp i l i p t f th sym p
,

ran a n er ve d th t t e r na n e rv e , a n e a n er o r ar o e a

h ic A sys t mic ch f ; b pl b chi l p l xus ;

,

t et . o, e d ar o ao rt a r . ra a e or

du d um H h t ; H y hy id th l y i
,

sa l t ;
ao r a au ,
’
o d by
en ea r o w i i l a n t e r o r an
?
,

p st i h s ; L lu g ; N s l b ; O bi t ; P l p ulm a y
,

o er o r o rn , n ,
na a o ne n or u ,
on r

y ; Sp A s p l ch ic
a rt e r . li m s t i c t y ; S t t m ch ;
,
an n o r c oe ac o -
e en e r ar e r ,
s o a

S y sym p h ic ; 1 1 d f p t ic O p h h lmic V m il
,

1 2
, t at et ; V , cu en o o n e rve t a ,
ax

l y d V m dibul b ch f fif h t ig mi us
, ,

3 1
ar ,
an , an ; V1 1 ar r an o t or r e n n erve ,

p l ti d V1 1 hy m dibul b ch f f ci l ; I X g l p h y
,

a a n e , an 2 o an ar ran o a a o sso ar n

g l gus i t h X c di c Xg g s t ic Xl l y g l
, ,

; X
’
ea n e rve va w oa , ar a as, a r ar ar n ea ,

d Xp l p ulm y b ch s ; Sp fi s t s p i l h y p gl ss l ;
, , , ,

an u , o n ar ra n e I r na n e r ve , o r o o a

p 5 p s c d fif h s p i l s (F m P k d P k s
,

2 —s s , e on to t na n er ve . ro ar er a n ar er
'

Z l gy sligh y m difi d f m H s )
oo o ,
t
‘
o e ro ow e .
29 6 TH E BIOLOGY OF TH E FROG CH A P .

the frog which a rise fro m the br ai n an d are k n ow n c o n se

qu e n tly as cr an i al n erves .

The first n erves cou n ti n g fro m be fore b ackw ard are

, ,

t h e ol fa cto ry They a rise from th e olfactory lobes by two

.

roots t h e an t erior o n e emergi ng from th e fro n t en d th e

, ,

pos t erior ru n n i ng alo n g the ve n t r al sid e o f the lobe n e arly

to i ts poste rior e n d The olfactory n erves p ass through
.

sm all for ami na in th e eth moid bo n e an d are distributed to

th e w alls o f th e n as al ch ambers .

Th e op tic or seco n d p ai r of cr an i al n erves ari se fro m th e

, ,

t h al a m e n ceph alo n an d a ft er crossi n g in th e chi a sm a em erge

, , ,

fro m th e skull th ro u gh for a mi n a in th e sides o f t he cho ndro ‘

cr an ium an d are distributed to the eyes B o th the olfactory

, .

a n d th e optic n erves ar e pu rely se n s o ry .

Th e third n erve ocu lo m o to r is sm all an d arises fro m

,
-

t h e ve n tr al surface of th e crur a cerebri I t emerges from the .

skull th rough a sm all for a m e n n e ar th e O pe n i n g from th e

op tic n erve a n d i n n erv ates four of th e muscles o f the eye
,

b all t h e rec t us sup erior r ectus i n ferio r rec tus m edi alis an d
, , , ,

obliquus in ferior A fter givi n g off a br an ch to th e rec t us

.

superior th e thi rd n erve becomes co n n e cte d wi t h t h e cili ary

,

g an glio n ; this gan glio n also receives a bran ch from th e

o p h th a l mic divisio n o f th e fi ft h n erve an d gives off th e s m all ,

cili ary n erves to t h e retr actor bulbi muscle an d th e tu nics

o f t h e eye .

The fourth n erve the t r ocb lea r is is very sm all ; i t arises

, ,

fro m the dors al sid e o f t h e br ai n bet w ee n th e optic lobes ,

a n d th e cerebellum I t le aves th e skull through a sp eci a l

.

for am e n a lit t l e abov e t h e optic n erve an d is distribu ted to

th e superior obliqu e muscle of the eye B oth the third and .

the four t h n erves are exclusi vely mo tor .

The fift h n erve the t r igem in u s or t r ifa cia l i s o n e o f the

, ,

l arges t o f t h e cr an i al n erves I t arises fro m t h e sides of the

.
29 8 TH E BIOLOGY OF TH E FROG CH AP .

to th e n as al ch ambers an d an terio r portio n o f the roof o f

the mo u th The seco n d br an ch t h e lzy o m a n zlibu la r ru n s
.
, ,

outw ard an d the n b ackward arou n d t h e audi t ory c apsule

, ,

p assi n g over the colum el la an d aft er receivi n g a br an ch

, ,

fro m th e glossoph ary n ge al ru n s ou tw ard gives some twigs

, ,

t o the ear an d mu scles of th e lower j aw an d th e n divi des ,

n e ar th e a n gl e of t h e i n to th e m a n clibu la r is in ter n u s
j aw ,

which ru n s forw ard close to t he m an dible an d th e by oizleu s , ,

which i n n erv ates th e su b hyo deus muscle an d the S ki n in t h e

regio n of t h e t hro at Th e s e ve n th n erve like the fifth co n
.
, ,

t ai n s both se n sory an d motor fibers .

Th e eighth or a u dito ry n erve i s di stribu ted e n tirely to the

, ,

i n n er ear .

The n i n t h or glossop /z a ry ngea l n erve arises fro m the sides

, ,

o f th e m edull a fro m a grou p o f roots in co mmo n with the

v agus t he s e roots e m erge fro m th e skull through a for ame n
in th e exoccipit al exter n al to the co n dyle an d e n ter th e l arge
j ug u la r g a nglio n S hor t ly a ft er its e m erge n ce from this
.

g anglio n the tru n k of th e glossoph ary n ge al be ars a sm all

g an glio n ic swelli ng an d the n soo n divide s th e o n e br an ch ,

p assi ng forw ard to j oi n the hyom an dibul ar di visio n o f the

fa ci al the o ther ru nn i ng forw ard in a si n uous course alo n g
,

th e floor o f th e m ou th and i nn erv ati n g th e m ucou s m em

br an e of th e to n gue an d ph ary n x .

The te n t h (vag u s or p n eu mog a s tr ic) n erve emerges from

the j ugul ar g anglio n ge ner ally by two tru nks th e sm all
an terior tru n k the r a m u s a u r icu la r is is d istributed to t h e
, ,

regio n o f the tymp an um the m ai n n er ve p asses b ack w ard ,

an d a ft e r givi n g off so m e sm all br an ches to th e mu scles

,

of t h e S houl der becom es d istributed t o th e l ary n x eso p ha

, ,

gu s stom ach lu n gs a n d h e art B o th the glossop h ary n ge al

, , ,
.

an d v agus co n t a i n se n sory an d mo tor fibers The a ccesso r iu s .

,

which is a sm all br an ch supplyi ng th e cucull ar is muscle ,

xv1 TH E NE RVO U S S YS TE M 299

become s in th e higher v ertebrate s an i n dep e n de n t cr ani al

n erve .

The S ymp at het ic — S y st em

The m ai n tru nks of th e .

symp ath e t ic sys t em co n sist o f a n ervous s t r an d o n either

S ide o f the spi n al colu m n A n teriorly e ac h t ru n k begi n s
.

in th e proo t ic g an glio n fro m which i t exte n ds b ackw ard

,

withi n the cran i al c avi t y le avi ng th e skull by th e j ugul ar

,

fo ra me n in co m mo n w i t h the v agus I t receives a br an ch .

fro m t h e j ugul ar g anglio n wh ich e n te rs t h e firs t ganglio n ic

,

e nl arge me n t o f t he m ai n tru n k E ach tru n k receives a .

bran ch ( r a m u s co m m u n ica ns) fro m e ach of t h e spi nal

n erves a n d where th e t w o j oi n t h ere is
,
a g a n glio n ic em

l argeme n t Th e eighth spi nal n erve however is co n n ected

.
, ,

with t h e symp at h e t ic by two commu n ic ati n g bran ches and ,

the n i n t h n erve b y thre e an d som eti mes four The l ast .

g anglio n o f the symp ath etic ch ai n co n sists o f th e n i n t h an d

te nt h g angli a fused i n to o ne i t receives besides th e br an ches ,

fro m the n i n th spi nal n erve a si n gle sm all br an ch fro m the

,

t en th .

A p art o f th e fibers fro m th e symp ath etic tru n ks e n te r

t h e spi n al n erves by w ay of th e co m mu n ic ati n g r a mi oth er
fi b ers form i n depe n d e n t sy mp at hetic n erves F ro m the a n .

t erio r portio n of t h e tru n ks br an ch es are give n off to th e sub

c lavian an d occipi to ver t ebr al a rteri es an d an terior e n ds of

‘

t h e ovid ucts F ar t her b ack ( fro m t h ird to sixth ga n gli o n)

.

several n erves are give n off which u n ite to form the cceliac
or sola r p lex u s in which sever al g anglio n ic m asses occur
,
.

F rom this plexus n erves ar e d istributed to th e sto m ach ,

i n testi n e liver p an cre as splee n ov aries oviducts an d

, , , , , ,

kid n eys F arther b ack s ever al br an ches anastomose to

.

for m the u r og en ita l plex us which supplies th e ki d n eys , ,

ov aries oviducts and testes

,
The g an glio n cells o f the
, .

symp ath etic system are o ft e n situ ated very far fr o m th ei r

3 0
0 TH E BIOLO G Y OF TH E FRO G CH AP .

poi n t o f origi n Th ey are fou n d embedded alo n g the

.

course of c er t ai n n erves su ch as the v agus an d fa ci al ; in

the w alls o f the i n t esti n e ( A u er ba c/z s a n d M eiss ner s ’ ’

n th e h e art ( Rem a b s B z cl zl er s D og iel s g a nglia

'
’

) i
’ ’

p lex u ses , ,

an d sever al s m alle r groups o f cells ) ; in th e w alls o f the

bl add er ; an d in the ski n .

Reflex A ct ion Th e re flex actio n s o f th e spi n al cord of

.

the frog m ay be ill u str ated by the followi n g experi me n t '

C ut t he spi nal cord o f a frog across j ust behi n d th e m edull a

an d aft er d es t royi n g th e br a i n
,
susp e n d th e an i m al u po n
,

a hook p assed th rough t h e u pp er j aw ,

No w pi n c h o n e o f
.

t h e to es the hi n d foot w ill be dr aw n u p to th e body I f a .

pi ec e o f blo tti n g p aper s a tur ated with dilute ac etic acid be

pl aced o n o n e side o f th e body the hi n d leg o f th at S id e
,

will be brough t forw ard an d the acid wiped away with th e

foo t I f the a cid is pl ace d n e ar the middle of the body
.
,

bo th hi n d feet m ay be e mployed to re move it The vigor .

o f th e respo n se d ep e n ds upo n the stre ngth of th e sti mulus ,

a we ak sti mulus produ ci n g o n ly a slight move me n t while a ,

very stro ng sti mulus w ill throw the an i m al i nto viole n t c o n

t o rt io n s . S imil ar respo n ses m ay be evoked from the fore
li mbs a nd if a ci d be pl aced o n the S i de o f th e body so me
,

dist an ce in fro n t of the hi n d l egs both fore an d hi nd limbs

,

m ay be e mployed to remove th e irrit an t Th es e actio n s are

.

of a very defi nite and m ech an ic al type an d i f the frog is in , ,

go od co n ditio n follow i n e vit ably u po n th e app lic atio n o f th e

,

s t i mulus They are termed refl ex actio n s o n accou nt o f a

.

cert ai n an alogy wi t h th e reflec t io n of light which m ay be

thro w n b ack u po n i ts origi n al source by a mirro r Th e .

spi n al cord n erves co n t ai n t w o ki n ds of fibers the affere n t ,

o r se n sory which co n duc t i mpulses fro m t h e periphery to

,

t h e cord an d effere n t
, ( usu ally mo t or fibers ) by m e ans ,

o f which i m p ulses are c arried ou tw ard to o th e r organ s .

30 2 TH E BIOLO G Y OF TH E FRO G CH A P
.

r egio ns in fro n t o f and be hi n d the po i n t stimul ated Th e .

'
stro nger th e sti mulus th e gre ate r is the p art o f the body
,

i nvolved in th e respo n se A we ak stimul us applied to the

.

foot produces move m e n t o nly in the me mber stimul ated ,

while a much stro nger sti mulus m ay c aus e a move m e n t o f the

opposite leg as well as other p arts o f th e body I f acid b e‘

pl aced o n o n e side of the body an d th e leg of t h at sid e h eld ,

the leg o n the opposite sid e i s so meti m es brought a rou n d

to wi pe th e acid away Thi s app are n t ly i n t ellige n t act is
.

obviously depe n d e n t upo n t he p ass age o f impulses from o n e

sid e of th e cord to th e o ther A nd i t requires a stro nger
.

stimulu s to bri ng i t about th an is n ecess ary to produce

a si mple u n il ater al refl ex .

Th ese reflex actio n s of th e spi nal cord are of a purposive

n ature ; they bri n g th e li mbs aw ay fro m i nj urious stimul i

an d remove i r r it ati ng subst a n ces fro m th e body Th ey are

.

depe n de n t u po n th e org an iz atio n o f th e an im al the n euro ,

muscul ar m ech anism bei ng such t h at a s ti mulus to any p art

o f the body bri ngs about th e appropri a te actio n s for rem o v
ing or esc api ng from th e source of i nj u r y .

Th e reflex ac t io n s o f a frog m ay be ch ecked or modified

by impulses fro m t h e brai n A frog with i ts br ai n in c o n
.

n ec t io n w ith the cord will n o t respo n d to sti mul atio n in t h e

s ame regul ar an d u n varied m an n er as a br ai nless i n dividu al .

I t s actio n s are much less m ech an ic al an d more spo n t an eous

an d u n cert ai n of predictio n The i n flue n ce o f th e br ai n i s
.

re n d ered possible by m e an s o f n erve tru n ks which p ass fro m

t h e br ai n dow n the spi nal cord an d form co nn ectio n s wi th

t h e n euro n es of the spi n al n erves I f th e an terior e n d

.

o f th e spi n al cord is s t ro ngly sti mul ate d at th e s am e time a

sti mulus is applied to the foot th e withdr aw al of th e l atter
,

m ay be e n tirely preve n ted I n this w ay the ordi n ary refl ex

.

a c t io n s of t h e cord are co n ti n u ally checke d an d modified by

i mpulses fro m the higher n e rvou s ce n ters .

XV I TH E NE RVO U S S YS TE M 30 3

Th eCroak ing Reflex the S id e of a fr og be st r oked

. If
w it t h e fi nger the an i m al o ft e n respo n ds by c r o aki ng Th e
, .

s am e re actio n freque n tly occurs whe n th e frog is picked up

in th e h an d . I t t akes pl ace m ore ofte n in the m ale frog
th an in th e fem al e e speci ally duri ng th e breedi n g pe r iod
, ,

whe n the c r o aki n g m ech an ism is very re adily put i n to

activity . A frog th at is seized m ay co n ti nu e to cro ak vigor
o u sly for some time an d whe n it h as ce ased to cro ak i t m ay
,

be i ndu ced to co n ti n ue by rubbi ng its side with the fi nge r .

The cro aki n g re actio n however i s very vari abl e an d in

, , ,

c er tai n i nd ividu als i t m ay n o t be perform ed at all A nd the .

s ame i ndivid ual re ac ts qui te differe ntly at diffe r e n t ti mes “

acco r di n g to wh at seem s i ts o w n c a price .

I t is quite otherwise with frogs fro m which th e ce r eb ral

h emisph eres h ave bee n removed I t w as fou n d by G oltz .

th at an im als thus oper ated o n cro ak with mech an ic al r egu

lar it y wh e n ever o n e strokes their side o r b ack They n ever .

cro ak whe n t h e hi n d legs or ve ntr al sid e o f th e body is

stroked an d th ey cro ak o n ly o nce aft er e ach st r oke o n the
,

b ack or side Th e experi me n t succeeds well with both

.

sexes bu t the cro ak of the m al e is n atur ally much loude r

,

th an th at of the fem al e “ I k n ow of sc arcely any physio

.

logic al experi me n t s ays G oltz which succ eeds in so ce r

,
”
,

t ai n an d regul ar a m an n er a s thi s cro aki n g experi me n t .

AS I w as de liveri n g a discourse u po n this subj ect be fo r e th e

m eeti n g o f th e n aturalists at H ann over in th e ye ar 1 8 6 5 I , ,

pl aced upo n th e t able for purposes of d emo n str atio n sev

, ,

eral frogs oper at ed up o n sever al mo n ths previously whic h ,

I h ad brought wi t h m e from K o n igsbe r g I n th e course .

o f t h e sessio n I asked H err V o n Wi ttic h who w as p r ese n t ,

in t h e audi e n ce how oft e n e ach of the frogs which we r e

, ,

resti n g sile n tly an d still S hould c r o ak Th e an swe r five

,
.

times w as give n an d e ach of th e frogs u po n bei ng give n five

, ,
3 4
0 TH E BIOLOGY OF TH E FROG CH A P .

strokes c r o aked ex actly five times to th e evid e n t gr atific atio n

, ,

o f the audi t o r s wher eu po n they all l apse d i n to S ile n ce

, .
”

The cro aki n g o f a br ai nless frog is a r efl ex se t in ope ra

tio n by a p ar ticul ar loc alized exter nal sti mulus I n a n orm al .

sp eci me n t his r e flex is subj ec t to co n t r ol fro m th e high e r

n erve c e n ters I t m ay be e ntirely ch eck ed by i m pulses
.

dow n th e spi nal cord from t he brain The n orm al frog .

shows a spo n t an ei ty an d freed o m of actio n whi ch is l argely

d estroyed in i ndividu als which h ave rece n tly lost th ei r
c e r eb ral h em ispheres The l a tte r oft e n cro aks wi t h out any
.

app are n t exciti ng c au se while a br ai n less frog c r o aks o n ly

,

upo n the applic atio n of an exte r nal sti mulus to cert ai n p ar t s

o f th e body .

The u tility of th e c r o aki ng reflex is n ot app ar e n t I t h as .

be e n suggested by B aglio n i th at si n c e in cro aki n g air i s

,

push ed b ack an d forth betwee n the m outh an d th e lu n gs ,

the swelli ng thus produced e nables th e an im al to push

ag ai n st an y t hi n g th at sei z es i t and m ake i ts esc ap e th e ,

sou n d bei ng m erely an i n ci de n t al accom p an i me n t o f th e

p r ocess Any o n e w h o picks u p a frog in the h an ds m ay
.

r e adily co nvi n c e himself th at th e cro aki ng move m e nts are a

m e an s o f e nabli n g th e cre atu re to S lip fro m h is gr asp In .

'

fac t t h ese moveme n ts o ft e n occur in t he fe m ale withou t

,

p r oduci ng an y sou n d wh atever I t is a S ig nific an t fac t th a t

.

th e refl ex is evoked by sti muli u po n those p arts which as ,

th e body swells press ag ai n st wh at ever seizes the an i m al

,
.

I t is n o t b r ough t about by seizure of the h e ad or hi n d legs ;

th e swelli n g in such c ases would be of n o av ail Th e fac t .

th at th e cro aki ng r eflex is brought abou t by ge n tly stroki ng

the sid e or b ack m ay i n dic ate n o t hi n g bu t t h e ext r em e readi
n ess with which the swelli ng reflex is i n i t i at ed Whe n the.

an i m al is sei zed an d th ere is a co n st an t pressure sti mulus ,

the r e is a r ep e ated swelli ng o f th e body and accomp an yi n g

3 0 6 TH E BIOLOGY OF TH E FRO G CH AP .

a n u mb er o f frogs of both sexes are pl aced togeth er duri ng

th e breedi n g s e aso n the m ales wil l be fou n d to be cl aspi ng
,

fe m ales i n ste ad o f o t her m ales . Th e questio n n atur ally

arises : H o w doe s t h e m ale frog dis t i nguish th e fem ale
fro m o n e o f his o w n sex ? Th at it is n o t as in m an y ,

an im als,
through the se n se o f smell w as show n by G oltz
,

by cu tti n g throu gh the olfactory lobes of sever al m ale

speci me n s ; afte r this op er atio n the mu til ated m ales were
pl aced amo ng several fem ales an d in a shor t time they were
,

all in copul atio n with m emb ers of t h e o ther sex The n .

several m ales which were bli n ded were pl a ced amo ng the
fem ales wi th th e s ame resul t as before Th at i t is n o t the
,
.

sou n d produced by th e fe m ale th at reve als h er sex w as ,

S how n by G oltz by pl aci ng sever al fe m ales whi ch h ad bee n ,

re n dered i n c ap able of u si n g their voice amo n g t h e m ales

, .

These were as re adily seized as n orm al fe m ale s Aft er t h e .

destructio n o f both sm ell and sigh t m ales were fou n d to

,

be still able to disti nguish th e fem ales although t h ere w as

,

a pp are n tly a ce r t ai n d imi n utio n of th eir a rdor Th e se n ses .

o f sight s mell an d he ari n g are n o t th erefore the exclusive

, , , ,

or i n dispe n s able m e an s o f sex recog ni t io n wh atever be th e

,

p art they pl ay u n der n orm al co n di t io n s Nei t h er are the

.

highe r n erve ce n ters n ecess ary G oltz cut th rough th e S kull

.

of a m ale so as to cut off th e cerebr al h emisph eres an d

eyes th e speci me n al most i mm edi a tely cl asped a fem ale th at
w as prese n ted to i t while a m al e th at w as offere d w as rej ected
,
.

The form of th e body differs in t h e t w o sexes especi ally ,

whe n the fem ale c arri es a l arge m ass o f eggs but this alo n e ,

does n o t e n able th e m al e to disti n guish t h e fem ale G ol t z .

fou n d th at if the bodies of m al es were filled wi th flesh and

sewed up so as to resemble t he form of the fe m ales they ,

would be cl asped for a shor t ti m e an d the n rej ected The .

bodi es of freshly killed fe m ales o n th e o t her h and were

, ,
xvr TH E NE RVO U S SYSTE M 30 7
0

h eld fo r a lo ng tim e th e te n sio n or cram p o f th e mu scles

,

i n cre asi n g the lo nger th e bod ies were h eld G oltz has sug
, .

gested th at th ere is som ethi ng th at e m anat es fro m the b o dy

of the fem ale th at acts as a speci al excit an t to th e m ale but ,

there is n o t su ffici e n t evide n ce for adopti n g thi s V i e w an d ,

th e o n ly re aso n for accepti n g it is th at othe r expl anatio n s of

s ex recog ni tio n d o n o t suffice .

The r e is so much v ari atio n in th e beh avio r o f diffe r e n t

i n dividu als an d so m any factors th at m ay de termi n e wheth e r
,

th e cl aspi n g re flexm ay or m ay no t persist in p articul ar c ases ,

th at it is h az ardous to dr aw co n clusio n s from si ngle exp eri

me n ts D iffere n ces in t actile s t i m ul at io n m ay pl ay a rOle in
.

sex recog nitio n an d differe n ces in t h e move me n ts of th e t w o

,

sexes m ay be an oth er factor The experi me n ts of G oltz

.
,

although very i n structive an d suggestive h ave n o t succeede d

,

in S howi n g sex recog ni tio n to be th e result o f an y o n e c ause ,

an d it is n o t improb abl e th a t i t m a d epe n d upo n sever al

y
factors e ach of whic h is n o t e n tirely i n dispe n s able
,
.

S ex discri mi nat io n in frogs is n o t very precise M ales .

are some times h eld by other m ales for a lo n g time an d ,

copul a tio n is n ot i n freque n tly k n ow n to occur betwe e n the

m ales of di ffere n t spe cies S p all an z an i records a m ale to ad
.

th at w as c arryi n g arou n d an i n dividu al o f his ow n sex th at

h ad d ied som e d ays previously an d w as in an adv an ced st at e
of dec ay Wh eth er or n o t a m ale will cl asp an obj ect oth er
.

th an th e fem ale d epe n ds upo n th e le n gth of time h e has

bee n sep ar ated fro m th e sexu al e mbrace I f a m al e is
.

r ece ntly torn fro m a fem ale he is very sluggish an d will

, ,

cl asp the fi n gers or almost any obj ect th a t is prese n ted to

him I f left for a t ime he becom es mo r e active an d sho w s
.
,

lit t le te n de n cy to cl asp all sorts of obj ects M al e frogs

.

which at first would be seized are n o w usu ally rej ected o r ,

held bu t for a short t im e but if a fem ale is prese n t ed sh e ,

is s ei z ed with eage r n ess .

3 0 8 TH E BI O L O GY O F TH E F RO G CH A P .

Compensat ory M ot ions frog i s rot ated o n a c ircu

. If a

lar disk with i ts h e ad poi n ti ng aw ay from th e ce n t er t h e he ad ,

of the an im al will t ur n opposite th e directio n o f rot atio n .

Th e tu r n i n g o f th e h e ad i s oft e n followed by loco motio n in

the s am e di r ectio n s o th at th e frog keeps circli ng arou n d
,

in o n e direc t i o n while th e d isk is ro t ati n g in an othe r If .

t h e frog is pl ace d o n a bo ard an d tilted eith er u p o r dow n

, ,

t h e he ad i s tur n ed opposite t he directio n of motio n C om .

b in ed m ove m e n ts of rot atio n an d tilti n g are followed by

correspo n di ng co mbi n ed m ove m e n ts of the he ad th e resul t ,

bei n g in all c ases to keep t h e he ad as n e arly as possible in

i ts origi nal positio n S uc h m ove m e n ts are therefo r e c alled
.

co mpe n s atory motio n s They t ake pl ace in respo nse to very

.

S ligh t ch an ges in the positio n of th e body an d they occu r ,

wi th rem ark able r egul arity app are n tly i n depe n de n tly o f the
,

an i m al s voli tio n They are performed aft er the d estructio n

’
.

of all p arts of th e br ai n in fro n t o f th e medull a or eve n aft er ,

t h e remov al o f th e an terior p art of th e l atter org an provided ,

the i nj ury does n o t exte n d so far b ack as th e trigemi n u s

group of n erves I f the se micircul ar c an als are d estroye d
.

or the auditory n erves cut compe n s a tory m otio n s acco r di n g

, ,

to S chr ader are n o lo nge r performed bu t S t ei n er fi nds th at

, ,

compe n s at ory motio n s o f the he ad at le ast still t ake pl ace .

The F u n ct ion s of t he B rain —Th e br ai n i s t h e g r e at

.

c e n te r o f com mu n ic a tio n betwee n the pri n ci p al org an s o f

se ns e an d th e rest of t h e body ; th r o ugh i t a re effec t ed the
n u merous c o Ordin a t io n s betw ee n a gre a t v ariety o f stimuli ,

sights sou nds odors etc an d th e appropri at e muscul ar

, , ,
.
,

a ctio n s which e n able the an im al to adj us t itself t o the e n

viro n m e n t . The n u mb er an d n ature o f th e co n n ec tio n s

es t ablished in the c e n tr al n ervous syste m determi n e to a l arg e
degree t h e ch ar ac t er of t h e an i m al s i nsti ncts an d th e possi
’
,

b ilities a n d the li mits of th e developm e nt of it s i n t ellige n ce .

310 TH E BIOLOG Y OF TH E F RO G CH AP .

sph eres we r e c are fully removed wi thout i nj uri ng th e th al ami ,

an d the an i m al kept some mo n t hs after th e oper atio n spo n ,

t an eit y o f actio n is by n o me an s lost ; t h e an im als move

abou t fro m l an d to w ater bury the mselves in th e mu d o n th e

,

a ppro ach of wi n ter an d s n ap a t flies which m ove wi thi n their

,

r e ach Their co ndu ct in fact rese mbl es very closely th at

.
, ,

of n orm al frogs and in j us t wh at resp ect i t differs S chrader

does n o t m ake cle ar A fter loss o f t h e cerebral hemi spheres
.

all fe ar a ccordi n g
,
to K a to dis appe ars an d the frog n o
, ,

lo n ger cro aks spo n tan eously but ho w lo ng aft er the oper a
,

ti o n these results were observed is n o t s t a ted S i n ce so .

m any of th e fu n ctio n s su pposed to h ave their s e at in th e

cerebral he mispheres are show n to be pe r form ed by other
p arts o f the br ai n it is n o t d efi n i t ely es t ablish ed j ust w h a t
,

r Oles th ese org an s re ally pl ay I n high er an im als they are

.

t he se at o f i n tellige n ce and volu n t ary co n trol ; an d it i s n o t

i mprob able th at wh atever i n tellige n ce the frog m ay possess
m ay h ave a si mil ar loc al h abi t atio n B ut this co n clusio n .
,

prob able as it m ay be c ann ot be r eg arded as est abli shed by

,

exp eri me n t .

Remov al of o ne ce r ebr al he misp here i s n o t followed by

so severe S hock effects as th e rem ov al of th e e ntire cere
br u m Th e frog thu s op er ated u po n is accordi n g to K ato
.
, ,

m ore active it s p ri n gs more re adily an d m an i fests fe ar upo n

, ,

s e ei ng t h e appro ach of l arge obj ects I t te n ds at first t o .

avoid obst a cles pl aced in its p a th by tur n i n g aw ay fro m

t he oper ated sid e b u t in four d ays a fter th e oper atio n it s
,

beh avior accordi ng to L o eser becom es e n t i rely n orm al

, ,
.

I f t h e p arie t al regio n of th e cerebr al he mispheres be s t im

u lat e d by a very we a k curre n t of elec t ricity move me n ts o f ,

t h e limbs are brough t abou t o n the opposi t e side of th e

body an d usu ally al so although t o a less ex t e n t o n th e s a m e

, , ,

sid e wh ere as stimul atio n of o t h er p arts o f t h e cerebr al

,
xvr TH E NE RVO U S S YS TE M 3 1 1

hemispheres produces n o such e ffect ( Kraw z o ff K ato ) No , .

defi n i t e loc aliz a tio n o f motor fu n ctio n s such as h as bee n

est ablish ed in the br ai n s of the higher m am m als h as ye t
bee n worked out
Tb a la men cep b a lo n — Re mov al o f th e th al a me n ceph alo n
.

alo n g wi t h th e cerebr al he mispheres results as before t e ,

m arked in an al most complete loss of spo n t an eous move

,

m e nt s Th e an im al r ests quietly an d seldo m m akes a move

.
,

u nless in r espo n se to some ex t er nal sti mulus an d shows a g en

-
,

er al i n se nsibili t y to touch ( L oeser ) S i n ce the optic n erves .

e n ter t h e t h al a m e n cep h alo n re mov al o f this p art c aus es

,

co mplete bli nd n ess ; the frog n o lo n ger avoids obst acles

pl aced in its p ath or shows any re ac t io n i nvolvi n g the powe r
,

of visio n Wh e n pl aced o n a pi ece of wood th at is slowly

.

tilted upward i t will n o t climb to th e upper S id e an d b al an ce

t here like a frog th at h as lost i ts cerebr al h emisph eres but
, ,

i t respo n ds o n ly by move me n ts o f the he ad loweri n g th at ,

p art whe n the an terior p art of t h e body is r ais e d an d r ais

ing it wh e n the revers e movem e n t is perform ed ( S tei n er ,

S chr ader ) I f the an i m al i s ro t a ted o n a horizo n t al disk

.

it respo n ds by tur n i n g soo n aft e r th e oper atio n it shows n o

co mpe n s atory mo tio n s ; l ater i t respo n ds by tur n i n g th e
,

he ad o pposite to th e directio n of rot atio n an d after co m , ,

p et e recovery p erforms circus m ove me n ts o f th e body in

l ,

the s ame directio n as those of the he ad The s a me move .

m e n ts are also performed after loss o f th e optic lobes Th e .

loss o f t h e power to b al an ce th e body o n the edge of a tilted

bo ard is app are n tly perm an e n t .

T h e Op tic L obes Accordi n g to S tei n er th e ce n ter for

.

c o Ordin at e d motio n is in th e op t i c lobes F rogs from which .

t his p art of th e br ai n w as removed were fou n d by S t ei n er to

swim wi t h alter n ate m ovem e n ts o f th e legs i n s t e ad of co m e

b ine d strokes S chr ade r sho w ed however th at this w as th e

.
, ,
31 2 TH E BIOLO G Y OF TH E F RO G CH AP .

result o f shock e ffects and th at frogs c ar e fully ope rated

,

u po n an d give n ti m e to r ecove r perfo r med c o Ordinat ed ,

j umpi ng an d s w i m mi ng m ove me n ts re m ark ably well Th e .

c e n ter for loco motio n c an n o t therefore be loc a ted in th e

, ,

optic lobes Re moval o f t h e optic lobes w as fou n d by L o eser

.

to produce forc ed m ove m e n t s an ab n o r m al r ete n tio n of

,

uri n e an d a slight loss o f sight but l ater th es e symp to m s

, ,

gr adu ally dis appe ared F rogs with the br ai n re moved as far
.

b ack as th e m edull a loco mo t e n orm ally perform the cro ak ,

ing re flex bre athe regul arly and S ho w the u su al co mp e n s atory

, ,

motio ns .

The optic lobes exercise an i nhibiti ng i n flu e nce o n th e

re flex ac tivi ty o f the spi n al cord Re m oval o f th e optic .

lobes resul t s in an i n cre a sed irrit ability of the cord while ,

sti mul ati o n o f th e optic lobes m ay gre atly ch eck spi n a l

reflexes I f o n e op tic lobe is sti mul ated the spi nal re flexes
.
,

o n the opposite side of t h e body a re i n hibited while those ,

o n th e s a me sid es are n o t n otice ably affected ( L an gen do rff

) .

T/z e Cerebellu m The cerebellu m o f th e frog is so sm all

.
-

co mp ared with th at of most oth er ver t ebrat es th at it would ,

n o t be expected to h ave an import an t fu n ctio n S tei n er .

fou nd th at frogs fro m which the cerebellu m had bee n

rem oved were app are ntly u naffected by th e oper atio n an d ,

o t her i n ve stig ators h ave co nfirm ed this co nclusio n G oltz .

,

ho w ever fou n d th at co Ordinat ed loco motio n w as disturbe d

,

after extirp atio n o f the cerebellum bu t as w as admitted l ater , , ,

som ewh at more th an the cerebellu m alo n e w as re moved .

Rece n tly however L o eser h as a rrived at resul t s simil ar to

, ,

t hose o f G ol tz Afte r re mov al o f the cerebellu m accordi n g

.
,

to L o eser “ th e j u mps bec a me very u n ste a dy an d were

,

we ake n e d in force bu t n o t much red u ced in r ate M ost o f

,
.

th e ti me th e frog lay s t re t ch ed in co n t ac t with the floo r

of th e vessel an d covered with moss Th e u nste ady an d .
3 14 TH E BIOLOGY OF TH E FROG CH A P .

m ai nly by creepi n g bu t th e frog is n everth eless c ap able

,

o f spri n gi n g in the ordi n ary m an ner I n th e w ate r i t swi ms

.

by alter n ate move me n ts o f th e li mbs The resti n g po siti o n

.

assu m ed i s n o t quite n orm al th e body i s m ore fl atte n ed

,

th an u su al an d th e h i n d limbs are n o t dr aw n u p to the S ide s

, .

Th e bre athi ng an d sw allowi n g refl exes are still n orm al b u t ,

th e cro aki n g refl ex is n o lo n ger p erformed The reflex o f .

S n appi n g at food is n o t d estroyed al though in the abse n ce

,

o f co n n ec tio n with the eyes it i s n o t p erformed in respo n s e

to visu al stimuli I f a pi ece o f m e at i s rubbed ag ai n st th e
.

frog s n ose th e an im al s n aps at i t an d uses the fore legs t o

’
,

stuff i t i n to the m ou th Th e s am e re actio n m ay be brought

.

about by u si n g th e fi n ger but a ft e r th e fi n ger is sei zed an d

,

i t is fou n d th at th e obj ect is too l a rge to be stuffed i n to th e

mo u th t he frog begi n s to rej ect th e morsel an d u ses th e fore
, ,

legs to push i t aw ay Truly a re m ark able combi natio n o f

.

reflexes !
Th e s nappi n g reflex is also bro u ght abou t by co n t act
with th e fore l egs o r eve n other p ar t s of th e body I f s ev
,
.

er al frogs whose br ai n s are cut through j ust behi nd th e

,

cerebellum b e pl aced together in a box they frequ e nt ly

, ,

s nap at e ach oth er whe n t h ey accid e n t ally co me i n to c o n

t ac t as if they were defe ndi ng themselves ag ai nst att ack
,
.

F lies cre epi ng over th e n ose are u su ally n o t s n apped at bu t ,

th e stimulus produc es an othe r reflex the fo re leg bei n g ,

brough t forw ard t o bru sh t h e i n sec t aw ay .

The s n appi n g reflex is preve n ted if t he faci al n erve is cut ,

or i f th e regio n o f th e m edull a fro m which t h e fi fth an d

seve n th n erves origi nat e is d estroyed These n erves form .

t he affere nt an d effere n t ch ann els respectively of the n ervous

i m pulses i n vol ve d in t h e s n appi n g refl ex Th e extrusio n of .

t h e to n gue w hich goes alo n g w i t h t his proc ess is depe n de n t

, ,

upo n the hypoglossus n e r ve which has its origi n fu r the r

,
XVI TH E NE RVO U S S YS TE M 315

b ack in th e m edull a D estru ct io n of the roo t o f t his n erve

.

preve n ts th e extru sio n of the t o n gue b u t th e s nappi n g o f t h e

,

j aws m ay still occur The two p arts of th e reflex of seizi n g

.

food m ay therefore t ake pl ace i n dep e n de n tly although they ,

are n orm ally almost alw ays associ at ed .

The sw allowi n g refl exes p ersist after d est r uctio n of the

ce n te r for s nappi n g They are depe n de n t u po n th e v agu s
.

group of n erves an d dis appe ar o n ly w h e n th e regio n of t h e

,

medull a is i nj ured fro m w hich th e roots o f th e v agus group

a rise. Norm al bre athi n g m oveme n ts still occur whe n the
medull a is cut ac r oss just behi n d the cerebellum an d th e p ar ts
in fro n t of the cut e n tirely removed They are n o t d estroyed
.

if the m edull a is also cu t across at the tip of the c al amu s

scriptorius but if the regio n betwee n these t w o cu ts is re
,

moved r espir atory moveme n ts e n tirely ce ase This regio n

,
.

i n cludes therefo r e the ce n ters for the move me n ts of

, ,

r espir atio n .

Th e posterio r regio n of the medull a co n t ai n s those p arts

of the b rai n m ost esse nti al for th e m ai n te nan ce o f life S o .

lo n g as this regio n rem ai n s th e an im al m ay live for a lo n g

,

ti me Re mov al of th e medull a far t her b ack th an the p ars

.

c o m m isSu ralis results in m aki n g l ocomotio n more di fficult ,

although it is still possible wh e n the br ai n is re m oved as far

b ack as the tip of th e c al amus scriptorius The restless

”

n ess of the an im al dis appe ars an d the body te n ds to ass u me

an ab n orm al attitude .

There is n o ce n ter for coordi nated locomotio n in the

medull a D isturb an c es of locomotio n begi n wi th th e fore
.

limbs I f the m edull a i s cu t across at the tip of th e c al amus

.

scriptorius the ani m al si nks o n its bre ast an d th e fore limbs

, ,

are for a co n sider able t ime h elpless a lthough the hi n d limbs

,

are c ap able o f performi n g vigo r ous c o Ordin a t e d move me n ts .

Th e r e aso n for this is th at th e i nj ury lies so n e ar the

316 TH E B IOLO G Y OF TH E F RO G CH AP .

regio n fro m which th e n e r ves o f the fore li mbs ar is e th at

the move m e n ts o f t hes e m e mbers are very n atur ally
affect ed .

Th e S egment al Char act er of t h e F u n ct ion s of t he

Nervous S yst em — Th e fun c t io n s of the c e n t r al n ervous
.

system o f th e frog be ar out th e segme n t al theory of th e

a ctio n o f th e h igher n erve ce n ters whi ch w as firs t suggested

by S chr ader an d mo r e fully el abora ted by L oeb The .

n ervous syste m o f such forms as worms crust ace an s an d

, ,

i ns ects co n sists of a series o f p airs o f g an gli a co nn ected by

a double n erve cord there bei n g typic ally o n e p air to e ach
,

s egm e n t o f th e body ; and as a rule e ach p air of g angli a

forms the ce n te r of th e moveme n ts o f th e p ar ts of th e
s egm e n t in which i t lies S chr ader at the close o f hi s
.

p aper o n th e fu n ctio n s of th e b rai n o f th e frog gives expres

“
sio n to th e segme n t al th eory as follows : Th e seri es of
experime n ts we h ave give n te aches u s th at the ce n tr al
n ervous system of th e frog c an be divided i n to a series o f

sectio n s e ach of which is c ap abl e o f per fo r mi ng an in dep end

,

e nt fu n ctio n . I t bri n gs the ce ntr al n ervous syste m o f th e

frog i nto closer rel atio n wi t h th e ce n tr al n ervou s syste m of
t h e lower forms whic h co n sists of a series o f disti nct g an gli a
,

th a t are co n n ected by com missu res I t sp e aks ag ai n st th e

.

absolu te mo n archy of a si n gle c e n tr al a pp ar a tus an d ag ai n st

th e exis te n ce of differe n t ki nds of ce n ters an d i nvites us to ,

seek for th e c e n traliz atio n in the m any side d coupli ng o f

-

rel atively i n depe n de n t st atio n s .

“
L oeb h as a ttempted to S how th at th e more complic ated
i nsti n cts are for the most p art n othi n g but a series of seg
m e n t al reflexes . I am i n cli n ed “
he s ays to reco m me nd
, ,

usi n g th e word ch ai n re flexes whereby the p erform ance of

-

o n e reflex a cts at th e s am e tim e as the sti mulus for

se tti ng fre e a seco n d reflex The t aki ng of food m ay se r ve
.
318 TH E BIOLOGY OF TH E FROG c n xp .

m achi n ery of th e lower n erve ce n ters of th e frog is guided

by those p ar t s whose organ s of se n se put the m i n to mo r e
i n tim at e co n n ectio n wi th t he ou ter world .

RE F E RE NC E S

Alb ert on i, P . E p er x i c s l c t s en e su r es e n re n e rve ux du c ra

pa ud .

A ch r . I t al de B . i ol .
, T 9 888 .
, 1 .

B e ch t erew , W . U b di F u c t i d V i
e er e n on er e rh u ge l . Ar ch .
g es .

P hys B d 3 3,.
, . 1 8 84 .

B ic el, A k . U b e er de n E in flu ss de r se n sib e len Ne rv e n u nd de n

La by i r nt h auf die B ew e gu g n der Th i e re . Ar ch .
g es . P hys .
, B d 6 7, .

1 897 . B e t ra i g e z ur Riick e n m a rk sp h ysio lo gie de r A m p h ibi en und

Rep t ili en, lc . .

,
B d 71 , . 1 8 98 .

B i ge , E A r . . No t e o n t h e F u n c t i o n s o f t h e Sp i n al Co r d of the F g ro .

A m M o nt h ic J u V l
.M Di Z h l d N
. fs d r
. o r .
,
o . 2 . e a er erve n a e r n un

d m t isch G gl i z ll im Rii k m k d F sch s A ch

er o or e an en e en c en ar es ro e . r .

A t u P hys
na p hys .A b t h 8 8. .
, . .
, 1 2 .

Dal H H O bs ti s
e, P ssib l E ff
. t F ib s i th
. erva on on o e ere n er n e

D s lN
or a R ts f th T d d F g
e rv e J u P hys V l 7 oo o e oa an ro . o r . .
, o . 2 .

D n il ky B a U b di H mmu g d R fl x d W illk
ew s , . e er e e n er e e un ur

b gu g
ew e A ch g P hys B d 4 8 8
n en . r . es . .
, . 2 , 1 1 .

Don l d on H H O bs a ti s t h W igh t d L g t h f t h
s , . . e r va on on e e an en o e

C t l N
e n ra us S ys t m d t h L gs i B ullf gs f D iff t S iz s
e rvo e an e e n ro o er e n e .

j C
o ur m p N u. V l 8 8 9 8
o ; c t i u t i by D lds
. d e r .
, o .
, 1 on n a on ona on an

S c h o em ach e r, l . c. , V ol . 1 0, 1 9 00 .

G as k ell, hy t h m f t h H t f th F g d t h
W . H . O n the R o e e ar o e ro an e

N tu a f th A c t ire o f t h V gus P h il T s V l 73 p t 4
e on o e a . . ra n .
,
o . 1 , ar .

G ol t F E i ig V such tib z ,
d N m h
. im lch n e er e er en erve n ee an s u s, w e er.

w ah d d B g t t u g d F O h th at ig i t C t m d W iss B d 3
ren er e a n er r sc e, s . en . e . .
, .
,

1 865 W it s ub d N
. m h e im t
ere l Bd 4 8 66 er en er ve n ee an s u s, e c .
, . c .
, .
,
1 .

B it g
e L hra d Fu k ti
e z ur d N t d F sch s
e r e vo n en n o nen er e rv e n c e n re n es ro e ,

l. Bd 4
c.
,
8 66 B i t ag . L h ,
d N
1 t d . e r e z ur e re vo n en e rve n c e n ren es

F sch s B l i 8 6 8
ro e . er n, 1 .

H rd t y I a T h Numb d A
es ng m t f t h F ib s f m
, . e er a n r ra e en o e er or

i g th
n S pi l N s f th F g ( R
e i na ) J u C m p e r ve o e ro ana v r escen s . o r. o

8 9 9 —9
.

N u V l 9 Fu t h
e r .
, bs t i s l V l
o . . r er o e rva on ,
. o .
, o . 1 0, 1 1 00 .

H rl E a U b di F u k t i ess , schi d P t hi e d
. e er e n o n en ver e e n er ar n es

R k m k d A m p hibi
uc en A ch A ar t u P hys s 8 46 er en . r . na . . .
, 1 .
xvr TH E NE RVO U S S YS TE M 319

H eub el, E . Das K ram p fcen t ru m deS F sch s u nd s i n e V e h l t en

ro e e r a

gg e en g iss
ew e Ar z is t ff
ne o e . Ar ch .
g es .P hys B d 9 1 8 74 U b .
, .
, . e er

die A b h an g igk e it des vo n w a ch en Ge h i zus t d s

rn an e au ssere n E rr e g u n

g e n . I bi d .
, Bd . 1 4, 1 88 7 .

K at o , T . V e such r e am G ro ssh i rn des F sch esro . I n aug . D iss .

B erl i n, 1 886 .

L an gen dorff, 0 . Die B e z i hu g d

e n es S eh o rg an es zu de n r e flexh em

m en d en Me ch ism an en des F r o sch g eh irn s . Ar ch . A n at . u P hys

. .
, 1 88 7 .

Ub e er Re flexh em m u n g . I bid p 9 6 .
, . .

J
L oeb , . Co mp arat v e i P hysi l gy o o of t h e B ra i n an d Co mp arat ve i
P sych l gy o o . Ne w Y o rk ,
1 9 00 .

L oese udy f t h F u c t i s f D i ff t P t s f t h
r , Wm . A St o e n on o eren ar o e

F gsB i
ro J u C mp N u
’
ra n d P sych V l.
5 9 5 o r . o . e r . an .
, o . 1 , 1 0 .

M n d l oh n M e U t such u g tib R fl
e s S it b d k , . n er n en er e e xe . z er . . .

p uss
re A k iss B li
. 8 8 d .883 w . er n, 1 2 un 1 .

M r b ch r L U b di B zi h u g d S i s g
e z a e d ,
. e er e e e n en er nne or an e z u en

R fl b
e ex
g g d F sch s A ch g P hys B d 8
ew e un 9 en es ro e . r . es . .
, . 1 , 1 00 .

U t suchu g tib di R gul t i d B gu g d W i b lt h i l

n er n en er e e a on er ew e n er r e er e , ,

B b ch t u g
eo F osch
a I bid B d 8 8
n 9 e n an r en . .
, .
,
1 02 .

Not h n g l H B g g h mm
a d M ch e ism im Ruc k
, . ew e un s e en e e an en en

m k d F sch s C t m d Wiss B d 7 8 69
ar e es ro e . en . e . .
, .
, 1 .

S ch lo er W U t suc hu g tib di H mmu g

ss , R fl x . n er n en er e e n vo n e e en

A ch A t u P hys p hys A b th 8 8
r . na . . .
, . .
, 1 0 .

S ch r ad r M Z P hysi l gi d F e h g hi
, A ch g P hys . ur o o e es ro sc e rn s . r . es . .
,

B d 41 ,. 1 88 7 .

S edgw ick , W . T . O n th e V ar i at i o n s of Re fle x E xci t bili t y i th a n e

F g i n duc d by Ch g s
ro e an e Of T mp e erat u re . S t udi s B i l L b J h s e o . a . o n

H op k i ns U i n v .
, V ol . 2, 1 88 2 .

S irot in in . Die p u nk t fOr m ig e b g zt ’

e ren e Re izu ng des F ro sch ru ck en

m ksar ch . Ar . A n at . u P hys p hys A b t h
. .
, . .
, 1 88 7 .

S t in r J e e , . Die F u n ct io n e n de r Ce n t raln e rve n syst e m und ih re

P hyl g s o en e e . A b th U t suchu g
. I ,
n er n en fi b er die P hysi o lo gi e des
F r o sch g e h irn s, B ra u sch ig 8 8 5
n w e , 1 .

V olk m ann , A . W Ub Rfl b . e er e ex e w eg u n g e n . M ul l s er
’
Ar h c i v,

1 8 38 .

V ul p ian . L eco n s su r la p hysi o lo gi e du sys t em e n e rve ux . P isar ,

1 8 66 .

W y m an , J . A n at o my f t h o e Ne rv o us S ys t m e of Ra n a p ip ien s .

Sm t ih so n an i Co n t r ibu t i s t K on o no w l dg e e, V ol 5 , . 1 853 .
3 20 TH E B IOLOGY OF TH E FROG CH A P .

Y er k es R M , h ibi t i
. d R i f c m
. In t f R ct i
o n an s i th e n or e en o ea on n e

F g R
ro , lana t J u C mp N u
c a nz z a n s . d P sych V l 4
o r
. o 9 4 . e r . an .
, o . 1 , 1 0 .

B h u g
a n n d H mmu g d
un R ti
e f t c t il R i z du ch k
n er ea c o nen au a e e e r a us

t isch R i z b i m F
e e eh A ch g
e P hys B d 7 9 5
ro sc e. r . es . .
, . 10 , 1 0 .
3 22 TH E BIOLOGY OF TH E FROG CH A P .

The ski n o f th e fr o g i s an org an o f u n usu al se n sibili t y I t .

is se n si t ive n o t o n ly to t ouch che mic al s t i muli an d differe n ces

, ,

F IG . 86 .
—T uch c
o o rp uscl e f g i t h i su p p lyi g
of th e ro w ts n n e rve . T he
b ran ch s f h
e o t e n e r ve m y b s
a emifyi g b
een r a h fl n et w e e n t e a tt e n e d c lls
e

o f th se s g
en e or an .
( F m G pp f
ro M k l )
au ,
a t er er e .

of tem perat ure but it is a ffecte d in n o sm all degree by

,

ligh t I t i s also re adily i n fl u e n ced by ch an ge s in the m ois

.

ture of th e air as is i n dic at ed by t h e be h avi o r of the an i m al

,

u n d er differe n t co n di t io n s o f th e at mosph ere .

S en se Or g an s of t h e M ou t h —I n th e m o n th there a re in
.

a dditio n to t h e n erve e n di n gs in the ge n er al e p i t h eliu m

n ume rous se n se org an s w hich h ave be e n su p pos ed to be

co n cer n ed wi t h t h e se n se o f t as t e Th ey occur o n t h e .

fl at te n ed surfaces o f th e fun gi form p ap iliae o f t h e to ngu e an d

also o n t h e floor a n d roof o f t h e m ou t h w here t hey assum e ,

a more rou n ded fo rm al though possessi n g esse n ti ally th e

s a m e s t ruc t ure in b o t h regi o n s ( B e t he ) The surface o f .

t h ese org an s i s covered by cyli n dric al e p i t h eli al cells be t wee n

which are sc at t er e d v ariou s forms o f elo ng a ted c ells which

X VI
I TH E S E NS E O RG A NS 3 23

ar e co nn ected usu ally by m e an s o f br an chi n g processes ,

wi th t h e ra mific at io n s of t h e supplyi n g n erve .

Th e Olf act or y Organ s —The se a t of sm ell is in the n as al .

c avity the w alls of which are supplied wi t h the olfactory

,

PI ex . su b b as.

FIG .
—S c t i h ugh f g l m p p i ll A bu dl f
e on t ro fib s a un t or a a. n e o n e rve er

t
e n er s t h p p ill f m b l b ki g p i fi s h sub b s l p l xus
e a a ro e ow rea n u n to r t t e a a e ,

d s c d h sub p i h li l p l xus u d h b s s f th c lls f

,

an e on t e e t e a e aro n t e a e o e e o

c lumo p i h lium
n ar Th el t p l xus f ms c
t e cti s i h h
. e at e r e or o nn e on w t t e

b ch s f h s s y c lls f
ra n e o t e d i gs ; b
en o r c lls ;
e . a, r e e n e r ve e n n
’
c , a , n e rve e

j ; c lls c ct d i h th sub p i t h li l p l xus ( F m G p p ft

,

e, e o nne e w t e e e a e . ro au ,
a er

Nie m ac k ) .

n erves The n as al c avi ty i s li n ed by a mucous m embran e

.

which is co mposed of a co n n ec t ive t issue subs t rat u m an d

an ou t er l ayer of olfac t ory epi t helium I n the regio n s t o .

w hich the olfactory n erves are d istribu t ed th e e p i t heliu m

co n sists of th r e e ki n ds of c ells : ( 1 ) in ter s t itia l cells which ,
3 24 TH E BIO L OGY OF TH E FRO G CH A P .

ar e very much elo n g ated an d cyli ndric al in form ( 2 ) ba sa l

( K r ause
) whic h are of stell a te fo rm an d li e n ext t o t h e
,

co n n ective tissu e ; an d ( 3 ) th e ol
fa cto ry cells The l a tter which .
,

represe n t the tru e se n sory cells ,

ar e v ery v ari able in form Th ey .

are u su ally very lo n g a n d n arrow ,

mu ch swolle n ou t where th e n u
c le u s occurs an d fur n ished at the ,

outer fre e e nd with a tu ft of fi n e

cili a At their i nn e r e n ds t hey are
.

dr aw n ou t i n to a fi n e process which
is co n n ected wi t h th e olfac t ory
n erve .

K n aue r fou n d th at ill sm elli ng -

subs t an ces held n e ar th e n os t rils

c aused t h e frog to t u r n aw ay the
he ad b u t wh e t h er t h e olfactory
,

se n se pl ays an i mport an t rOle in

the life o f t h e an im al is n o t k n ow n .

Th e E yes — The eyes o f the

frog are n e arly sp heric al in form ,

a n d t hey are lodged in l arge c avi

ties in the S ide s o f t h e h e ad c alled

th e orbits withi n which t hey are ,

FIG f th lf freely mov able The outer or ex

O e o ac

y p i t h l i um O t h l ft
.

tor e e n e e
posed f th e
.

s s y c lls i t h l p t O Il O ey e I S CO V
'

tw o en
O I I '

or e w Cl l

t d ds l g d ered by a tr an sp are n t membr an e

a e
u t o er en ,
en ar e

hz
f
.

d
gi fgg zg ii: fiiiiff;
a'

r
k n ow n as t h e co r n ea th e rem ai n

c lls ( A ft D gi l )
e . er in g portio n which co nsti t utes about
o e .
,

two th irds o f t h e su rface is forme d ,

by th e scler o tic coa t This i s an op aque w hi t e coveri n g

.
,

m ai n ly i n clos ed withi n the orbit ; it i s composed o f an

3 6
2 TH E BIOLOGY O F TH E FRO G CH A P
.

g ai n th e impressio n th at t he eye is pressed i n t o t h e h e ad by

th e closure of the lids The ope n i ng of th e eye o n t h e
.
,

other h an d whil e ge n er ally accom p an i ed by th e pro t rusi o n

,

of th e eyeb all through the co n tractio n of the leva to r b u lbi

muscle i s p artly effec t ed by m e an s o f an i nd epe n de n t
,

muscle th e dep ressor m embr a n x n ict ita n tis I n t h e closure

,
.

o f t h e ey e th e lo w er lid an d n ic t i t ati ng m embran e are pulled

over all bu t a sm all p art of th e exposed surface .

Th ro ugh th e tran sp are n t cor n e a m ay b e see n th e colored

ir is in th e ce n ter of w hich i s an ov al ap er t ure th e p up il
, ,
.

Just beh i n d t h e pupil lies the tr an sp are n t crysta lline len s ,

which u n like th e le n s of high er an i m als i s o f a n e arly

, ,

sph eric al form al t h ough som e w h at m ore fl at t e n ed o n t h e

,

an terior face I t i s m ad e u p of co n ce n t ric l ayers like t h e

.

co ats o f an o n io n O n i ts ou ter surface i s a S i n gle l ayer of

.

epitheli al cells t h e re m ai ni n g portio n co n sisti n g o f tran s

,

p are n t fibers whose ge n eral direc t i o n is p ar allel wi th th e

op t ic al axis o f th e eye Th e w h o l e le n s is su rrou n ded by
.

a very delic a te tr an sp are n t c apsu le which is a t t a ched a lo n g

t h e outer m argi n to th e cili ary body by m e an s o f r adi a ti n g

fibers .

Th e c avi t y o f th e eye i s divided by t h e le n s an d its su p

porti ng fibers i n t o an ou t er ch amber w hich co n t ai n s a t ran s
p are n t w atery fluid t h e a q u eo u s h u m o r an d a l arger i n n er
, ,

ch a mber filled by a more solid t r an sp are nt subst an ce k n o w n

as t h e vit r eo u s li u m o r Th e vi t reous hu mor is perm e a t e d
.

by n u merous fibers which form a sor t of su ppor t i n g n e t w ork

which co n t ai n s free cellul ar ele m e n ts an d n u m erous bl ood
vessels Th e circul atio n in th e l at t er as see n by the o p h
.

t h alm o sc o p e h as bee n d escribed by sever al observers an d

wi t h especi al full n ess by H irschberg an d S chleich .

The w all of th e pos t erio r ch a mbe r o f t h e eye is form ed

‘

o f three tu n ics t h e scler o tic co at which h as bee n describe d

, ,
 XVI I TH E S E NS E O RG A NS 3 27

a bove the c/z o r o id an d t h e r etina

, ,
. Th e is a vas
clz o r o id

c u lar pigm e n ted l ayer lyi n g n ext

,
to t he sclerotic Next .

Corn ea
Cam . an t er .

Z on . cil .

L en s cr y st .

Cor p . vi t t .

S clerae

F IG . 89 .
—S c t i
h ugh th p ic l xis f h y f th f g C m
e on t ro e o t a a o t e e e o e ro . a .

t
a n er u ch mb c i i g h qu us hum
o ter a C mp t i
er o n ta n n t e a eo or ; a os n n er

ch mb c i i g t h i us b dy ; C p il cili y b dy ; C p t
, ,

a er o n ta n n e v t re o o or c ar o or . vi r ,

i us b dy ; P t il d P fi b S l c t il g i us d fib us
.
,

v t re o o . ca r . an r os a c er ae, ar a no an ro

l y s f h scl t ic c t ; Z il z ul cili is ( F m G p p f
.

a er o t e e ro oa on c .
,
on a ar . ro au ,
a t er

G R t zius )
. e .

wi t hi n the c h oroid li es th e r etin a which is co n ti nu e d ante

r io rl
y over t h e posterior su rfa c e of th e iris as fa r as th e pupil .

I t is compose d o f t w o pri n ci p al l ayers an outer thi n l ayer

.

o f pigm e n ted c ells an d an i nn er much thicker l ayer w hich ,

forms the se n si t ive portio n of th e reti na O ver th e cili ary .

body an d posterior S ide o f t h e iris th e reti n a is much thi n n er

t h an it is elsewhere an d does n o t co n t ai n n ervous tissu e .
3 28 TH E BIO LOGY OF TH E FROG CH A P .

Th e opti c al por t io n o f the reti na w hich li n es the whole p o s

t e rio r regio n o f the eye an d ex t e n d s as far for w ard as th e

cili ar y body is a co m plex structure formed m ai n ly of n ervou s

,

elem e nts I t is th e p a r t of t h e ey e which i s es p eci ally sen

.

sit ive t o ligh t an d upo n whic h im ages of ex t er n al obj ec t s are

throw n I n addi t io n to th e ou t er pigm e n t ed epi t heliu m it i s

.

c o m p o se d a s in t h e high er vert e b ra t es o f n in e l ayers the l atter

, , ,

fo rmi ng the i nn er or thicker str at u m o f t his structure These .

l ayers t ake n in ord er fro m wi t hi n outw ard ar e as follows

1 . The in ner lim iting mem b r a n e a thi n suppor t i ng m e m ,

bran e lyi n g n ext t o t h e vi t reous hu m or I t i s co nn ec t ed wi t h .

elo n gat ed c ells ( M uller s fibers) which exte n d to th e ou ter

’

li mi ti n g m embra n e an d give o ff br an ches which form a sor t

o f supporti n g n et w o rk for t h e n ervous ele me n t s of th e reti n a .

2 . A l ayer of n er ve fibers forme d by the r a mific atio n s o f

,

th e o p t i c n erve .

3 . Th e in n er g a ng lio n ic la er form ed by comp a r a t ively

y ,

l arge g an glio n cells w hose processe s ex t e n d i n t o th e pr eced

in g an d follo w i n g l ayers .

4 Th e in n er r eticu la r lay er formed by a n etwork of

.
,

n erve fibers .

5 . Th e in n e r n u cle a r lay er composed,

of sever a l l ayers of
gan glio n cells whose processes ex t e n d i n to bo t h of th e
adj ace n t l ayers .

6 The o u ter r eticu la r lay er a n arrow l ayer formed by a

.
,

n e t w ork o f n erve fibers d erived from t h e i n n er n ucle ar l ayer

an d th e l ayer o f rod s an d co n es .

7 . The o u t er n u clea r lay er co n t ai n s th e n u cle ated por

t io n s of th e cells formi n g th e rods an d co n es an d a few

isol ated g an gli a .

8 . Th e o u ter lim iting m emb r a ne a very thi n supporti n g ,

m e mbran e p erforat ed in n um erous pl aces by t h e cells o f th e

,

outer l ayer .
 TH E BIOLO G Y OF TH E FRO G CH AP .

por t io n o f t h e rods Thi s i s follo w ed by a refractive le n s .

sh aped body beyo n d w hich t h e co n e bec o me s co n s t ricte d

,
.

F IG .
91 c i s h ugh h u p t f h f g s i i t h it p ig
-
Se t on t ro t e o t er ar o t e ro
’
re t n a w s

d p i h l i um T h figu h l f is k f m f g hich h d
.

m en te e t e . e re o n t e e t ta en ro a ro w a

b x p s d d yligh f fi h u s T h p c ss s f h h u t
ee n e o e to a t or ve o r . e ro e e o t e t ree o er

p igm t c lls xen di d h x

e l lim i i g m mb
e te n d h nw ar to t e e t e rn a t n e ra n e , an t e

p igm t th y c en i h st m d ie d su u di g h ds d
o n ta n as re a e nw a r rr o n n t e ro an

c s ly t h i b s T h c s s gly c d h t
,

one n e ar to e r a e . e o ne a re t ro n r e t ra te so t a

h i u s gm s li h l i mi i g m mb T h fi gu
,

t e r o ter e en t h e n ea r t e t n e ra n e . e re o n t e
righ t is k f m f g th h d b k p f f y i gh h u s i
ta en ro a ro at a ee n e t or o rt -
e t o r n

d k ss T h p igm i h p c ss s f h p igm t c lls is d

ar ne e en t n t e ro e e o t e en e ra w n

b ck t d h c f h c lls d h c s x dd u d
.

a o vr a r t e e n te r o t e e an t e o ne a re e te n e o t w ar
b ucl i f h d d c c lls sh b l
,

h
et w e e nd Th
t e ro s e n e o t e ro an one e a re ow n e ow

h l i mi i g m mb ( F m G pp ft V d
.

t e t n e G S )
ra n e . ro au ,
a er an en e re n t o rt .

Th e n ucleu s lies in an e n l arge m e n t o n t h e i n n er S ide o f t h e

ou ter li mi t i ng membr an e Th e i nn er e n d of t h e c o n e like .
,

t h a t o f the rods ru n s ou t i n to a fiber w hich e n t ers t h e ou te r

,

r eticul ar l ayer The co n es v ary in S ize an d fo r m as m an y

.
,
XV II TH E S E NS E O RG ANS 33 1

as three v arieti es bei ng disti n guish e d ( M S chul tze van .

,

G e n dere n S tort ) S ome of thes e are m o v able exp an di n g

.
,

F I G 92 I n n ersu f cr a e of i
th e n n er h lf f th y f R
a o e e e o a n a es cu len ta sh ow

c t lis ligh c l d b d b ill ic

. .

in g t h e t o ve t h e a opt
'

e n ra as a o o re an a
a re a p ap a
-

w h ere th e o p ict n e r ve e n t er s .
( A ft e r G a upp . )

in the d ark u n t il t hey m ay re ach the periph ery an d co n tr act ,

in g u n der t h e i n flu e n ce o f ligh t O rdi n arily mo s t o f th e .

co n es do n o t re ach t h e outer surface of t h e re t i n a .

Th e r eti n a is m arked by a t hicke n i n g or p apill a w h ere th e

optic n erve e n t ers Abo ve this p apill a t h ere ex t e n ds in an
.

a n tero posterior di rec t io n a t hi cke n ed portio n o f the re t i n a

-

kn ow n as t h e a r ea cen tr a lis w hich prob a bly m arks th e ,

regio n of most acu t e visio n like the y ellow spot in t h e eye ,

o f m an The co n es are rel at ively more n u merou s in thi s

.

regio n t h an in o t her p ar t s ( 1 c 2 4 1 i n s t e ad o f 1 c 3 . . . .

or 4 r as elsewh ere ) an d th e rods are of rel atively sm all

.
, ,

size .

Th e actio n of ligh t u po n t h e re ti n a results in t h e produc

ti o n of sever al visible ch an ges I n addi t i o n to the co n tr ac t io n .
33 2 TH E BIOLOGY O F TH E FRO G CH A P .

of th e co n es w hich h as bee n me n tio n ed above th ere i s a ,

ble achi n g o f th e pigm e n t in th e ou t er e n ds o f t h e I O dS ‘ '

this t akes pl ace more r apidly in th e visu al purple t h an in the

gre e n pigme n t Aft er a short exposure in t h e d ark bo t h pig
.

me n ts are restored eve n in the extirp ated eye I t is prob

, .

a ble th a t the ch emic al ch an ges produced in the re ti nal

pigme n ts t hrough t h e i n flue n ce o f ligh t pl ay som e p art in
,

t h e s t i mul at io n of t h e se n sitive cells L igh t also affects th e

.

cells o f t he ou ter pigm e n t ed l ayer of th e reti na c ausi n g t h e ,

pigme n t to s t re am o u t i n to the processes which exte nd

betwe e n th e rods and co n es Wi t h i n te n se illumi na tio n the
.

pigme n t m ay spre a d as far as th e i nn e r li mi t i n g m embr an e ,

in fro m t e n to fift ee n m i n utes which is a bou t t h e t i me which

,

i t t akes th e visu al purple to dis appe ar u n d er the s am e

co n di t io n s ( B oll Angellu c ci)
,
.

Th e amou n t of ligh t falli n g upo n th e re t i na i s regul ated by

the iris whic h acts as a sor t o f scree n or curtai n ad mit t i n g ,

t h e r ays o n ly through its c e n tr al aperture the pupil Th e

,
.

iris lies direc tly in fro n t o f th e le ns I t s ou ter m argi n is

.

j oi n ed to t h e cili ary body behi n d an d th e j u n cture o f th e

sclero t ic an d cor n e a in fro n t I t co n t ai n s n umerou s bl ack
.

pigme n t cells and several golde n o n es whic h gives i t i ts

brigh t yellow spots Ne ar th e m argi n o f t h e pupil there i s a
.

ri ng of smoo t h mu scle cells formi n g a sp /z in cter p up illce which

c auses the pupil to co n t r act S t ro ng light c auses a c o n t rac
.

tio n o f the pu pil an d co n seque n t ly a dimi nu tio n of the

a mou n t of light t h a t falls o n the reti n a thus checki n g t h e
,

excessive s t i mul at i o n o f the se n sitive p art o f the eye This .

effect is produced at l e ast in p ar t th rough t h e direc t actio n

, ,

o f t h e ligh t upo n th e m uscle cells of th e iris si n ce it occurs in

,

eye s th a t h ave be e n removed from the body ( S t ei nach G u th ) ,

.

The o ptic al arran gem e n ts o f the eye of t he frog are suc h

as to th ro w i m ages of obj ects upo n t h e re t i n a Th e se n si.
33 4 TH E BIOLO G Y OF TH E FROG CH A P .

which r ot ate s the eye arou n d i ts optic axis so th at its uppe r

m argi n tur n s forw ard an d an infer ior obliqu e which pro , ,

du ces the r everse moveme n t Th e eye is pulle d i n to t he .

orbit by me an s o f th e stro n g r et r a cto r b u lbi muscle which ,

arises fro m t h e an gle be twee n the l a ter al an d medi an por

tio n s of the p ar asphe n oid bo n e and i s b ro adly i n serted upo n

the posterior an d m edi an sides of th e eyeb all The eyes .

FIG 93 .
—Righ t m mb us l by i t h
e f t h f g f m th i
ra n o sid a r n o e ro ro e n n er e

i x l d p s i m p ull ; u f
. .

t ter o r a
aa, pa o, a a n t e r o r, e er na
p , an o a an , a e rt re o

u t iculus ; i x l d p s t i s mici cul c ls ;

,

r p ca , ce , c ,
an t e r o r , e t ern a ,
an o e r or e r ar an a

t i l s ccul
cus , u r c u o c l ; d duc t us d lym p h icus ; l l g ; m
-
a ar an a e, en o at a en a u

m cul cus ic f h c ssus u t iculi s cculus d p s

, ,

m ms, a, a a a t a o t e re e r , a an ar

gl c t sp c t i ly ; pb p s b sil is ; p l pp b p p i ll cust ic l g
,

ne e a re e ve ,
ar a ar a a a a a e n ae

d b sil is ; b b ch s f ud i y
, ,

an a p
ar l
r aa , ra , t
r s, rn, r , r ra n e o a to r n e r ve o

i m p ull p s t i m p ull s cculus m cul gl c t l g

,

t
an e r o r a a, o er o r a a, a a a ne e a, a en a ,

d p s b sil i s s p c t i ly ; s cculus ; p m u t icul i su p i

,

an ar a ar re e ve s, a ss , s s us r er o r

d p st i u t i culus ( A ft G R t z ius )
,

an o
; er o r u, r . er . e .

are pushed outw ard by the co n tr actio n o f th e leva to r b u lbi ,

a bro ad sheet of muscle r u nn i n g obli quely across th e v e n tr al

sid e o f the orbi t .

Th e E a r — I n th e ear of th e frog there are t w o sets o f

.

org an s which are fu n d ame n t ally differe n t in origi n an d in

fun ctio n T h e o n e co n sti t utes th e in ner ea r which fo r ms
.
xv1 1 TH E S E NS E O RG ANS 33 5

th e se n sory app ar atus th e other the middle ea r bei n g com , ,

posed of accessory s t ruc t ures for th e tran smissio n of sou n d

w aves to th e se n sitive p art o f th e org an .

Th e i nn e r ear is co mposed of th e m embr an ous la by rin th,

which lies w ithi n the auditory c apsule which is form ed by ,

the pro lifi c an d exoccipit al bo n es of th e skull Th e l aby .

rint h i s a complic at ed sac formed origi n ally by i nv agi n atio n

of th e ectoderm of th e sur face of the b o dy fro m w hich i t is

subseque n tly co n s t ricted off I t is divisible i n to an upper
.

por t io n t h e u tr icu lu s an d a lowe r sm aller p ar t th e sa ccu

, , , ,

lu s. The form er is an oblo ng sac lyi n g n e a rly h orizo n t al .

I t gives rise to the three semicir cu la r ca n a ls wh ich lie in ,

pl an es approxim ately at rig ht an gles t o e ach other Thes e .

c an als are m embr an ous t ubes embedde d wi t hi n th e c ar til age

o f the auditory c apsule an d co mmu n ic ati n g with the utriculus
at each en d .Ther e is an an t erior c an al w hich lies in a
v ertic al pl an e a l at er al c anal which lies in a n e arly horizo n
,

t al pl an e o n the ou ter S ide of t h e utriculus and a posterio r ,

c an al which exte nd s tr an sversely n e arly at righ t an gles to the

t w o oth ers . At o n e end e ach of th e c an als is fur n ish ed with
an e n l argeme n t or a mp u lla whic h co n t ai n s an i mport ant
, ,

se n so ry app ara tus .

Th e saccu lu s or lower divisio n o f th e l abyri n th is an

, ,

irregul ar ov al sac whi ch proj ects d ow nw ard an d forw ard

fro m th e utriculus wi th which its i n terior is co nn ec ted
,

through an aperture There are fou r sm all o u tp o c ke t ings

.

from the b ase o f th e s acculu s an d from the m edi an S ide ,

th ere is give n off a n arrow tube t h e du ct u s en doly mph a ticu s

, ,

which exte n ds dors ally wh ere i t pe n etr at es t h e cran i al c avi t y ,

a n d e n ds in a l arge sac th e s accu s en dol m h a ticu s which

, y p ,

is filled w ith a milky white fluid co n t ai ni ng n u merous w hite -

crys t als o f c arbo n at e of li me Th e utriculus the sac .

,

cu lus an d the l arge r o u t p o c ke t in s of th e l tter co n t ai n

, g a
3 3 6 TH E BIOLOGY OF TH E FROG CH A P .

m asses of si mil ar cryst als which ar e commo n ly c alled th e

The differe n t p arts of the ear are all supplied by th e

bran ches of th e audito ry n erve I n th e regio n s o f the n e rve
.

e ndi ngs t he epi theliu m co n t ai ns se n sory cells with h airlike

p roc esses a t thei r outer e n d s These ar e esp eci ally abu n
.

d an t an d well d eveloped in t h e w alls o f th e a mpull ae Th e .

whole l abyri n th i s filled with a fluid c alled th e en doly mp h ,

a n d it is su rrou n ded by an oth er fluid the p er ily mp h which

,

fills the sp ace betwee n th e l abyri n t h an d th e w all o f th e

a udi t ory c apsule O n th e ou ter side t h e w al l o f t h e audi
.

tory c apsule is perforat ed by an aperture the fenes t r a ova lis , ,

which is closed wi t h a s m all plug of c artil age .

Th e accessory auditory app a ratus co n sis t s o f th e structures

which in higher forms co n s t i t u t e th e middle ear th ere bei n g ,

n othi n g in th e frog which correspo n ds to the exter n al e ar of

m am m als . The c avity o f the m iddle ear com mu n ic at es

through th e E ustachi an tube wi th the mouth c avity n e ar th e
an gle o f t h e j aw I t i s closed exter n ally by th e ty mp a n ic
.

m e m bra n e which is re adily se e n at th e side of th e h e ad

,

behi n d t h e eye This m embr an e is n e arly circul ar in form

.

an d is att ached by i ts ou t er m argi n to a ri n g o f c artil age the ,

a n n u lu s ty mp a n icu s Ne ar i ts middle it gives at t achm e n t

.

to the outer h e ad o f a rod th e colu mella w hich exte n d s

, ,

across th e c avi ty of th e middle e ar a n d j oi n s th e sm all c a r

,

t ilage lyi n g over th e fe n es t r a ov alis I t is through this rod

.

t h a t th e vibr a tio n s which ar e produced by th e sou n d w aves

i m p i ngi n g upo n th e tym p anic m e mbran e are c arried t o th e

i n n er ear H ere th ey se t t h e co n te n ts of th e l abyri n th i n to
.

vibrat io n an d thus sti mu lat e the se n sory en d org an s of t he

aud it o ry n erve . Th e n ervou s i mpulses se t up in this w ay
a r e c o n v eyed to t h e br a i n wh ere they give ris e to the se n s a
,

t io n o f S ou n d .
338 TH E BIOLOGY OF TH E FROG CH A P .

rem ai n in this at ti tude w i t h i ts muscles in a s t at e o f t o n ic

,

co n tractio n oft e n for h alf an hour A n orm al frog of t h e .

sp ecies experi me n ted with ( R es cu len ta ) do es n o t S ho w

.

this respo n se except in a slight degre e a n d th e n o n ly aft er ,

it h as beco m e fat igued F rogs d evoid of otoli t h s were

.

fou n d to u t ter t h e so c alled p ai n cry upo n S ligh t provo

-
”

c atio n where a s thi s respo n se is r arely elici t ed fro m n orm al

,

s p eci m e n s L oco motio n m ay t ake pl ace in a n orm al m an

.

n er b u t th e muscul a r t o n us seems to be lo w a n d the an i m al

, ,

quickly becomes fatigued The compe n s atory mo tio n s are

.

u n affec t ed .

F rogs h ave a well d eveloped s e n se o f h e ari ng al t hough

-

th ey do n o t usu ally m an ifes t i t by a ny cle arly evid e n t S ig n .

I n cro aki n g o n e frog frequ e n t ly respo n ds t o t h e cro ak of

,

an o t h er i n divid u al so th a t o n e o ft e n h e ars a p a ir an s w eri n g

,

e ach other by regul arly alter n ati n g n otes I f aft er a co n cer t .

,

o f frogs has bee n S ile n c ed by som e o n e who i n t rudes too

n e ar their h au n ts o n e i n divi du al ve n t ures to resu me its
,

cro aki ng i t is speedily follo w ed by o n e a ft er a n o t her o f its

,

comr ades L an dois tells of a tree frog whic h h e kept in

.

c aptivi t y whic h would give an answeri ng cry every ti me th at

he w ould i mit ate it s n ote Y erkes observed t h a t wh e n h e
.

c aused a frog t o cro ak by rubbi ng it s sides th e other frogs ,

u nder observ at io n occ asio nally g av e sig n s o f at te n tio n by

s t r ai ghte n i ng up an d r aisi ng th e h e ad as i f lis t e n i ng The .

s ame observer n oticed whe n c arryi n g o n experi m e n ts with

,

frogs in a l abyri n t h th at t h e an i m als oft e n g ave sig n s of

,

he ari ng th e sou n d m ad e whe n o t her i ndividu als j umped i n to

the w at e r Th ey would str aigh t e n u p an d hold the liste n
.

in g o r at t e n t ive at ti tude for so me seco n ds As the an i m als .

could n o t se e o n e an ot her there i s good evid e n c e o f their

,

abili t y to h e ar th e spl ash m ade by a frog w he n i t strikes the

w ater . This accordi ng to Y erkes expl ai n s the fac t t h at

, ,
 TH E S E NS E O RG ANS 339

“i t is n ever poss ibl e to get n e ar to an frogs in t h e s am e

y
regio n aft er o n e has j u m p ed in The spl ash sou n d is sig
.
”

n ifi c an t to t he m an d pu t s t h e m o n th eir g u ard .

O n th e o ther h and m an y other S ou n ds v aryi ng gre atly

, ,

in loud n ess and pi t ch do n o t elici t any m arked respo n s e

,
.

O n e m ay appro ach t o withi n a fe w feet o f a gree n frog or

a bullfrog an d m ake all s o rts o f n ois es wi thou t c ausi n g i t to

give any S ig n s of u n e asi ness Just as soo n however as a

.
, ,

quick m oveme n t is m ade by the observe r the an i m al j umps ,

.

S ou n ds like t h e spl ash o f a plu n gi n g frog or t h e cro ak ,

or p ai n scre a m of an other m e mber of the s am e species ,

serve as w ar n i ngs but th e an im als do n o t j u mp i n to t h e

,

w a ter u n til t hey see so m e S ig n of an u nusu al or d an gerous

obj ec t
.
”

I t mus t n o t be i n ferred th at frogs do n o t he ar a g r e at

v ariety o f sou nds sim p ly bec ause they give m an ifest sig n s
o f atte n di n g t o o n ly a fe w sou nd s in w hi ch they h ave som e
p articul ar i n teres t Y erkes fo u n d t h at frogs whi ch give n o
.

o t her S ig n s of perceivi ng sou n d S how a differe n ce betwee n

,

th e r ates of th eir respirat ory move me n ts be fore an d aft er th e

sou n d is m ad e The sou n d of a tu n i ng fork falli n g w ater
.
, ,

a shrill w his t le th e ri ngi n g o f a bell an d oth er n oises were

, ,

em p loyed S ome of thes e produced little o r n o effec t T h e

. .

shrill w histle an d th e ri n gi n g of a bell c au sed a decre ase in

th e r ate of respira tio n o w i ng perh aps to fe ar while t h e
, ,

sou n d of falli ng w at er c aused th e r a te o f resp ir atory move

me n ts sligh t ly t o i n cre ase Th e gree n frog (Ra n a cla m ita ns)
.

w as fou n d to respo n d to sou nds v aryi n g in p itch betw ee n

fifty an d t en thous an d V ibratio n s per seco n d .

Th e re ac t io n t i m e o f frogs to visu al stimul i i s also in flu

e n c e d by sou n d . Y erkes fou n d th at frogs which were pl aced
in a gl ass aqu ariu m so surrou n d ed t h at t h e moveme n ts of th e
obse r ve r could no t be detected would j u mp vigo r ously at
,
340 TH E BI OLOGY OF TH E FR O G CH A P .

a sm all red c ard w hich w as m oved n e ar the m I f a tu n i ng .

fork w as sou n ded j u st be fore t h e c ard w as prese n ted it ,

b ec a m e evid e n t th at th e sou n d put the frog o n th e alert ,

an d wh e n the obj ect c am e i n to v i ew it j u mped at it more

, ,

quickly an d a gre ate r n umber o f ti m es th an whe n th e visu al

s t imulus w as give n withou t t h e auditory Wh e n the red .

c ard w as S how n i t w as ofte n sever al seco n ds before the frog ,

would n otice it an d a ttempt to get it but w h e n the sou n d ,

a lso w as give n th e an im al usu ally n o ticed an d j um ped tow ard ,

t h e movi n g c ard almos t i mm edi ately I t is prob able th at

”
.

t his h abit of ge t ti n g i n to r e adi n ess for a spri n g u po n he ari n g

a sou n d n e ar by i s o f v alue t o th e frog si n ce i n sects an d ,

other cre a tures th at serve aS fo o d ofte n m an ifes t t heir pres '

e n ce by som e sort o f n oise before th ey co m e i n to th e frog s ’

fi eld of vi sio n I f t h e frog prep ares hi mself he is more ap t .

,

to sei z e his prey wh e n it appe ars .

RE F E RE NC E S

A ch , U b di O t lit h f t i
N . d d L b y i t ht e er e o e n un c on un en a r n o nus .

Ar ch g P hys B d 8 6 9
. es . .
, .
, 1 01 .

B r T Di A cc m m d t i d A g s b i d A m p h ibi
ee , . A ch e o o a on es u e e en en . r .

g es . P hys B d 73 8 9 8 .
, .
,
1 .

G ir ar d H R ch ch s l f
, c t i d c ux s m i ci cul i s
. e er e su r a on on es an a e -
r a re

de l ill i t ch z l g ui ll A ch P hys N m t P t h
’
o re e n e rn e e a re n o e . r . . or . e a .

T .
4, 1 892 .

G olt z , F d u tu g d B g g g d
. Ub e er die p hysi l gisch o o e Be e n er o en an e es

O h l by i t hs A ch g
r a r n 87 B i t ag L h
. r . es P hys B d
. .
, . 2, 1 0 . e r e z ur e re vo n

den Fu k ti d N n t d F sch s B li 8 6 8
o n en er e rv e n c e n r e n es ro e . er n, 1 .

H ir ch b er g J Z V gl ich d O p h t h l m s k p i A ch A t
s , . ur er e en en a o o e . r . na .

u P hys p hys A b t h 8 8 Z D i p t i k d O p h t h lm sk p i d
. .
, . .
, 1 2 . ur o r un a o o e er

F isch d A m p h ib i un
g l 8 8 e nau en . . c .
,
1 2 .

L an dois , H . K On n e n F ro sch e h Ore n ? 2

5 J ah re sb e r . w est fal . P r o v.

V er .
,
1 8 97 .

L au den b ach . Z ur O t o lit h e n frag e . Ar ch .

g es . P hys .
,
B d 7 7, . 1 8 99 .

L y on , E . P . A Co n t r ibu t i on to the Co mp arat v e i P hysi l gy o o of

Co mp s e n at o r y M ot i s on . Am .
Jou r . P hys .
,
V ol 3, . 1 8 99 .
3 4 2 TH E B I OLOGY OF TH E F ROG cH A P .

C H A PTE R XV I I I
I NS TI NCT S AND T ROP I S M S A S RE L AT E D T O RE F L E X
A CT I O N

W E h ave al r e ady tre ated of so me o f th e re flex actio ns of

the frog an d h ave show n th at th ey exhibit a purposive ch ar
a cter oft e n in a v ery striki n g d egree
,
Th ey are far fro m
.

bei n g m ere r an do m respo n ses to stim uli ; n otwithst an di n g

t h e fact th at they m ay t ake pl ace i n depe n de n tly of in t elli

ge n t co n trol I t is ge n er ally recog n ized th a t t here i s a

.

close co nn ectio n betwee n re flex actio n s an d i n s t i n cts it ,

bei ng in fact very difficult to dr aw th e li n e betwee n th e m

, ,
.

Th e beh avior o f an an im al is u su ally c alled i ns ti n ctive whe n

it t akes pl ac e withou t previous i n struc t io n an d with n o c o n
sc io u sn ess of the en d to which i t is directed A digge r w asp
.
,

for i n st an c e m akes a n est of a p articul ar type in t he grou nd

, ,

c a tches o nly cert ai n species of i n sects which it sti ngs in the

,

ve n tral gangli a so as to produce p aralysi s wi thou t c ausi ng

,

de at h an d a fter d eposi t i n g an egg u po n it buri es its prey

, , ,

in the n est as provisio n for it s future o ff spri n g The w asp .

i s u t terly u n co n scious of the sig n ifi c an ce o f its comple x

be h avior ; o f the w riggli ng grub which it h as l abored so
i ndustriously to provisio n i t k n ows n othi n g an d c ares less ;
i t is i mpelled by bli n d impulses to a p ar t icul ar li n e o f ac t iv
it y which al t h ough o f n o servi ce to th e i n dividu al w asp is
, , ,

esse n t i al to th e co n t i n u an ce of the r ace I t i s guided n eith er

.

by previous experi e n ce n o r by i mi t atio n an d has n o b asis

,

for dr awi ng any co n cl usio n r eg ardi n g the u t ility of its co n

XV I II I NS T I NCTS A ND T ROP I S M S 34 3

du ct eve n were i t c ap able of so doi n g S uch beh avior

,
.

affo rds a t ypic al illus t r a t io n of i n sti n ct ; an d throughou t t h e

an im al ki n gdo m it is i n s t i n ct which i s the domi n an t el eme n t

in co n duct .

The beh avior o f th e frog is almost e n tirely m ad e up o f

i n sti n ctive actio n s Nature has equipp ed this an im al wi t h
.

t h e m e an s of getti n g through th e world wi thout relyi n g to ,

an
y gre at exte n t upo n t h e lesso n s o f exper i e n ce
,
The frog
.

h as i n tellige n ce of a rudim e n t ary sort to be sure but it , ,

pl ays a very subordi n at e rOle in sh api n g the cre atu re s c o n ’

duc t . I t is truly a m arvelous thi ng t h at an an im al sho uld

be e n dowed wi t h the power o f su ccessfully ad apti ng its c o n
d uct t o a complex e n viro n me n t withou t any perce p t io n of
th e co n seque n ces of its actio n s H o w c an th e exis t e n ce
.

of such a po w er be expl ai n ed or brough t i n to rel ati o n with

,

our k nowle dge o f th e o t he r fe atures of th e an i m al s life ? ’

I t w as formerly custo m ary to reg ard i n s t i n c t as a property

su ig ener is somethi n g h avi n g n o n ecess ary affili atio n with th e
,

other fu n ctio ns of t h e org an is ms a sort of power with which

,

an im als are mys t eriously e n dowed for their guid an ce .

I t is bu t an oth er illustr at io n o f the effect of i n cre asi ng

k n o w ledge in bri ngi n g differe n t fields o f biologi c al i n quiry
i n t o closer an d more org an ic co nn ectio n th at th e i n sti n c t ive
b eh avior o f an i m als is n o w show n to be more i n ti m at ely
co nn ected wi t h their structure an d physiologic al activi t ies .

I n s t i n ct is but a ph ase of th e ge n er al li fe pro cess exhibiti n g

,

t h e s a me purposive n ess th at is S ho w n in the activi t ies of th e

he art or alime n t ary c a nal All p arts of th e body are c o n

.

t in u ally respo n di ng to stimuli in w ays th at ar e b e nefici al t o

the org an ism Whe n t he sto m ach pours out i ts secretio n

.

an d begi n s it s perist al t i c moveme n ts u po n th e receip t of

food an d allo w s t h e m at eri al whe n digested t o esc ape
through the p y lorus i t is p erformi n g actio n s which we do
,
344 THE BIOLOGY OF TH E FROG CH A P
.

no t com mo n ly c all i n s t i n ctive but w hich are as well ad apte d

,

to t h e e n d ach ieved as t h e d ivi ng of a frog u po n th e a p

pro ac h o f an e n emy or i ts burrowi n g i n to the mud in cold
,

we ath er All o f thes e actio n s are purposive respo n ses to

.

s t imuli Those of t h e stom ach we c all re flex acts while th e

.
,

divi n g an d burrowi ng i n t o the m u d are u su ally c alled in

st inc t s . Th e ch ief disti n ctio n be t wee n t h e two is th at th e
o n e i n volves th e actio n m erely o f a p art whil e in the oth e r
,

th ere i s a respo n se by t h e org an is m as a w hole There are .

so m any i n termedi ate ty p es o f re actio n howeve r th at i t , ,

i s n o e asy m atter to decide how som e of the m should be

cl assed I f a frog wi t h draws it s foot wh e n i ts t o e i s st im u
.

l ated w e c all th e ac t re flex b u t how sh all w e d esig nat e t h e

, ,

a c t of bri n gi n g t h e foo t for w ard t o wipe aw ay a drop of acid

fro m t h e S id e o f the body ? AS w e h a ve see n th e l at t er ac t ,

m ay be performed by a frog whose S pi n al cord i s cut across

n e ar t h e br a i n ; i f t herefore we c all the ac t io n re fle x wh at
, , ,

S h all we say of t h e strugg les of a frog whe n a fter bei n g ,

picked u p in th e fi nge rs it uses bo t h hi n d legs to push

,

ag a i n st th e h an d a n d a t th e s a me ti me i n fl ates th e lu n gs
,

w i th a ir c ausi n g the body to s w ell ? Th ese struggles to es

,

c ape will t ake pl ace in a frog which h as lost th e gre a t er p art

o f i ts br ai n b u t n ever t h eless t hey woul d I t hi nk ge n er ally
, , , ,

be reg ard ed as i n sti n c t ive actio n s The use o f th e hi n d

.

limbs a nd t h e swelli ng o f the body m ay be reg arded as t w o

co m plex re flexes excited by the s am e c ause The frog is so .

org an ized as to respo n d to seizure by two me t hods w h ich

c o Op erat e to effect it s esc ape both of which are reflexly
,

brought i n to pl ay .

M an y o f the m ore compl ex i n sti n cts o f the frog m ay be

resolved i n to a series of re flex ac t s We h ave see n t h at t h e
.

te n de n cy of th e m ale frog to cl as p th e fe m ale duri ng t h e

bre edi n g se aso n d epe nds u po n t h e re flex i rri t abili t y o f
3 46 TH E BIOLOGY OF TH E FROG CH A P .

m an n er by pl aci ng th eir bodi es so th at they face th e r egio n

o f stro ngest illumi natio n an d freque n tly also movi n g tow ard
,

i t A n i m als which ori e n t th emselves to light in this m ann er

.

a re s aid to be p h o to t act ic o r h elio t r op ic Those whic h move

.

to w ard th e light are c alled positive ; those whic h move

a w a y fro m i t n eg ative .This orie n t atio n is ge n er ally r e
g arded as brought about in a re fl ex m ann er through the
u n equ al stimul a tio n of t he t w o S i des of the body ei t her ,

t hrough th e eyes or other p arts o f th e org an ism Wh e n t h e .

body is obliqu e to t h e r ays i t receives more sti mulus from

,

th e ligh t o n o n e S id e th an t h e oth er an d i f th e ligh t h as any

,

directive e ffec t u p o n th e an im al s m ove m e n ts i t will n atu

’
,

r ally c au se th e body t o t ur n u n til equ ally illu mi nated o n the

t w o S id es ; the n as bo t h sid es receive th e s ame am ou n t of
,

s t i mul atio n t h e an i m al te n d s to go ei t her tow ard or a w ay

,

fro m t he light in a s t raight li n e .

G r aber w ho experi m e n ted w ith R a n a escu len ta c a m e to

, ,

th e co n clusi o n t h at t his speci es is n eg at ively photot a ctic .

The speci m e n s w ere pl aced in a box in which were two

comp artme n ts o n e o f which w as d arke n ed while th e oth er
,

w as exposed t o diffuse dayligh t Th e an im als S howed a

.

te n de n cy to collect in th e d arker o f t h e two comp artm e n t s .

I f t he frogs were give n a choice bet w ee n red an d blu e they ,

collected u nd er t h e red light .

Th e respo n ses of R a n a p ip iens an d R cla mita ns t o .

ligh t h ave be e n s t udied by M is s T o relle w h o arrived a t re ,

su lt s qui t e differe n t fro m th ose o f G r a ber F rogs pl aced in .

a box o n e h alf of w hich w as exposed t o d iffuse ligh t w h ile ,

th e o t her h alf w as sh aded m oved i n to th e ligh t e n d of th e

,

box an d orie n ted t h e body so as t o fac e t h e i n co m i ng r ays .

Whe n o n e h al f o f t h e box w as exposed to di rect su n ligh t ,

t h e frogs fi rst m oved i n to t h e illumi n ated are a an d t he n , ,

aft er a sho r t tim e re tre a ted i n t o the S h ad e where th ey sat

, ,
XV I II I NS T I NCT S A ND T ROP I S M S 3 47

with th eir h e ads poi n t i ng to w ard the light The s ame .

result occurred whe n th e he at r ays were elimi n ated by

p assi n g the ligh t through a vessel of w a ter be fore i t e n t ered
the box showi n g th at it w as n o t th e h e at alo n e t h at c aused
,

th e frogs t o retre at i n to t h e sh ade I f light be admi t ted .

fro m below which m ay be do n e by m aki n g t h e floor of t h e

,

box of gl ass th e frogs le ap i n to th e ligh t ed are a as before

, .

I f t h e whole lo w er side o f th e box be ex p osed the ani m al ,

t akes a n orm al resti n g p osi t io n but if a h alf or two t hirds of

,

i t be covered the frog m o ve s tow ard t h e light an d the

,

body assu mes a gre at er an gle to the horizo n t al th e an gle ,

i n c re asi n g the sm aller th e are a through which th e light

,

e n ters Wh e n light is thro w n u po n the frog from above

.
,

t h e an terior p ar t o f th e body beco mes r aised M iss T o r elle .

fou n d th at wh e n a frog w as pl ace d in a t all gl ass cyli n der

the botto m and sid es o f which were cov ere d with bl ack
cloth “ the b o dy w as r aised so th at the fore legs were as
,

n e arly as possible at right an gles to th e horizo n t al bo t to m

o f the j ar This m ade th e i n cli natio n of th e body 6 0 or

.
°

over F reque n tly the frog assu m ed a n al most erect posi

.

tio n by m e an s of pl aci ng the fore feet ag ai n st the sid e o f

,

th e j ar .

As these an d other exp eri me n ts S how the frog h as a ,

stro n g t e n de n cy to pl ac e its body so a s to face t he light ;

y et n ot w i t hst an di n g i ts m arked orie n ti n g respo n se it m an i ,

fests a stro n g proclivi ty to seek the S h ade F rogs pl aced .

out of doors n e ar the sh adows of trees or buildi ngs soo n

, ,

hop i n to the sh ade an d rem ai n there eve n if th ey h ave to

travel a t right an gl es to th e r ays of light M iss T o relle tried .

th e exp erime n t o f pl aci ng d ark obj ects in t h e vici n ity o f t h e

-

frog to fi n d if the an i m al showed any te n de n cy to appro ach

t hem The S id e of a l arge woode n box w as covered w ith
.

bl ack cloth an d th e frog pl ac ed n e ar the bl ack p erpe n dicul ar

,
3 48 TH E BIOLOGY OF TH E FRO G CH A P
.

su rfac e . I t hopped close to thi s rem ai n ed a couple of ,

mi nutes the n moved close t o th e w all o f the gr ay colored

,
-

buildi ng wh ere i t re m ai n ed a t res t in th e an gle formed by

, ,

t h e w all an d th e grou n d Whe n pl aced n e ar the u ncov

.

ered box ( pi n e ) o n th e sid e in fu ll su nligh t there w as n o ,

move m e n t t ow ard it Whe n th e box w as r ais ed on o n e

.

edge an d propped so th a t t he o t h er edge w as about four

,

i n ch es fro m the grou n d t h e frog m oved to w ard th e sh ado w

,

t hus formed crept well u n d er th e box pl aced i ts body b e

, ,

twee n t he flo o r an d th e grou n d where i t rem ai n ed with i ts ,

h e ad directed ou tw ard A bl ack clo th w as faste n ed close

.

t o th e grou n d in t h e c e n ter o f a su n illu mi n ated are a and a -

frog pl aced n e ar i t m oved o n to i t crep t alo ng the edge ,

as i f se eki n g cover th e n h opped o ff ,

A seco n d frog also .

hopped o n to t h e clo th bu t al most i m medi ately moved o ff

,
.

Ap p are n tly a d ark su rface brigh t ly illumi nat ed do es n o t

,

produce the e ffec t of a sh adow or o f di ffuse ligh t I t m ay .

be however th at frogs are attracted to such surfaces j ust

, ,

a s t hey are to S h adows but fi n di n g differe n t co n di t io n s o f

,

s t i mul at io n whe n t h ey ge t th ere they do n o t re m ai n .

I t i s cl e ar th at t h e frog m an ifests two quite d iffere n t

respo n ses in it s beh avio r to w ard light The orie nt i ng .

respo n se in which the a n i m al puts i t self in li n e wi t h t h e

,

di rec t io n of the r ays affo rds a good illu st rat io n o f photo

,

t axi s Th e proclivity t o se ek an d rest in the sh ad e is more

.

n e a rly a ki n to w h a t is co mmo n ly c alled p h o top a thy U n der .

o rdi n ary co n di t io n s t h e frog m ay be co n sid ered as positively

photo t actic bu t n eg at i vely ph otop at hic M any ani m als col
,
.

lect in th e sh ad e n o t bec aus e th ey are n eg atively p hoto

,

t ac t ic but bec ause wh e n th ey h appe n to re ach th e sh ad e in

,

t h e course of th eir m ovi n g abou t t hey co m e t o res t there ,

.

The collecti o n of frogs in S h ady spo t s m ay be p artly ex

pl ai ned in t his w ay bu t th ere a pp e ar s to be also a percep
,
3 50 TH E BIOLO G Y OF TH E FRO G CH A P
.

th ey go to w ard th e gree n ; in ge n er al i t m ay be s aid th at

where they are abl e to go t o w ard o n e o f t w o colors o f equ al
i n te n sity they m ove to the color lyi ng n e arest t h e violet e n d
o f t h e spec t ru m .

The eyes would n atur ally be reg arded as the org ans
through wh ich th e photot actic re spo nse is effected an d in ,

fac t t h ey pl ay an i mport an t p art in t h e process bu t as , ,

D r Parker has rece n t ly S ho w n orie nt at io n m ay be brough t

.
,

abou t m erely by the pho t ic stimul atio n of the ski n I n .

ord er to dete r mi n e th e rOle pl ayed by the eyes in photot axis

D r P arker covered a frog w i t h th e ski n of a som e w h at
.

l arger i n dividu al le avi n g o nly th e eyes feet an d s n ou t ex

, ,

posed F our specime n s thus covered were tes t ed an d it

.
,

w as fou n d th at they
“ tur ned tow ard the light an d jumpe d
to w ar d it much as n orm al frogs do Whe n a n orm al fro g
.
”

w as i n troduced for comp ariso n it w as fou n d th a t in most in

,

“
s t ances i t respo nd ed more quickly th an th e covere d o n e ,

b u t the differe n ce w as n o t so gre at th at i t migh t n o t h ave

b ee n due to the purely mech an ic al i n ter f ere n ce o f th e cover
ing ski n Whe n th e eye s o f the frog were covere d as well
”
.

as th e ski n th ere w as n o lo n ger an y respo n se t o light thu s

, ,

showi n g th a t i t w as n o t t h e ligh t wh ich m ay h ave pe n e

,

t rat e d t h e coveri n g o f de ad ski n th at effec t ed t h e orie n t a

,

tio n .

Th at photot axis m ay be produced through th e ski n alo n e

as w ell as through the eyes al o n e w as show n by P arker in

th e followi n g experi me n t E leve n frogs we re t ake n and

.
,

“by a S i n gle vertic al tr an sve rse o u t j u st beh i n d t h e eyes

, ,

t h ese org an s an d the cerebr al he mis p h eres were re moved

w i t h th e s n out o f the an i m al I t i s well k n ow n th a t frogs in

.

t his co n ditio n m ay wi t h a li t tl e c a re be kept alive for m an y

w eeks ,
a n d th a t th e ch ief differe n ce betwee n th ese an d
n o r m al frogs is the gre at reductio n in spo n t an eou s move
XV III I NS T I NCT S A ND T ROP I S M S 35 1

m e n ts show n by t h e form er O f these frogs n i ne S howed

.

an u n mist ak abl e pho t ot ac t ic r espo n se tur n i n g by th e shor t est ,

course to face th e ligh t where they re m ai n ed for a c o n “

,

siderab le p eriod usu ally termi n at ed by a j u mp tow ard th e

,

light .This re a ctio n occurs whe n th e exposed por t io n s o f

”

th e br ai n are covered by an op aque obj ec t or a sh ado w c ast

ov er the h e ad The ski n therefore is in all prob ability the
.
, ,

organ whi ch is se n sitive to sti mul atio n by light I n the .

urodeles T r iton cr ista tus and P r o teu s a ngu in us it h as bee n

sho w n th at the S ki n is si mil arly s e n sitive to light .

Thigmot axis — M an y an im als te n d t o re m ai n in situ a

.

tio n s w hich afford co n t act sti muli over a co n si derable

surface o f t h e body S u c h forms ar e c alled posi t ively
.

t higmot actic t hose which avoid co n t act are c alled n eg a

t ively thigmo t ac t ic The te n de n cy so com mo n amo n g

.

in s ects a n d w orms to cr awl u n der sto n es an d lie u n d er

crevices is to a gre at exte n t the m an ifest atio n of a t higmo
,

t ac t ic respo n se al t hough in so me c ases it m ay be due in

, , ,

p ar t to a n eg ative ph o t o t axis o r pho top athy .

F rogs oft e n S ho w a prope n sity to crawl u n der sto n es or

to ge t be t w ee n obj ects w here t hey rem ain q uiet The s ame
,
.

te n de n cy seems t o be so mewh at more m arked in t o ads .

I t i s app are n t ly stro n ger whe n the temper ature i s lowered .

M iss T o relle in experi me n ti n g with frogs pl aced in a j ar of

“
cold w at er fou n d t h at whe n a rock w as lo w ered in to th e
j ar i n such a w a y t h a t a sm a l l sp a ce w as form ed betwee n it
an d th e w all of the ar th e frog cr awl ed i n to th is sp ace an d
j ,

rem ai n ed t h ere Whe n a sp ac e w as formed betwee n the

.

bo t t om of the j ar and th e rock i t cr aw led i n to th at This ,

.

w as tested sever al t imes a n d w as al s o observed whe n the

,

temper at ure o f the w ater in th e aqu ariu m in which th e frogs

were kept w as l o w ered to 1 0 C an d belo w Wh e n t his
°
. .

w as do n e all the frogs respo nded either by fl at t e ni ng their

, ,
35 2 TH E BIOLOGY OF TH E FROG CH A P .

bodies agai n st th e sto n e floor or by c r eepi ng u n de r th e

,

rocks usu ally kept th ere I t th erefore see ms th at t h e

.

frog is ste r eo t r opic [thi gm ot ac tic ] in tempe r atu r es betwee n

1 0
°
C . and 4
°
C .
”

RE F E RE NC E S

G rab er , V . Gr u dl i i
n f sch u g d H
n en z u r Er or n es elligk eit s un d F ar
Thi
b e n sinn es der e re . P g d L i p zig 8 8 4
ra an e ,
1 .

Lo b J e C mp , . o ara ti P hysi l gy f t h B i
ve o o o e ra n , an d Co mp i
ara t ve

P sych l g y N Y k
o o . ew or , 1 9 00 .

P ar k er , G H . . The S k i n an d E y s R c p t i O g s i th R
e as e e ve r an n e eac

t i s
on of the F ro
g t o L igh t A m . . J u P h ys V l
o r .
9 3 .
,
o . 1 0, 1 0 .

T orelle, E . T he R sp s
e on e o f t h F g t L igh t
e Am J u
ro P hys o . . o r . .
,

V ol 9 ,
. 1 90 3 .

Y er k es R M , . . T he I n t s i c ts
n , H a bi t s , and Re a cti s on of th e F g
ro ,

M o n o gr S pp l . u . Psych l o . Re v , V o l 4, 1 9 03
. . .
354 TH E BIOLO G Y O F TH E F RO G CH A P
.

frogs would go hu ng r y S u b seq u en t ly I pl aced a l ar ge fly

.

u po n a piece of thi n mic a and surrou nd ed it wi th a circle

,

o f fi n e n e edles pierci ng the pl ate The fly thus protec ted

,
.

could o nly be seized by the frog sufferi n g a severe pricki n g

of the j aws This I fou n d a frog would suffer i nde fi n itely
.

in i ts attempts to secure th e fly I n o n e i n st an c e th e frog

.
,

which h ad bee n fasti ng for seve n ty t w o hours co n t i n ued to -

s n ap at the n eedle protected fly u ntil it h ad e n tirely ski n n ed

-

its upp er j aw . I co n cluded fro m this th at th e wits of a frog

were t oo limited to be de mo n str ated .
”

Kn auer fi nds th at frogs p ersist for a lo ng ti me in s n ap pi n g

at w orms fro m which th ey are sep a r a ted by a gl ass p arti tio n

without beco mi n g aw are of the fu t ili t y of their e fforts They .

will keep u p t hei r e n de avors at i n terv als all day how much
more t i m e w ould be required to co n vi n ce th em th a t thei r
efforts are v ai n is u n cert ai n .

Wood frogs accordi n g to Abbott exh ibit much more

, ,

i n ge n uity in the pursui t of prey t h a n th e ordi nary aqu at ic

species . I h ave frequ e n tly n o ticed s ays Abbo t t “ w h en
”
, ,

I pl aced flies in the c as e th at th e w o o d frog si ngled ou t

,

o n e an d a ppro ach ed i t iii a very s t e al t hy m ann er squ atti n g ,

closely to the moss hidi ng behi n d fer n s an d dr aggi n g itself

,

alo n g,
u n t il i t h ad re ached a posi tio n sui t able for m aki n g
a successfu l le ap I f th e fly moved th e frog would al ter
.
,

i ts positio n accordi ngly an d follo w u p th e ch ase with gre at

,

p ati e n ce and u n qu estio n able S kill A t t i mes it would h appe n .

th a t so me o n e of the s m aller b atrachi an s kep t in the c ase

s n apped at the cove ted prize w he n t he d isgust of the w ood,

frog would be pl ai nly show n by its m ann er but such an ,

occurre nce n ever l ed to a qu arrel .

F rogs wh e n liber ate d are ge n er ally credi t ed wi t h m aki n g

, ,

fo r w ater by the n e ares t rou te b u t Abbo t t fi n d s t h at this is

,

by no me an s al w ays the c as e F rogs te n d to ke ep n e ar .

XI X TH E T
I N E L L I G E NCE OF TH E F ROG 35 5

w ater i f th ey c an perceive it eVen whe n th ey c ann ot get ,

i n to i t although they m ay e asily re ac h an other body o f

,

w ater at a som e w h at gre ater dist an ce Abbott buri e d a .

p ail of w ater to th e brim in dry gro u n d wi thi n fifty y ards o f

a ru n n i n g brook an d covered it with a sieve,
H e th e n .

“took seve n frogs of three species an d pl aced t he m o n th e

F IG 94 by i nt h us d i s udyi g th f m i f h bi t s by h f g
La r e n t n e or at on o a t e ro .

F mA h f g s E G gl ss p l i i ; R d su f c
. .

ro t e ro P p
e n t er at . a at e ,
a rt t on re r a e

hi t su f c ; T t k ( A ft Y k s )
, ,

lV
/
,
w e r a e , an . er er e .

si eve wh ich w as about h al f an in ch above th e sur face of the

,

w at er H ere five of the m re m ai n ed duri n g the whole day

.
,

expos ed to the gl are an d h e at of a cl oudl ess su mm e r day .

The ev apor at io n fro m the w ater b e n e ath b arely kep t the m

alive ; an d ye t wi t hi n so short a dis t an ce w as a ru n n i n g

brook wi th all th e a t t r active fe a tures o f ide al frog li fe

,
.

An an im al like a dog aft er fi n di n g th at i t could n o t ge t

,

a t t h e w ater in t h e p ail would w as t e l ittle ti me in m aki n g ,

for th e brook ; but the frog is so li mited in it s resources ,

th at such a course is ou t o f re ach o f it s powers .

Y erkes h as studied th e po w er of formi n g associ at io n s in

frogs an d h as co m e to th e co n clu sio n th at th eir le arn i n g is
,

slow but th at h abits o n ce formed are h ard to ch ange Th e

, .

irogs experime n ted with were pl aced in a l abyri n th (F ig 9 4) .

formed by a box 7 2 cm lo n g 2 8 cm wide an d 2 8 cm deep .

,
.
,
. .

The frog e n ters th e box th rough a sm all op e n i n g at o ne en d

A . At the other end o f the box an ope ni n g at o n e S ide l e ad s
35 6 TH E B I OL OG Y OF TH E F RO G CH AP.

to a t ank o f w ater i nto which the frog is n atu rally desi r ous of
,

getti ng Ne ar A th e box is divid ed so th at a ch oice o f t w o

.

p aths is give n I f th e frog p asses to th e r ight its cou r se is

.
,

blocked o ff by the p artitio n P Ne ar th e oth e r end of th e

.

box t w o alter natives are also p rese n t ed in th at th e frog c an

,

go ei t h er to the l eft whe r e its course is cut off by th e gl ass

,

pl ate G or to th e right p ath wh ich le ads to th e w at er The

, ,
.

sid es of th e box were fi tted so th at colored c ardbo ard could

be pl aced in th e posi tio n s m arked W an d R an d t h e color ,

o f th e sides of th e l abyri n th could t hus be v a ri ed at will .

The p artitio n P w as also m ov able an d could be sh i fted to

th e other side o f the box so as to reverse the closed an d
free p ass ages A frog e n teri n g th e box at A u su ally do es
.

n o t go at first by th e m ost direct rou te to the w ater bu t ,

a ft er sever a l tri al s it co m es to avoid th e clos ed p ass ages an d

tr avels to th e w a ter by the s ho r tes t rou te The frog le ar ns.

this p at h very slowly as it w as fou n d to t ake fro m fi ft y to

,

o n e hu n dred tri als be fore i t would t ak e th e direct r oute

withou t bei ng l i abl e to m ake a mist ake Associ atio n s o n ce

.

formed however were fou n d to p ersist for ov e r a mo n th

, , .

I f a ft er th e frog h ad le ar n ed to go to th e w a ter by th e
,

n e arest p ath th e colored c ardbo ards li ni n g the sides of

,

th e box were exch an ged so th at the sid e th at w as re d

,

before w as m ad e white th e an i m al would b ecom e c o n

,

fused an d freque n tly t ake t h e wro ng rou te Y erk es com es .

to th e co n clusi o n th at t h e frog i s guided by color visio n

as well as by
“ com pl ex se n s atio n s o f t ur n i ng .
”

F e ar exerci ses a s t ro n g i nhibi t i n g effect o n the form atio n

o f associ a t io n s. Th e fr o gs experi me n ted wi t h by Y erkes ,

although they g ave little evid e n ce o f fe ar by m ovem e n ts ,

aft er bei n g kept in the l abor at ory for a fe w weeks they ,

w ere r e ally very t i mid an d the prese n c e of

,
a ny s t r ange

o bj ect i n flue n ced all th ei r r e ac tio ns Q ui esce n ce i t is to b e

.
,
 B I OLOGY OF F RO G
'

35 8 TH E TH E CH AP . XI X

em o tio nal feeli ngs While the r e is o fte n riv al r y amo n g .

th e m for th e possessio n o f food their struggl es are p e ace

‘

able an d betr ay n o ill temp er

“
Th e a ngry an d e nvious .

gl ances wi th whi c h accordi n g to K n auer to ads reg ard o n e , ,

of thei r n u mber which is fortu n ate e n ough to seize a wor m

for which th ey are all st r uggli n g are more m atters o f sub ,

c t ive i n terpret atio n o n t h e p art o f th e observer th an an

j e
y
re al e motio n al exp r essio n o n th e p art o f th e an im als .

H o w the cou n te n an ce of th e to ad is m odified to express an

an gry gl an ce w e ar e n o t i n formed .

O f symp athy or a ffectio n for its ki n d th e frog o r th e to ad

S hows n o t r ace C are for offspri ng i s almost of n ecessity
.

abse n t owi n g to th e m eth o ds e mploy ed in reproductio n ,

S i n ce th e you n g shift fo r the ms elves in e n tire i n depe n de n ce

o f their p are n ts The form atio n o f groups in hiber natio n .

i s doubtless brought abou t ei ther by the an i m als h app e n i ng

to get i n to the s am e n ooks or throu gh the te n d e n cy to seek ,

th e sligh t degre e of w ar mth afforded by e ach othe r s bodies ’

.

The frog an d the to ad are pu re egoists Thei r o nly acts .

w hich h ave an y refere n ce to oth er m e mb e r s o f thei r species

are those p r ompted by th e bli n d i mpulses to r eproductio n

which n ature has i mpl an ted in th es e an im als in the in

t erest s o f thei r p oste r i ty .

RE F E RE NC E S
Ab b ott , C C . . llig c f B t ch i s S ci c V
Th e I nte en e o a ra an . en e, o l. 3,
66— A N t u l ist s R mbl s b u t H m d d 8 94 ’
pp . 67 . a ra a e a o o e, 2 . e .
, 1 .

B reh i
n . T h ie rleb en . Bd 7 . .

J o r dain
u , S . De l in t ellig e n ce des
’
b at r a ci sen . C R A ss
. . . fra n c .

A v S ci ,
. . 29 m e Se ss .
, 1 9 00 .

K nau er b ch t u g
, F . R pt ili
K . d A m p h ibi
B eo i a n en an e en un en n

d G f g
er e ansch ft W i 8 75
en a . en , 1 .

Y rk e R M T h I s t i c t s H bi t s
es , .
d R cti
. s f th F g e n n , a , an ea on o e ro .

M g
ono S u pp lrP sych
. l R V l 4 9 3
.
I h ibi t i d R i foc . ev
.
,
o .
,
1 0 . n on an e n or e

m t
en f R c ti
o s i th F g R
ea l m iton J u C mp N u n e ro ,
a na c a a ns . o r . o . e r.

and P ych V l 4 1 9 4
s .
, o . 1 , 0 .
 I NDE X O F A U T H O RS

Abb tt o 353 35 8 1 3 1 9, 21 , , . uck 9

Br e, 1 2 , 277 ,

B uckl d 4
, ,

Ab l e o u s , 224 2 27 ; A b e lo u s a n d B il an ,
1 .

l d g
,

ar , 224 2 27 ; A b e lo u s a n d L an

l is C lm ls
,

o 2 26 , 227 a 83 e 1 .

C me
. ,
,

a ra n o , 1 20 .
A c h 3 37 3 40
C i cci
.

d mi
, .

A a 1 0 21 1 8 a o, 1 1
, 2
.
.
,

A lb e rt o n i, 3 1 8 .
Co n tej e a n , 1 63 .

ll
A e n 1 5 , 20 2 1 Co p e ,
1 8, 21 23 .

l d
. ,
, ,

A ng e llu ch i 3 22 , .
Co p e an , 1 20 .

A sch s
er o n 1 84 1 98
l
.
, ,

A ssh e t o n 9 5 1 0 2 , ,
.
Da e 3 1 8 , .

A t h an as iu 1 6 1 , .
Da n ile w sky 3 1 8 , .

D e k h u yz e n 2 6 3 2 8 2 , , .

Ba b er , 227 .
De w evre 1 6 3 , .

Ba gli i on 1 7 2, 1 77 3 04
Dissa rd 1 7 7 , .

lds
.
, ,

B arf u rt h , 5 5 1 20 , 1 61 Do n a o n 5 8 1 8 8 1 98
318 , .

sch
. , , ,
,

ch
B e t e re w 3 1 8 , .
Dra 1 98 , .

B e e r , 3 33 3 40 ,
.
Diir ig e n ,2 1 25 , .

B e issn e r 20 5 , 3 1 1 .
Du m e ril 2 1 2 1 0 , , .

sl y 4
,

B en e , 1 .
Du t art re 1 95 1 9 8 , , .

B er g 77 1

b h 5 4 5 9 64 8
.

d
,

B ern ar an d B rat u sch ek 8 4 E ert 1 , 1 ,

1 , 1 1 1 92 .

E ck
, . , ,

B ert 1 77 , 1 98 49 5 78
e r , 22 , 1 , 2 2, 2 .

E id m 44
.

h
,

B et e, 3 22 a , .

Eh m
.

B ib ro n 21 r 8 9 98 an n , 1 0, 1 2, 1 .

E g lm
.

ick l 3 8
,

B e 1 n 8
e 83 a nn , 1 1 1 1 84 1 85 ,
—
. , , ,

Bi d m
,

1 98 1 98
e er9 an n , 1 0, 1 9 2 , 1 94, 1 9 5 .

ld 9
.

B ill d ar4 7 , 22 ,
22 . Ew a ,
1 0, 3 37 .

Bi g 3 8
r e, 1

F i gt
.

B h o 76r, 1
77 1 a rr n o n , 21 1 .

F ti 87
.
,

B l o au , 22
7 a o, 1 .

F i ld 3
.

B ll 3
o 2 2. e , 1 1 .

F ish S ig
,

B o rn 5 1 . er -
w art , 22

ui
,

B o n 21 8 .

Fixs 7
,

B ou l g en4 er, 1 21 1 20 , 1 9 8 . en , 0 .

h m 35 8 F iss 5 5
, ,

B re 21 . ra e, .

F kl
, ,

B re u 3 37
er , . r an , 21 1 .

B ro w S qu dn -
e ar ,
21 9 . F rear , 6 .

359
3 60 IN E X D

F uchs ,
28 2 . K lug 1 74 1 77

Fu nk
, , .

e, 34 ,
1 60, 21 8 . u
Kn a e r 26 3 24 3 5 7 , 3 5 8 , , , .

K nau t h e 5 7
d
.
,

Ga ow , 3 7 9 , , , 1 1 , 22 , 1 99 . K ra w z o ff 3 1 1 , .

C ag e , 1 1 ,
15 , 22 . K ro gh 1 68 1 75 1 7
7
m 4
, ,

uk b g
, .

G ar an , 1 . Kr en e r 1 90
sk ll 3 8
.

h
,

Ga e ,
1 . K u n e 1 90
ul 3 4 6
, .

Ga e, , 1 2 , 2 27 ,
28 2 .

G au p p , 26 , 1 39 ,
1 71 , 1 77 , 1 85 ,
222 La bb e 43 , .

2 24 , 225 27 8 28 5 . L ah o u sse 154

mmill
, , , .

Ge 1 15 . L an d is 3 3 8 3 40
o

G igli T
, , , .

o -
o s, 2 1 7 , 2 1 8 . L a n g e n do rff, 3 5 3 1 9 ; an d M o z eik
Gi d 3 37
, ,

rar 3 40 1 61
—
.
, ,

G l tz 3 3 gl y 4 —48 5 4
.

o , 0 , 3 05 3 7 0 , 3 1 2, 313, 31 8, L an e ,
1 2 1 1 1 61 1 64

L ud b ch 3 4
, , , .

33 7 3 40 0
a en a 0.

b
.

L idy 4 —
,

G ra e r, 3 46 3 5 2 , . e 4 ,
0 2 .

G rut z n er , 1 4 2 , 1 45 1 46 , 1 64 L eo n ar d 34 5 4 5 9 1 1 1 60 1 64
ut h
, .
, , ,

L ydig
, , .

G 332 e 83 86 22 , 1 1 1 87, 1 89, 1 90 ,

.
, , , ,

1 99
ll 5
.

H a 1 1 .
L is ter , 1
95
mm
, .

H a a r , 2 24, 2 2 8 .
L oe b J 3 6 3 9 35 1 1 2
d s ty 3
.
, , , .

b L dS g
,

H ar e 1 8 .
L oe an t ro n 1 80
l ss 8
, , .

s 3 39
,

H ar e 1 1 1 92 L o e e r, 1 0, 1
t g
, , , .

H ar o 6 ,
2 .
L mb d
o
54 ar , 2 .

H ay , 2 2 .
L oo s 4 1
id h i
.
,

H e en a n, M . 1 64 1 81, 1 83 , 1 99 . L uchsi g 6 n e r, 1 1
id h i
, , .

H e en a n, R .
,
1 49 , 1 5 0 , 1 6 4 .
Ly on,34 0

s 3 37
.

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22 . M c llum 8
a a ,
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H e u b e l, 319 M B id
c r 5 e, 1 1 .

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.

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n e 1 1 20 ar u 2 , 2 2 .

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.
,

H i schb g 3 6
, ,

r er 2
3 33 3 40 ar 95 a , 22, , 1 09 , 1 15 1 1 9,

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,

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ss
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,

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, ,

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1 78 , .

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1 83, 1 85 1 99 . M ivart 2 2 .

ubi i
, ,

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19 , . M o o re a n d V n e n t 2 26 228 i c , , .

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, , . M o ra c z ew sk i 1 6 4 .

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, ,

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-

, .
 I ND E X OF S U B J E CT S

A bs p t i f f d
or on o oo , 156 o f w at er , A u icl r e, 279 .

58 87 97 1 1 A t d
u o l g ax u u b ris , 9 .

Ac h c p h li 4 A xis cyli d
.
, ,

a nt o e a , 1 . n er , 1 3 1 ,
28 8 .

A c t bulum
e a 43 2

l idium 4
.

A c is 7 ; A g yllus
,

r 1 . r Ba an t ,
2 .

A di p s t issu B s i di b lus
, ,

o e 5 e, 1 2 . a o m 44 o ra n aru , .

Ad l b dy
re n a o 20 2 , 2 2
5 . B t ch s p s 9
a ra o e , .

B l s m
,

A ffe re n t x
n e rve s , 3 00 . e o to
39 a, .

A gl ss o a, 1 1 . B cus 39
en a .

B idd sc l 5
,

Al li 6 8 '

ve o 1 . er an a 20 .

B il 5 5 ; c p ill i s duct
, ,

A ly t s b t t i
e o s e r ca n s , 1 2 .
5
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A mblys t m
,

7 o a, , 8, 46 ; A m b lys to 1 07 ,
152 .

m at in ee 7 . Bldd a 7 er , 1 0
9 ,
20 .

Bls c l 93
,

A m p hium A m p hiumid a, ae , 5 . a t o oe , .

A m p ull f 33 5 a o e ar , . Bls p 93
a to o re , .

A b lism 3 4
na o 1 . Bl
s t ul 93
a a, .

A gul gul sp l i l
,

n a r e , an o -
en a 233 . Bl doo 5 8 ; c p uscl s
1 21 58 2 or e 2

ulus tym p icus 33 6

, , , ,

ss ls 6 8
,

A nn an . 6 ; 2 2 ve e 2 .

A u B dy c iy
, ,

n ra , 2 , 1 1 . o av t l m s e e c ce o .

A us 64 7 4 s uc u f 6 ; c lls 7
,

n , ,
1 01 . B o ne , tr t re o 12 e ,
12 .

m s c p sul 3
, ,

A t 64 7
’
o r a, 2 ,
2 0 . B ow an a e, 20 .

A t ic
or ch s ar e , 1 12 . B chi l p l xus 8 9
ra a e 2

i 8 9 ; fu c t i s f
.

Ap d
,

o 3 a, . B ra n , 3 0, 10 , 2 1 n on o ,

A qu duc t f S yl ius
e 3 o v , 10 .
3 8 ; i flu
0 c n c d 3 en e on or 02 .

A qu us hum 3 6 chi l ch s 8 cl fts 9 8

,

eo o r, 2 B ra n a ar e ,
1 1 e , 10 , 1 1 .

di g h bi ts 45 5
.

A rac li n o id , 28 5 . B ree n a 0 .

s p t 6 4 94
,

ch
,

Ar e n t ero n , 95 . B ro w o 2 .

B ucc l c i y 69
,

cif
,

Ar e ra , 1 1 . a av t , .

A re a c e n t ra lis o f ret n a , i 331 . B uf l t igi

o sus 5 4 46 99
en no ,
1 1 , , .

y icul i s B uf id
,

A rt e r 26 8 F o r p a rt ar ar t e r e on 3 ae , 1 1 1
—
. .
,

B uf q
,

see 2 68 27 2 . o i 6
u e rc c u s , 1 .

y id c til g 65
A r te n o ar a e, 1 . B ulbus c dis 64 7 7 or 2 2 .

A sc is
, ,

ar t m l 4 en o e as , 0 . B ullf g 8
ro 9 7 5 61 1 2, 2 1 21

A s t g lus 3
.
, , , ,

ra a ,
2 0.

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22 . C lc a um 3an e 2 0 .

A udi t y g 63 3 34 ;
,

or or an , ,
n e rve , 2 98,C l c
an a ,
l f c d 3 86 ;
e n t ra o or , 10 ,
2

3 36 . s mici cul 3 3 5 3 3 7
e r ar, , .
3 64 I ND E X

C an a liculi 1 27 , . C o ra o c id
240 .

ill i s ium
,

Ca p ar e 269 2 8 0 , . Co r 1 79 , .

hyd s
,

C a rb o ra t e 1 37 , . C o rn ea 3 24 , .

Car adi c gl d an 1 40 , . us di sum 77 7 Co rp a po 21 .

id y gl d
, ,

C a ro t art e r 26 9 ; an 26 9 , , 78 2 uscl s . bl d 5 8
Co rp e of , oo 2 26 2 ; of

us
, ,

lym p h 6
,

C a rp 2 42 , . 2 2

il g c il g b s
.

i l s 95
,

C a r t a e 1 25 ; a rt a e o ne 35 C 2 . ra n a n e rve 2

ud qui
, .

ium 9
,

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ll h y
.
,

C e t eo r 82 1 21 , . C ick f g 7 r et ro 1

um
, .

C ic id c il g 65
,

Ce n t r 23 7 , . r o ar t a e, 1

b llum
.

C e re e 1 0 3 29 1 293 29 6 , , . C ki g 3 3 ro a n 1 02

b l h mis h s c b um
,

c b i 3 93
, , , .

C e re ra e p e re e re r , 3 C , 10 ,
i t ira e re r ,
10 ,
2 .

2
9 193 2
94 3 9 2 0 C yp t b chid 5 r o ra n ae ,

C hi sm p ic
, , . .

a 94 9 6
a o t ,
2 ,
2 . C yp t b chus 5 6 3 9 5 9
r o ra n .

C ys t ll i l s36
, ,

Ch N s
, ,

69
o a n ae , S l . ee a so are r a ne en 2 .

C u icl
.
,

Ch o n dro t u s 7 .
79 t e , 1 22 , 1 .

C u icul d t is 7
,

h
C o ro o at id c of eye , 23 7 ; p l xus e t a en 2.

Cu t is 7 9
,

,
1 .

C h o ro p h ilu s , 17, 46 . C y ig hid 8 st n at ae , 1 1 ,

1 .

C h m t ph s
ro a o 88 o re , 1 80 , 1 . Cyt a m oeb a 43 ,
.

C ili a , 1 22 .

C ili t d p i t h lium 7
a e e e 1 1 22 c yl s m
Da t o o a 43 .

l
, .
,

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,

r 58 a on, 2 De n t a e 2 3 4 .

i
. ,

C is t m g 79 8
er n a a na, 2 0. De n t n e 7 2 , .

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,

av 4 e, 2 0 . D er a p ae 6 7 , .

Cl eava g 89 e, . De sm o g n a t hin ae 7 8 , , .

C l p si
e
39 ne , De s m o g n a t h u s f u s c u s 8 .

l m
.
,

C l c 64 7 4
o a a, . De ve o p e n t 8 1 .

C gul t i i id sc s
, ,

oa 6 a on, 2 1 . Die rn y c ty lu s v r e en I O , .

C cc i dium 43
o . ig s
D e t i o n 1 3 6 1 42 1 48 , 1 5 1 ,
15 6 .

ig s i g s
, ,

mu t bilis
, ,

C d
o ph l
o n o ce a us a , 42 . D e t ve o r a n 1 3 4 , .

C oecilian s 3 . Dip lo discu s s ub c lavat u s 42 , .

isc gl ss d
,

Coe o l m 73 97 , , , 1 10 . D o o i ae 1 1 1 2 , .

l mic fluid 7 9 D is t mum

,

C oe o . o

h h im s fi lds 3 Di i g 9
, ,

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'
n e e ,
1 0 . v n ,
2 .

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, e e o ,
-

, 1 , 1 D p i dium 43
re an .

9 6 ; i flu c bl d 6 3 5 D um 6 3
, , ,

1 n en e on oo , 2 2, 1 . r , .

Co l ch g s 3 7 5 8 8 7 97
or an e , , 1 -
1 . Duc t bil g l] 7 3 e or a , 152 ; cys tic ,

5 3 ; h p t ic
,

lum ll 63 3 3 6
, ,

Co e a, , . 1 5 e a , 1 2 .

Co mmissu s f c d 8 6 re o or ,
2 . Duc us t d lym p h ticus en o a
3 35 .

mp s t y m ti s 3 8
,

Co en a o r o on 0 . Du d um 7 4 48
o en , , 1 .

dyl s cci p i t l 9 Du m t 85
,

Co n e , o a , 22 . ra a er, 2 .

Co n e s f ti 3 9 o re na, 2 .

Co n ju c i n 3 5 t va , 2 . E ar , 63 3 3 4 ; d l pm , eve o en t o f, 1 02 .

Co n n e c i t issu t ve3; b t e , 1 21 , 12 su cu a E chi hy chus 4

nor n 1 .

s 8 E c d m 96
,

ne o n ,
1 2. to 3 er , 10 .

p ul t i E ff s 3
,

Co a47 5 3 5 on, ,
2, 0 . t
e re n n er ve , 00 .
3 66 I ND E X
H m p h di tism 5 9 6
er a ro ,
-
1 . La cu n ae, 1 25 , 1 27

H ib
.

ti 53
e rn a on, . L an kes t rella 43 .

H i d b i 9 ; limb
,

n 3
-
ra n , 10 2 1 , 1 19 . L arva 1 0 2 1 1 6 .

y x 7
, ,

H is t l gy
,

o o ,
1 21 . L ar n , 10 1 65 .

H l t m m i idum 4
,

o os o u n t 2. La te ral pl t a e, 110 ; ve n tr icl e, 1 04 ,

H um us —
,

39 4 er ,
2 2 1 . 294 .

H ya lo k lo ssia L ieb e rku h n i 43 , . L ave ran i a, 43 .

yl
H a 1 7 1 94 ; g o e ldii 1 7 ; p
, , ,
ick er L en s o f ey e , 3 26 3 3 3 .

sic l
,

in g ii 46 ; ve r o o r, 1 7 . L eo p a r d f g ro 20 . S ee a lso Ran a
ylid p ip i s
, ,

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,

7 0 23 4
o . L ep t o t h era O h lm a ch e ri 43 ; r a n ae ,

cl ft
, , ,

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Le uc cy t s 50 1 5 4, 8, 280 .

ism 5 9—
2 97 . o e 1 , 22 1 25

yp
, ,

H no t 6 1 . L e u CO p h o re s ,
1 88 .

H yp bl s t E t d m igh t ff c s cti s
,

o a ,
see n o er . L e e t o f, 1 7 6, 1 94 ; rea on

H yp gl ss l
,

o o89 a n e r ve , 2 . t 0 , 3 45
H yp o p hysis 94 2 . L imbs 6 5 .

Li lb 5 5
, ,

nea a a, 2 .

I l um 7 4 48
e 1 . L ip s 68, .

Li
, ,

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73 7 5 10 , 1 2, 157 -
1 63 , 20 1 207

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, , , .

n n 94 ar o e, 2 . n 7 67
,
10 1 .

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, ,

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58 6 2 2 2, 28 1 ; ea r t 28 1 ;
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.
, , ,

n p o r a, a ra ,
2. a e , 1 ,
2 0 .

I sc i p t i
n r s t di 55 o ne en n ee , 2

—
.

I s ti
n er f muscl on o
47 e, 2 . Ma l i s ti c ts
e, n n o f, 31 46 5 2 ; or g s an

I sp i t i
,

n 7ra on, 1 0 . o f, 2 1 5 ; p ro n ucl us e 88 .

I s i c 34
,

n t n t, 2 . Ma p l ighi an b dy c o or o rp uscl e, 1 13,

I llig c 3 5 3
nte en e, .

I m xill y gl d 7
n t er a ar an , 1 . M an d i bl
e 23 3 .

b l f mi 3 8
,

I n t e rvert e ra o ra na , 2 . M arro w 1 27 263 , .

su i l f g
,

I s i 7 4 48
n te t n e , ,
1 . M a r p a ro 17 .

gi i 9 4 u i
,

I n va n at o n , . M at rat o n o f e g g 8 6 .

I is 3 6 xill y b h
,

r 2 Ma ar o n e 23 3 ; t ee t 23 3

ar t a e 23 3 —
.
, ,

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, .

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'

43 2 . 23 4

dull bl g
, , .

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31 3
dull y f lds
, .

sh h
, , ,

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g l b chus
.

m ximus
, ,

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'

.
7
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, ,

83 . M e an n 1 8 8 .

l h s
, ,

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is mb b s
,

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e el an
-

o ne 23 4

s y
.

b lism
,

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id ys
, ,

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,

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us s m mb s gs
, , , , ,

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'

e , . , .

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by i t h 3 3 5 s s um
,

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240 .

s ium
,

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2 . M e o va r 21 2 ,
.
 IN E X D 6
3 7

M et a b lism 3 4 o 1 .
31 1 ; n erve , 1 04, 296 ; t h l mus a a

M t c p us 4
,

icl
,

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.

e a 6 or o 1 1 . Or 62
sus 44
.
, ,

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2 . s bl s s 7
O t eo a t 12 .

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,

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2 1 ,
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eo a , .

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21 2 .

y 74
.
,

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. ,
,

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,

en , 1 v 3 21
44 , , .

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.

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3 9 49 , 1 .
v o o n, 50 ,
21
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.
.
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.
,

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a a n e , 233 .

55
2 56 ; f y 333 ;2 f hi d l g
,

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,

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,

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5 9 1 1 21

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,

55 ; s s l ch g s f 3 4 ;
,

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, ,
,

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t on e, 0, 22 .

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.

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ara a a ,
ara en o , 2 1 .

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o or a,
P idi m P
ara c o c c t i 43 u revo

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.
.
,

ara ,
2 .

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ar e 1 02 P si t s 4
ara e 0 .

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,

a a o n e, 2 2 a e, 2 2 ar e a 94 n e rve , 2
3 .

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.

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e r e o ra r e, 2 .

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.
, ,

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e a o x s, 1 e o at ae , 1 1 , 12 .

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.

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e ro 3 4 o e, 1 1 20 e v r e, 2 2 .

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.
,

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erve , 1 1 e 1 2 e n, 1 2, 1
3 .

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.
,

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er vo e 0, 1 2 e no 47 en , 1 .

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.
, ,

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e ra ar37 2 n e, 2 .
e one , 1 2 .

t i c l 5 P ic dium 7 3
,

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2 .

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,

e r 3 e a, 1 1 . er on r 12 .

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,

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,

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,

, 6 3 3 25 , . er o e ,
1 2 .

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to ne 3 9 48 3 , , 11 1 , 1 ,
1 21 .

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,

, . P 65
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,

, . a an e 2 2 .

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, ,

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, ,

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cci p i t l c dyl 9 Ph t xis 3 46

,

O a on e , 22 . o o ta .

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,

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2 . Pi m ta 85 a e r, 2 .

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o E s p h gus
a ,
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4 1 . 88
1 .

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,

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t ar o 10 .

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,

er 8 9 ,
1 1 , 1 1 . a a, o oo , 2 2 .

O p t i c chi sm 94 ; l b 3 a a, 2 o e, 10 . 293 , Pl is t p h D il kyi 43

e o o ra an ew s , .
3 68 I ND E X

P l th d n 9 ; Pl th o d tin 7 9
e o o e on ae , Ran a cla m it a n s , 1 9 , 20 , 4 1 , 339 3 46
—
. .

scul
, ,

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,

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, , ,

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,
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2 1 o a ra 2 0 e n er
9
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fusc
,

99 ;
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2 0 o ar , 2 Ra n a a, 25 , 44 48 ,

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ro en ta ,
2 . 87 , 1 75 , 1 83 , 1 86, 1 87, 20 5 , 20 6 , 2 1 6 ,

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or on an ,
1 .

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r en a , 2 .
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or a 5 3 77 vein , 1 2 Ran a o xy rh in a , 44 , 1 87 .

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.

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,

o ra n a o 22 Ran a pa tr ,
21 , 99 .

p ipi s
.

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,

re a ,
2 .
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20 , 2 3 , 24 , 27 ,

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,

re a a, 2 44 , 6 6 21 5 21 6, 2 22,
3 46 .

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. , ,

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r 99 ve ro o v e , Ra n a va t a, 20 , 2 4 , 45 3 5 4 .

t mp i
. ,

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2 .
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ro o ra o a o , 22 .

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,

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1 1 .

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1 .

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l p t l sys t m l
.

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,

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i s 6 74
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ro t e 1 ve n ,
20 , 2 .

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.
,

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ro t e e ro t ve o r a n
i 6 8 —7 7
, , ,

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,

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.
,

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. ,

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,

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4 3 7 2 .

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. ,
,

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.
,

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27 2 .
o an o ne o re t na , 2 .

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cculus 33 5
,

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,

v nar ro ra e , . Sa en o a , .

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a an ra a ra , a o a, 9 .

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.

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. S ci ic at 9 ; p l xus
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s l ch g s 3 3 5 9
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21 6 .
3 70 I ND E X

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.

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r ,
r ,
2 .

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yg hys s
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The Development Frog s Egg ’
of t he
A N I NTRO DUCTI O N TO E XP E RI M E NTA L E M B RYO L O G Y

B y T H OM A S H UNT M ORG A N, P h D . .

P rofesso r of B io l o gy in B ry n M aw r Co g ll e e A tho r of “E vo
. u l u t on i an d

A dap t a t o n ,
” “ i
Reg e n e rat i on ,
”
etc .

Clo t h n et

P r o fe ss
g s b k gi s us m uch d d t x t b k f
or Mor an
’
ot r ve a -
n ee e e -
oo or

b t h st ud t d i st uc t
o en d it sh uld s t imul t
an n r d g t ly id
o r, an o a e an rea a

i s t ig t i by p i t i g t t h id fi ld t h f g s gg s t i l l ff s t
nve a on o n n ou e w e e e ro
’
e o er o

emb y l gic l s ch — A m i j
r o o a re e ar l f S i .
”
er ca n ou r n a o c en ce.

“A cl succi c t d c m p h sibl cc u t f ll th k
e ar, n , an o re en e a o n o a e now n

p h s s
a e f t h f to iliz t i e d d l
er
p m t f t ha f g s ongg an ev e o en o e ro
’
e .

Th t th s t t m t s
a e a ecl d i t lli g ib l
en are
p ssibl th dea r an n e e as o e e r ea er

m y f l su
a ee T h m dic l s t ud
r e. t h ul d m s t it eTh g e a en S o a er . e en

e ral s t ud t t h en d h i s i t s t d i th m tt f t h p hysi
or e re a er w o n e re e n e a er o e

ca l b sis f h a di t y o ill fi d h t h fu d m t l f c t s g di g t h e
e re w n er e e n a en a a re ar n

fi s t b gi
r i gs f lif d t h st uc t u f th gg
e nn n o e an l l t h sp m e r re o -
e e as w e as e er

c lle . Th I d p d te n e en en .

A Text bo o k !
of Ent o mo lo gy
I ncl d u i g th n e si
Anat omy , P h y ol ogy , E mb ry ol ogy , and M et a m orp h oses of In
sect f s, or iu u
A g r c l t ral a nd T ech nical S ch ool s and Col l eg es a s w ell as f or
W ork ing E nt omol og is t s .

B y A L P H E US S . P A CK A RD, M D . Ph D . .

P RO F E OR O F Z O O O Y
SS L G A ND G EO O L G Y ,
B RO W N UNI E R I Y V S T

Clo t h n et

h g y dmi d y u c t t x t b k m l gy
.

I av e r e a tl a re o r re en e -
oo of E n to o o .

Th is e re on ly on e w ord t x p ss my dmi t i f t o e re a ra on o i ,

P f ss
ro e or S . W . W N U i si t y f K s s
I L L I STO , n ve r o an a .

TH E M A CM I L L A N CO M PA NY
6 4 6 6 F I F TH A VE NUE NE W Y ORK
-

,
 A MANUAL OF Z OOLOGY
By T . J E FF E RY P A RKE R
P RO F E OR O F B I O O Y
SS L G IN T H E U NI E R I Y O F OVO D NE S T TAG , U D IN N Z , . .
, A ND

W I L L I AM A H A S W E L L .

P RO F E OR SS O F B I O L O G Y I N T H E UNI V E R S IT Y O F S I DNE Y N S W , . . .

Re vis ed a n d a dap t ed f o r th e us e o f A m er ica n s c h o o ls a n d co l leg es

Clo t h 1 2m o n et

Th h b l g f l t g t d i t his c u t y f g d
er e as ee n on e a re a n ee n o n r o a oo
Z l gy d p t d f
oo o a sch ls
a S f
e I judg f m
o r o ur oo . o ar as ca n e ro ex
ami t i f t h b k it s ms t m t b
na on o lle g d d pl oo d , ee o e o e w e a rran e an an n e

d li k ly t b y us ful i c ll g s ”
an e o e v er e n o ur o e e .

H W C NN W sl y U i si t y . . O , e e an n ve r .

I t hi k it ill su pp ly
n d t b f m t f b i f b t cl
w a n ee , no e o re e , o a r e u e ar
an di t s t i g u t li f c l ss s i Z ol gy I h
n e re n o s it
ne or a e n o o . ave n ev e r een s

eq u l da i t
an d t i t duc it
n enx t y o n ro e ne e ar .
”

H D D N MOR B l i t C ll g . . E S E, e o o e e .

A S yno pt ic Text bo o k !
of Z o o lo gy
F o r Colleg es a nd S chools
B y A RT H UR W I S S W AL D W E Y S S E
I ns t r u in Z o Olo g y a t t he M as s ac h u s e t t s I n s t it u t e o f T e c h n o o gy ,
ct o r l
i f
an d A ss o c a t e P ro e s s o r o f P h s i o o g y at B o s t o n Un i v e r s t
y y l i
Clo t h 8 vo n ot

This b k i clud s i si gl lum f c

oo n e i t siz ll t h t is
n a n e vo e o o n ve n e n e a a

ne ec ss y f ar l m t y c u s i z ol gy i
or an e e A m ic
e n ar c ll g s o r e n o o n our er an o e e

sui t bl f t h g
a e or l s t ud t e d t th s m t i m
en e ra
p p i t en an a e a e e a ro er n ro

duc t i t x t d d t t is s t h b k s dis t i c t i ly c m p t i
on o e en e r ea e , or e oo n ve on o ara ve

t my f
an a o th ,s t u d t h ish s t p u su t h subj c t fu t h
or e en w o w e o r e e e r er .

Th is t p s t
ere a th b k
re en z l gy t h t x c t ly m t s t h is
no o er oo on oo o a e a ee

n ee d t h t t t s t h subj c t i
or a rea
y sui t d t t h edult st ud t h e n a w a e o e a en w o

is t s m x t t c p bl f t hi k i g f h ims lf d f mi g his
, o o e e en , a a e o n n or e an or n ow n

ju dgm t s H c t h t t m t h is
en . en e ; t h s l ct i
e f f cts
r ea en e re n ew e e e on o a

an d th i g m t diff f m t h t f xis t i g t x t b k s ; i f c t
e r a rra n e en er ro a o e n e -
oo n a ,

t his is t t xt b
no k t h t t m is p p ly us d t d y h
a e -
oo as pp li da er ro er e o -
a w en a e

to x t d d t t is s usu lly i m t h
e en e rea e lum hich t
, a n o re an one vo e, w e n er

i t t h subj c t xh us t i ly H c th h l subj c t is p i t miz d ;

n o e e e a ve en e e w o e e e o e

t h fu d m t l f cts sci c th i s f t h ut h
.

e n a f th
en a t th a o e en e, n o e eo r e o e a o r,

are
p s t d t
re e n e t h s t ud t; ch subj c t is h dl d sy pt ic lly t
o e en ea e an e no a , no

t t d f gm
re a e t ra ily i t h m j i t y f th l m t y b k s hich
e n ar , as n e a or o e e e e n ar oo , w

are f t h m st p
or e t d p t d t ch ild
o ly ; t hus t h p u p s h
ar a a e o re n o n e r o e as

b t p s t é sum é d bl it is h p d hich sh l l b sui t d t

,

een o re e n a r , r ea a e o e , w a e e o

th l g cl ss f s t ud t s f
e ar e a h m such t t m t is
o en d d or w o a r ea en n ee e .

H c t h t i t l f th
en e e k e o e w or .

TH E M A C M I L L A N CO M P A NY
6 4 66 F I F TH A V E NUE NE W Y ORK
-